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: ILLINOIS 


atural History Survey 


BULLETIN 


WATUAAL WISTORY SURVEY 
FEB 19 198] 
LIBRARY 


Contents 
Volume 31 
1973-1977 


4 


STATE OF ILLINOIS 

ILLLINOIS INSTITUTE OF NATURAL RESOURCES 
NATURAL HISTORY SURVEY DIVISION 
URBANA, ILLINOIS 


CONTENTS 


ARTICLE 1.——THE EFFECTS OF SUPPLEMENTAL FEEDING AND FALL 
DRAWDOWNS ON THE LARGEMOUTH BASS AND BLUEGILLS AT RIDGE LAKE, 
ILLINOIS. By George W. Bennett, H. Wickliffe Adkins, and William F. Childers. January, 
1973. 28 p., frontis.,.8 fig.; bibliog., index:.0%6.0 2-0 2se0s cae cuen > on erate eae 1-28 


Acknowledgments 3, The 1963 restocking of Ridge Lake 3, The feeding program 3, Fall 
drawdowns 6, Population dynamics of fishes during the feeding-drawdown period 8, The 
effects of early drawndowns in reducing bluegill numbers in Ridge Lake 13, A comparison of 
the effects of various management techniques on angling yields 14, The effects of 
supplemental feeding and drawdowns on the condition of largemouth bass and bluegills 17, 
The growth of bass and bluegills in a feeding-drawdown program 19, Discussion 22, 
Summary 25, Literature cited 26, Index 27 


ARTICLE 2.——THE REPRODUCTIVE CYCLE OF THE RACCOON IN ILLINOIS. 
By Glen C. Sanderson and A. V. Nalbandov. July, 1973. 57 p., frontis., 13 fig., bibliog., 


htc eee ae ee ae a eine nee ene eC a eat tie Seca 29-86 
Acknowledgements 29, Methods 30, Results and discussion 35, Summary 79, Literature cited 
82, Index 84 


ARTICLE 3.——NUTRITIONAL RESPONSES OF PHEASANTS TO CORN, WITH 
SPECIAL REFERENCE TO HIGH-LYSINE CORN . By Ronald F. Labisky and William L. 
Anderson. July, 1973. 25 p., frontis., 3 fig., bibliog., index ................-....0000 87-112 


Acknowledgments 87, Methods 88, Findings 90, Discussion 100, Summary 107, Literature 
cited 109, Index 111 


ARTICLE 4. ——AN URBAN EPIPHYTOTIC OF PHLOEM NECROSIS AND DUTCH 
ELM DISEASE, 1944-1972. By J. Cedric Carter and Lucile Rogers Carter. May, 1974. 30 p., 
frontis: ; 12:fig.; bibliog. ; index 2%. Sees =. salaavae elaine wean bsiaitts) crete py ote oe ae 113-144 


Acknowledgments 114, Literature review 115, Materials and methods 116, Results 122, 
Discussion 137, Summary 139, Literature cited 141, Index 142 


ARTICLE 5.——LARVAE OF THE SERICOTHRIPINI (THYSANOPTERA: 
THRIPIDAE), WITH REFERENCE TO OTHER LARVAE OF THE TEREBRANTIA, 
OF ILLINOIS. By Thomas C. Vance. August, 1974. 64 p., frontis., 89 fig., bibliog., 
anicex:” VA5-20B rie siete ih ties ie. es Sealer oe REN ioe Re ene I ee 145-208 


Acknowledgments 145, Materials and methods 146, Analysis of characters 147, 
Metamorphosis 149, Life history of Sericothrips variabilis (Beach) 150, Phylogeny 156, 
Systematics 166, Literature cited 204, Index 207 


ARTICLE 6.——ROOT INFECTION OF WOODY HOSTS WITH VERTICILLIUM 
ALBO-ATRUM. By Gerald L. Born. August, 1974. 52 p., 18 fig., bibliog., index........ 209-250 


Acknowledgments 209, Literature review 210, Code of Verticillium albo-atrum isolates 212, 
Relationship of root wounds and age of wounds on infection 213, Penetration and 
development of V. albo-atrum in roots of woody hosts 215, Effect of root infection on growth 
response of redbud & green ash seedlings 226, Effect of temperature & heat treating on 
development of V. albo-atrum in roots 234, Evaluation of systemic fungicides against V. albo- 
atrum 237, Summary 245, Literature cited 247, Index 249 


ARTICLE 7.——THE MECOPTERA, OR SCORPIONFLIES, OF ILLINOIS. By 
Donald W. Webb, Norman D. Penny, and John C. Marlin. August, 1975. 66 p., frontis., 186 
His, DIDO. , HAGER 52). = coe set take eatin oj saat ceca HED ara eee acs eee le eet erence 251-316 


Acknowledgments 252, Natural history 252, Distribution and dispersal 260, Collecting and 
preserving mecoptera 265, Morphology 266, Monographs on nearctic mecoptera 268, 
Taxonomic treatment 268, Literature cited 311, Index 315 


ARTICLE 8.——AN ELECTROFISHING SURVEY OF THE ILLINOIS RIVER, 
1959—1974. By Richard E. Sparks and William C. Starrett. August, 1975. 64 p., 10 fig., 
IP OP Te AMGERG f-teiaye le Scissor state eenstay =the fine Suetajerai cg eree tes win wowace aks hatnene ted e ave eats 317-380 


Acknowledgments 317, Procedure 318, Results 319, Discussion 332, Summary 344, Literature 
cited 377, Index 378 


ARTICLE 9.——PESTICIDES AND ENVIRONMENTAL QUALITY IN ILLINOIS. 
By Robert L. Metcalf and James R. Sanborn. August, 1975. 56 p., 3 fig., bibliog., index. . .381-436 


Acknowledgments 381, Use of pesticides 381, Need for surveillance 382, Benefit-risk of 
pesticide use 383, Early-warning technology 383, Model-ecosystem technology 385, 
Herbicide test results 386, Organophosphorus insecticide test results 389, Carbamate 
insecticide test results 392, Miscellaneous insecticide test results 393, Organochlorine 
insecticide test results 394, Fungicide test results 399, Discussion 400, Literature cited 433, 
Index 436 


ARTICLE 10.——THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS: SYSTEMATICS 
AND DISTRIBUTION, AND LIFE HISTORY IN WOLF LAKE, ILLINOIS. By Brooks M. 
Burr. September, 1977. 30 p., 7 fig., bibliog., index.............. 0.0000. c eee eee eee 437-466 


Acknowledgements 437, Methods and materials 438, Systematics 439, Distribution 447, 
Conservation status 448, Life history in Wolfe Lake 449, Literature cited 461, and Index 465 


a ILLINOIS 
tural History Survey 
= BULLETIN 


RUATURAL (USiuAY SURVEY 
JUN 29 1973 
IRRAPY 


The Effects of Supplemental 
Feeding and Fall Drawdowns 
on the Largemouth Bass and 
Bluegills at Ridge Lake, Illinois 


rge W. Bennett 
Nickliffe Adkins 
iam F. Childers 


THE LIBRARY OF THE 
: OF ILLINOIS MAY 30 1973 
ARTMENT OF REGISTRATION AND EDUCATION UNIVERSITY OF 
AT URBANA.CHAMPAIGNS 


URAL HISTORY SURVEY DIVISION | ISL 
JANA, ILLINOIS 


VWALiaarcr 941 ADOTICIVOC 7 


ILLINOIS 


atural History Survey 
BULLETIN 


The Effects of Supplemental 
Feeding and Fall Drawdowns 
on the Largemouth Bass and 
Bluegills at Ridge Lake, Illinois 


liam F. Childers 


E OF ILLINOIS 
ARTMENT OF REGISTRATION AND EDUCATION 


TURAL HISTORY SURVEY DIVISION 
BANA, ILLINOIS 


STATE OF ILLINOIS 


DEPARTMENT OF REGISTRATION AND EDUCATION 


BOARD OF NATURAL RESOURCES AND CONSERVATION 
WILLIAM H. Rosinson, Chairman; THOMAS PARK, Ph.D., Biology; L. L. SLoss, Ph.D., Geology; (VACANT), 


Chemistry; RoBeRT H. ANDERSON, B.S.C.E., Engineering; CHARLES E. Oumstep, Ph.D., Forestry; 


Wek. 


Everitt, E.E., Ph.D., Representing the President of the University of Illinois; RoGER E. BEYLER, Ph.D., 
Representing the President of Southern Illinois University. 


NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 


SCIENTIFIC AND TECHNICAL STAFF 


GEORGE SPRUGEL, JR., Ph.D., Chief — 
Auice K. ADAMS, Secretary to the Chief 


Section of Economic Entomology 


Wittiam H. LUCKMANN, Ph.D., Entomologist and 
Head 

WILLIS N. Bruce, Ph.D., Entomologist 

Wayne L. Howe, Ph.D., Entomologist 

STEVENSON Moore, III, Ph.D., Entomologist, Exten- 
sion 

Howarp B. Perry, Ph.D., Entomologist, Extension 

James E. ApPLeBY, Ph.D., Associate Entomologist 

Epwarp J. ARMBRUST, Ph.D., Associate Entomologist 

Marcos KoGAN, Ph.D., Associate Entomologist 

JosEPH V. MaAppox, Ph.D., Associate Entomologist 

RONALD H. MEYER, Ph.D., Associate Entomologist 

Rosert D. PAUSCH, Ph.D., Associate Entomologist 

RALPH E. SecuRIEST, Ph.D., Associate Entomologist 

JouN K. BouSEMAN, M.S., Assistant Entomologist 

GeorGE L. GopFREY, Ph.D., Assistant Entomologist 

WILLIAM G. RUESINK, Ph.D., Assistant Entomologist 

JAMES R. SANBORN, Ph.D., Assistant Entomologist 

Douctas K. SELL, B.S., Assistant Entomologist 

CLARENCE E. Wuite, B.S., Assistant Entomologist 

KEUN S. ParRK, M.S., Assistant Chemist 

Sue E. WATKINS, Supervisory Assistant 

DonaALD E. KUHLMAN, Ph.D., Assistant Professor, 
Extension 

RoscoE RANDELL, Ph.D., Assistant Professor, Exten- 
sion 

Tim Cootey, M.A., Assistant Specialist, Extension 

JEAN G. WILSON, B.A., Supervisory Assistant 

NATALIE EKt, B.A., Research Assistant 

MartTHA P. MILLER, M.S., Research Assistant 

ANNEMARIE ReppoRG, B.S., Research Assistant 

KETURAH REINBOLD, M.S., Research Assistant 

Nancy TSUNG, M.S., Research Assistant 

STEPHEN Roperts, B.S., Junior Professional Scientist 

JoHN T. SHAw, B.S., Junior Professional Scientist 

DENISE A. Cope, B.S., Technical Assistant 

Lowe. Davis, Technical Assistant 

Marcia JANES, B.S., Technical Assistant 

Lu-Pinc KAN, M.S., Technical Assistant 

Mary KATHRYN MCCLENDON, B.S., Technical Assist- 
ant 

CHING-CHIEH YU, Ph.D., Technical Assistant 


Section of Botany and Plant Pathology 


J. Cepric CARTER, Ph.D., Plant Pathologist and Head 

Rosert A. Evers, Ph.D., Botanist 

Junius L. Forsserc, Ph.D., Plant Pathologist 

EUGENE B. HIMELIck, Ph.D., Plant Pathologist 

R. DAN NEELY, Ph.D., Plant Pathologist 

D. F. SCHOENEWEISS, Ph.D., Plant Pathologist 

J. LELAND CRANE, Ph.D., Associate Mycologist 

WALTER HARTSTIRN, Ph.D., Assistant Plant Pathol- 
ogist 

Betty S. NELSON, Junior Professional Scientist 

GENE E. Reip, Technical Assistant 


Section of Aquatic Biology 


See W. BENNETT, Ph.D., Aquatic Biologist and 

ea 

D. Homer Buck, Ph.D., Aquatic Biologist 

R. WELDON LARIMORE, Ph.D., Aquatic Biologist 

Rosert C. HILTIBRAN, Ph.D., Biochemist 

WILLIAM F. CHILDERS, Ph.D., Associate Aquatic 
Biologist 

DONALD F. HANSEN, Ph.D., Associate Aquatic Bi- 
ologist 

RICHARD E. SPARKS, Ph.D., Assistant Aquatic Bi- 
ologist 

ARNOLD GNILKA, Ph.D., Junior Professional Scientist 

RicHARD J. BAuR, M.S., Research Assistant 

DENNIS L. DooLey, Technical Assistant 

LINDA KLIPPERT, B.S., Technical Assistant 

Mary FRANCES MARTIN, Technical Assistant 

KENNETH R. WALKER, Technical Assistant 

C. RUSSELL Rose, Field Assistant 


CONSULTANTS AND RESEARCH AFFILIATES: 


Section of Faunistic Surveys and 
Insect Indentification 


Puitip W. SMITH, Ph.D., Taxonomist and Head 
WALLACE E. LABERGE, Ph.D., Taxonomist 

MILTON W. SANDERSON, Ph.D., Taxonomist 

Lewis J. STANNARD, JR., Ph.D., Taxonomist 

Larry M. Pace, Ph.D., Assistant Taxonomist 
JOHN D. UNziIcKER, Ph.D., Assistant Taxonomist 
DoNALD W. Wess, M.S., Assistant Taxonomist 
BERNICE P. SWEENEY, Junior Professional Scientist 


Section of Wildlife Research 


GLEN C. SANDERSON, Ph.D., Wildlife Specialist and 
Head 

FRANK C. BELLROSE, B.S., Wildlife Specialist 

RICHARD R. GRABER, Ph.D., Wildlife Specialist 

Haro_p C. HANSON, Ph.D., Wildlife Specialist 

WILLIAM L. ANDERSON, M.A., Associate Wildlife 
Specialist 

W. W. CocHRAN, JR., B.S., Associate Wildlife 
Specialist 

WILLIAM R. Epwarps, M.S., Associate Wildlife 
Specialist 

Jack A. ELLIs, M.S., Associate Wildlife Specialist 

RONALD F. LaABisKy, Ph.D., Associate Wildlife 
Specialist 

CHARLES M. NIXON, M.S., Associate Wildlife 
Specialist 

Rosert E. GREENBERG, M.S., Assistant Wildlife 
Specialist 

G. BLAIR JOSELYN, M.S., Assistant Wildlife Specialist 

Davip R. VANCE, M.S., Assistant Wildlife Specialist 

RONALD L. WESTEMEIER, B.S., Assistant Wildlife 
Specialist 

RONALD E, DuzAN, Junior Professional Scientist 

HELEN C. SCHULTZ, M.A., Technical Assistant 

HILDA WIESENMEYER, Technical Assistant 

ELEANORE WILSON, Technical Assistant 

Rosert D. Crompton, Field Assistant 

JAMES W. SEETS, Laboratory Assistant 


Section of Administrative Services 


RosBert O. WATSON, B.S., Administrator and Head 


Supporting Services 


Witma G. DILLMAN, Property Control and Trust 
Accounts 

Rosert O. ELLIs, Assistant for Operations 

Lioyp E. HUFFMAN, Stockroom Manager 

J. WILLIAM Lusk, Mailing and Distribution Services 

MELVIN E. ScHWarRtTz, Financial Records 

JAMES E. SERGENT, Greenhouse Superintendent 


Publications and Public Relations 


OwEN F. GLISSENDORF, M.S., Technical Editor 

Rosert M. ZEWADSKI, M.S., Associate Technical 
Editor 

SHIRLEY MCCLELLAN, Assistant Technical Editor 

Lioyp LEMERE, Technical Illustrator 

WILMER D. ZEHR, Technical Photographer 


Technical Library 


Doris F. Dopps, M.S.L.S., Technical Librarian 
Doris L. SUBLETTE, M.S.L.S., Assistant Technical 
Librarian 


SysTeMATIC ENToMmoLOGy, RopERICK R. IRWIN, Chi- 


cago, Illinois; WivpLire RESEARCH, WILLARD D. KuimstRA, Ph.D., Professor of Zoology and Director of Co- 
operative Wildlife Research, Southern Illinois University; PARASITOLOGY, NORMAN D. LEVINE, Ph.D., Profes- 
sor of Veterinary Parasitology, Veterinary Research, and Zoology and Director of the Center for Human 
Ecology, University of Illinois; ENTOMOLOGY, Ropert L. METCALF, Ph.D., Professor of Zoology and of En- 
tomology and Head of the Department of Zoology, University of Illinois; and GILBERT P. WALDBAUER, Ph.D., 


Professor of Entomology, University of Illinois; Statistics, HORACE W. Norton, Ph.D., Professor of Sta- 
tistical Design and Analysis, University of Illinois. 


CONTENTS 


AV OTE DOH YL DDEITOINTOS, Wy cP Ry eucs DIS DOK Osbtel OENG Cle eo to ot oho ROME EOI meet a aR eT Renee ic seer core 3 
Sige OAPI MESTO CIGIN GHO Re RIDGE Ss GAKR!  avcnet Shits, ccets sos io ldichacens once aia Sha. oa. avaeleybuawuacanace ayia 3 
“Tigee TPEvsipiisve; JEROeRUNN G3) cig ties Search tech tr RETO cen Bre ern pe 3 
TAiLi, LD RUN WADIERIANISh: Ont Sy lead Sala 0a epee: ORIENT GI Ee ee ey ei ees reece cee 6 
PoputaTion Dynamics oF FisHes DuriING THE FEEDING-DRAWDOWN PERIOD.......... 8 
IL@ueS MOVIN LS so 5g A Se os nice BIOs DOR CIDR eee inne dee Orgran 8 
mecily sak dee dpoMOGoe ASSO CU RE CoCmee CUCM Soman AEs Wo Rees sic Rome m os or 10 
VAY San PTOLLG NS 4s Syery dog eacie ented o- tut ERED. © On HORSE REESE RCACL, NHPSEE oe NE IPR ne RTI ope eat il 
(Clrewaal CABINS 2% aleased oS Han & TO EIST O GEE ONS OICSS ae Bia DIES G DIS Ie mee reat 12 


Tue Errects oF Earty Fatt DrRAwpowNs IN REDUCING BLUEGILL 
CINE MR GREEN ROI He GA KEN eee apanne 2, aieisy sss cee NERS eee Hi Ge Creed sta caeeenane eee 13 


A CoMPARISON OF THE EFFECTS OF VARIOUS MANAGEMENT TECHNIQUES 
ONE ANIGUUN GUYCTELD Stmteeys ruta at Net ci mebs uce (teemgc tee Sate tains Soecaee ye ashe e Shia 14 


Tue EFFECTS OF SUPPLEMENTAL FEEDING AND DRAWDOWNS ON THE 


ConpITIon oF LARGEMoUTH BASS AND BLUEGILES...........0.cc0eeceseseese 17 
THE GrowTH OF Bass AND BLUEGILLS IN A FEEDING-DRAWDOWN PROGRAM eeenneone 19 
IDES GIERSSIIOINE cea era otic ehee Gk stich © FG CIRO IG SSR DRES = CIPIEy CR Mir iiet Mite By ener ar NETS 22 
‘SHUMENDAENY a, aiatd cid RA o-cx6 Enlace S ORE TDM CPT ESLER pe hc, cA te Pena aes Eee re 25 
PER AG UREA CITED bees ary coy weer acer ny eee esata acto nuakiis onic aie a alt i savas ee arecane 26 
TESTBYESS. . 9 o opie Dtott Sia NL SIAES iS 9 6.5 Ri IO OO ERE aC es Ens ee arn emery eee 27 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It is 
a contribution from the Section of Aquatic Biology of the Illinois Natural History Survey. 


George W. Bennett is Aquatic Biologist and Head, Section of Aquatic Biology, Illinois 
Natural History Survey. H. Wickliffe Adkins is Head, Science Department, Benjamin Franklin 
Junior High School, Champaign, Illinois, and has been employed by the Illinois Natural History 
Survey each summer since 1952 as Resident Biologist at Ridge Lake. William F. Childers is an 
Associate Aquatic Biologist, Illinois Natural History Survey. 


(45582—5M—1-73) 


JAWS 


vat 


TRH. 


The Effects of Supplemental Feeding 
and Fall Drawdowns on the Largemouth Bass 
and Bluegills at Ridge Lake, Illinois 


IN 1963 WHEN THE PROGRAM 
described here was begun, studies of the 
fish population of Ridge Lake had been 
going on for 21 years (Bennett 1954a and 
19546; Bennett & Durham 1951; Durham 
& Bennett 1949 and 1951; Bennett, Ad- 
kins, & Childers 1969). These studies 
involved annual controlled public fishing 
during June, July, and August and drain- 
ing censuses (usually in the spring) to 
gain estimates of the total population of 
fishes in the lake. Between these draining 
censuses we applied several types of pri- 
mary or secondary population manipula- 
tion, or none at all, to explore the effects 
of these manipulations upon the fish pop- 
ulations and the yields of fishes. This 
period included 10 years of biennial drain- 
ing of the lake and culling of small fishes; 
5 years of fall drawdowns of the lake, 
with one draining census after 2 years and 
one after 3; 4 years of stable water levels 
and no manipulation of the fish popula- 
tion; and 3 years of testing the value of 
hybrid sunfishes for angling (Childers 
1967:189). In the period 1941-1970, 
Ridge Lake has been completely drained 
and the fishes have been censused 10 
fmes in! 1943" 1945; 1947) 1949) 1951), 
1953, 1956, 1959 (in the fall) , 1963, and 
1970. On the basis of complete creel 
censuses in all years (except 1942, when 
the lake was closed to fishing) we were 
able to measure with some degree of cer- 
tainty the type of fish population the lake 
would support and the effects of various 
management efforts on that population. 

The fishes included in this investiga- 
tion were largemouth bass, Micropterus 


Frontispiece—A 3-meter (10-foot) drawdown at Ridge Lake. 


George W. Bennett 
H. Wickliffe Adkins 
William F. Childers 
salmoides (Lacépéde) ; bluegills, Lepomis 


macrochirus Rafinesque; warmouths, Le- 
pomis gulosus (Cuvier) ; lake chubsuckers, 


Erimyzon sucetta (Lacépede) ; and chan- 
nel catfish, Jctalurus punctatus (Ra- 


finesque). Any other fishes that gained 
entrance to the lake through fishermen’s 
minnow buckets or from the- drainage 
basin were removed during the draining 
censuses. All of the fishes in the Ridge 
Lake population descended from 435 
largemouth bass stocked in 1941, 129 
bluegills stocked in 1944, 138 warmouths 
stocked in 1949, 558 lake chubsuckers 
stocked in 1960, and several groups of 
6- to 12-inch channel catfish stocked in 
1951, 1952, 1957, and 1969. The several 
stockings of catfish were necessary be- 
cause channel catfish usually cannot re- 
produce successfully in Ridge Lake. All 
of the other fishes (bass, bluegills, war- 
mouths, and chubsuckers) have main- 
tained adequate populations through nat- 
ural reproduction and survival with no 
stocking but the original one. 
Experimental drawdowns were begun 
at Ridge Lake in 1951. In the spring of 
that year the lake was drained; the fishes 
were censused; and selected numbers of 
largemouth bass, bluegills, and warmouths 
were returned to the partially filled basin. 
The lake refilled before June 1 and was 
opened to controlled public fishing during 
the summer until September 1. Fishing 
was then terminated and the lake level 
was lowered 4.6 meters (15 feet), reduc- 
ing the surface area from 6.9 to 2.0 ha 
(17 to 5 acres) and the maximum depth 
from 7.6 meters to 3 meters (25 to 10 


About one-third of the 


lake bottom, mostly in the upper end of the impoundment, is exposed by such a drawdown. 


1 


2 Ittrnoris NaTuRAL History Survey BULLETIN 


feet) without allowing any fishes to es- 
cape with the water. ‘Thus, the fish 
population that had developed through 
natural reproduction and growth to fill 
a volume of water represented by a sur- 
face area of 6.9 ha’ (17 acres) became 
concentrated in a volume represented by 
2 ha (5 acres), a minor fraction of the 
full lake. 

In 1952, after the lake had been open 
to public fishing during the summer, the 
lake level was again drawn down 4.6 
meters (15 feet) in early September. In 
late March of 1953 the lake was complete- 
ly drained, the fish were censused, and 
selected individuals were returned to the 
lake. These studies were reported in 
several papers (Bennett 1954a and 19545; 
Bennett et al. 1969). The effect of the 
4.6-meter drawdowns in 1951 and 1952 
upon the bluegills was severe, reducing 
their numbers to the point that fewer 
small bluegills survived than did large 
ones. To insure the survival of enough 
bluegills to maintain successive year 
classes for fishing, it was decided to limit 
drawdowns to 3 meters (10 feet), leaving 
a maximum lake depth of 4.6 meters (15 
feet) near the dam and a lake surface 
area of about 4.5 ha (11 acres). This 
procedure was followed in early Septem- 
ber of 1953, 1954, and 1955, and the lake 
was drained again in the spring of 1956 
and a census was made of the fishes. The 
lesser drawdowns of 1953-1955, inclusive, 
allowed the survival of a greater number 
of bluegills (both larger and smaller than 
150 mm, or 6 inches, the length at which 
this fish was considered useful) than the 
+.6-meter drawdowns allowed. 

Following the spring lake draining and 
fish census of 1956, selected bass, blue- 
gills, warmouths, and a few channel cat- 
fish were returned to the lake; the basin 
refilled by May. From March 1956 until 
October 1959 the water level in Ridge 
Lake was allowed to fluctuate around the 
crest of the tower spillway, i.e., without 
any drawdowns and with only minor fluc- 
tuations caused by runoff from rains in 

1 Ridee Lake oyateeinelity had a surface area of 7.3 ha 
(18 acres); silt deposits in the upper lake had re- 


duced the area to about 6.9 ha by 1953 and to 6.5 ha 
hy 1963. 


Vol. 31, Art. 1 


the lake watershed. In October 1959, 
after four growing seasons for fishes, the 
lake was again drained completely and 
the fishes were censused. As in the years 
of the drawdowns, the lake was open to 
controlled public fishing during the sum- 
mers of 1956-1959. 

Thus, as a background for the experi- 
ment reported here, the authors had in- 
formation on anglers’ catches and total 
fish populations from 10 years of biennial 
draining of the lake and culling of the 
fishes, 5 years of drawdowns, and 4 years 
of stable water levels (including that part 
of 1959 important for fish reproduction 
and growth). From the lengths, weights, 
and scales of individual fishes taken by 
fishermen in 1951-1959 and from similar 
data gathered from fishes during the 
draining censuses of 1953, 1956, and 1959, 
it was possible to compare the growth 
rates of bluegills and their relative plump- 
ness (condition) under a program of an- 
nual fall drawdowns and under another of 
stable water levels. 

In the fish censuses of 1953 and 1956, 
many of the bluegills were of exceptional 
sizes but appeared to be comparatively 
thin. The supplemental feeding proposed 
at the beginning of the experiment re- 
ported on here was in part related to this 
observation. 

Our laboratory production and culture 
of hybrid sunfishes (Childers 1967) had 
demonstrated that most species of sun- 
fishes (Centrarchidae) quickly learn to 
feed upon commercial trout food if the 
pellets are small enough for them to swal- 
low. Bluegills in laboratory aquaria be- 
came plump and grew rapidly on a trout 
pellet diet. If bluegills in Ridge Lake 
could be trained to eat fish food pellets 
to supplement their diet of natural foods, 
this additional food supply should be re- 
flected in improved bluegill growth and 
condition. We wished to discover wheth- 
er enough improvement in bluegill yield 
would occur to make artificial feeding 
practical. In 1963 following the spring 
fish census, we decided to combine Sep- 
tember drawdowns of Ridge Lake (for 
the control of bluegill numbers) with 
supplemental feeding to increase the 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FisHEs 3 


growth rate, condition, and yield of these 
fish. Results would determine whether 
such a program was practical. 


ACKNOWLEDGMENTS 


Mr. H. Wickliffe Adkins, stationed at 
the Ridge Lake Laboratory during the 
summer months, supervised the fishermen 
and recorded their catches, fed the fishes 
twice daily, made daily observations on 
schools of bass fry and on the nesting of 
bluegills, assisted in age and growth analy- 
ses from fish scales, and recorded many 
biological happenings of importance to 
this study. Dr. William F. Childers 
planned and supervised the draining cen- 
suses in 1963 and 1970. Many people 
assisted in the fish censuses; these included 
Mr. Robert O. Ellis, Mr. Howard Crum 
(deceased), Mr. Robert T. Crompton, 
Mr. Dennis Dooley, Dr. D. Homer Buck, 
Mr. Richard Baur, Mr. Russell Rose, Dr. 
R. Weldon Larimore, Mr. H. W. Adkins, 
Mr. Ronald Havelka, Mr. David Mower, 
Mr. Edward Doyle, and Dr. George Spru- 
gel, Jr., of the Ilinois Natural History 
Survey staff; Mr. Alvin C. Lopinot, Mr. 
Arnold Fritz, and Mr. Rudy Stinauer of 
the Illinois Department of Conservation; 
.and Dr. Leonard Durham and Scott Buck 
and other students from Eastern Illinois 
University. The manuscript of this paper 
was read and criticized by Dr. Horace W. 
Norton, Professor of Statistical Design 
and Analysis, Department of Animal Sci- 
ence, University of Illinois, and it was 
edited by Mr. Robert M. Zewadski of 
the Natural History Survey. 


THE 1963 RESTOCKING OF 
RIDGE LAKE 


Following the draining census of April 
8-13, 1963, Ridge Lake was restocked 
with 2,270 small bass and 116 large ones, 
4,492 bluegills, 1,335 warmouths, 1,020 
lake chubsuckers, and 11 large channel 
catfish, a total of 9,244 fishes weighing 
510.6 kg (1,125.5 pounds) (Table 1). 
The weight of these fishes was 78.8 ke 
per hectare, or 70.3 pounds per acre. Be- 
fore the restocking, the lake contained 
287 kg per hectare (256 pounds per 


acre), almost four times the weight of 
fish returned to the lake. 

A total of 1,000 channel catfish were 
stocked on May 21 and 29, 1969. These 
were Age III fish with an average total 
length of 259 mm (10.2 inches) and an 
average weight of 127 grams (0.28 
pound); their total weight was 127 ke 
(280 pounds). On October 20 and 21, 
after the 1969 growing season was nearly 
over, an additional 1,000 channel catfish 
were released. These were also Age III 
fish, averaging 234 mm (9.2 inches) in 
total length and 113 grams (0.25 pound) 
and having a total weight of about 113.4 
kg (250 pounds). All of these catfish 
originated in Arkansas in 1968 and were 
held in ponds on the Sam A. Parr Co- 
operative Fisheries Research Center in 
Marion County, IIl., until stocked in 
1969. 

After the spring draining and restock- 
ing of 1963, the fish population was fished 
by the public during the summers of 1963 
and 1964 under the regular creel census- 
ing system. Otherwise, the restocked fish 
population of Ridge Lake was allowed to 
expand for almost 2 years before any ex- 
perimental management program was ap- 
plied. During the summer of 1963 fish- 
ermen caught 299 largemouth bass, 358 
bluegills, 49 warmouths, and 11 hybrid 
sunfishes, weighing a total of 113.4 ke 
(250 pounds) in 1,816 hours; the catch 
in 1964 was composed of 554 bass, 1,287 
bluegills, 108 warmouths, and 65 miscel- 
laneous hybrid sunfishes, weighing a total 
of 232.2 kg (512 pounds) in 2,346 hours 
of fishing. While the total yield per 
hectare in 1964 was twice that of 1963 
(1963, 17.5 kg per hectare, or 15.6 pounds 
per acre; 1964, 35.9 kg per hectare, or 
32 pounds per acre), both must be con- 
sidered much below the average of the 
hook-and-line yields of fishes from Ridge 
Lake in the 1941-1963 period. 


THE FEEDING PROGRAM 


In May 1965, a supply of Splash Ex- 
panded Fish Food was purchased from 
the Ralph G. Wells Company of Mon- 
mouth, Ill. This food consisted of fish 


Vol. 31, Art. 1 


Inuino1is NaTuRAL History Survey BULLETIN 


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Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FIsHES 5 


meal, corn distillers’ dried solubles, meat 
and bone meal, soybean meal, cottonseed 
meal, wheat shorts, dehydrated alfalfa 
meal, brewers’ dried yeast, yellow hominy 
feed, salt, vitamin A and D oils, vitamin 
A palmitate, D-activated plant sterol, d- 
alpha tocopherol acetate, thiamin hydro- 
chloride, riboflavin supplement, calcium 
pantothenate, niacin, choline chloride, 
vitamin B-12, and trace amounts of nine 
additional compounds. The food was 
32 percent protein and 4 percent fat. 
Total calories of energy per pound of fin- 
ished feed were recorded as 1,884. 

The original pellets were too large for 
most bluegills to swallow, but they soon 
learned to pick at the pellets until they 
could break off pieces small enough to 
swallow. After the first season, we pur- 
chased smaller pellets. About half of the 


pellets would float for several hours. The 
rest would become waterlogged and sink 
almost at once. 

Feeding was begun in late May or early 
June, and bluegill spawning beds were 
selected as feeding areas along with the 
area around the boat dock, where blue- 
gills were observed to congregate (Fig. 1). 
The fishes were fed twice each day at 
10:30 AM and 6:30 PM; the dry fish 
food pellets were broadcast by hand from 
a boat. The authors assumed that 
food pellets that sank into bluegill nests 
would be picked up by the guarding males 
for removal from the nests, at which 
time these fish would discover that the 
pellets were edible. This was exactly 
what happened, and bluegills were active- 
ly foraging for pellets after less than a 
week of daily feeding. In less than 2 


Fig. 1.—H. W. “Wick” Adkins scattering food for bluegills from the laboratory pier. 


6 Ixttinois Natura History Survey BULLETIN 


weeks it became possible to distinguish 
bluegills that were eating pelleted food 
from those that were not by their obvious- 
ly plump condition. Bluegills were more 
interested in the pellets that floated and 
those in the process of sinking than those 
that had reached the bottom. Probably 
most of the latter were picked up by cat- 
fish after dusk. 

The quantity of pelleted fish food pur- 
chased and fed each season amounted to 
1,360.5 ke in a 6.48-ha lake (1.5 tons per 
16 acres of lake). This represented 210 
kg per hectare per season (187.5 pounds 
per acre per season) or a little more than 
2.2 ke per hectare per day (2 pounds 
per acre per day). The cost of the pellet- 
ed food used in this experiment was 6 
cents per pound when purchased in lots 
of 1,000 pounds or more. With the feed- 
ing rate given above, the cost was $11.25— 
$12.19 per acre per season or $27.80- 
$30.12 per hectare per season. As men- 
tioned above, feeding was begun in late 
May or early June, and it was continued 
through August. 

It became evident that not all of the 
bluegills were feeding on the pelleted 
food, either because they had not learned 
to eat it or because they had not ranged 
into areas where the food was available. 
These fish appeared to be quite thin. 
Some fish appeared to be feeding almost 
exclusively on “Splash,” and when we 
dissected them, we found that their diges- 
tive tracts were gorged with this food. 
These fish rather quickly became very 
plump and developed fatty deposits in 
the mesentaries between the loops of the 
intestine. After bluegills had fed exclu- 
sively on Splash for a month or more, the 
livers of these fish lost their dark red 
color and became pink, suggesting fatty 
degeneration. 


FALL DRAWDOWNS 


The early fall drawdowns proposed for 
Ridge Lake were similar in extent and 
timing to those performed there in the 
period 1951-1956. The objectives were: 
(1) to concentrate the fishes that had de- 
veloped in a 6.5-ha lake (16 acres) with- 


Vol. 31, Art. 1 


in a much smaller volume of water to 
cause selective mortality among the small- 
er fishes by stranding and by predation; 
(ii) to expose a significant portion of the 
lake bottom to oxidation and drying; 
(iii) to time the drawdown so that it 
would coincide with at least a month of 
warm weather during which water tem- 
peratures would remain at 18° C. (64° 
F.) or above. In the 1951-1956 period 
our draining censuses in 1953 and 1956 
indicated that drawdowns within the 
range of 3.0-4.6 meters (10-15 feet) 
would reduce the number of bluegills in 
the Ridge Lake population by 80—90 per- 
cent. 

Early fall drawdowns were conducted 
each year in early September, 1965-1969, 
inclusive. In 1965 the lake level was 
lowered 4.6 meters (15 feet) over a period 
of 15 days (August 30-September 13, in- 
clusive). This slow drawdown was the 
result of some intermittent rains and our 
concern about the poor condition of the 
road beyond the boundary of the park, 
where the outlet channel from the lake 
became a ford for several farm families. 
By October 3, the water level was back 
up to within 3.4 meters (11 feet) of the 
full level. 

In 1966 and 1967 the lake level was 
lowered 3 meters (10 feet) below the 
full level. In both years draining was 
started on August 28 and completed by 
August 31. The road ford was regraveled 
in 1966 so that automobiles could pass 
through a greater flow of water, and little 
or no fall precipitation occurred. 

In the summer of 1968 there was visual 
evidence ef an abundant supply of small 
bluegills. Consequently, in the fall of 
1968 the lake level was again lowered 
by 4.6 meters (15 feet) ; in 1969 the draw- 
down lowered the lake level 4.3 meters 
(14 feet). The drawdown operation re- 
quired 3 days in 1968 and 5 days in 1969. 

In every year the lake had completely ~ 
refilled by April. There was no evidence 
of loss of fish from winterkill, as even 
when the lake level was lowered by 4.6 
meters (15 feet) there was always an 
area of water above the dam where the 
water was 3 meters or more in depth. 


EFFECTS OF FEEDING AND DRAWDOWNS ON FISHES 7 


Jan., 1973 BENNETT ET AL. 


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8 Inurnois NaTuRAL History SurvVEY BULLETIN 


POPULATION DYNAMICS OF FISHES 
DURING THE FEEDING-DRAWDOWN 
PERIOD 


Table 2 shows the yields of the four 
species of fishes taken by anglers in the 
fishing seasons 1963-1969, inclusive. 
This table also shows the averages of 
the yields for the 2 years when no sup- 
plemental feeding or drawdowns were 
conducted and for the 5 years of the feed- 
ing-drawdown program. From these aver- 
ages it was obvious that a large difference 
occurred in the yield of bluegills, so large 
a difference that the average bluegill 
yield during the feeding-drawdown (f-d) 
period was about six times that for the 
pre-experimental 2 years. This difference 
occurred partly because the fish popula- 
tion was expanding in 1963-1964, and 
many of the bluegills were too small to 
interest anglers. This lack of interest in 
the small bluegills was further demon- 
strated by the light fishing pressure in 
those years (46 man-hours per hectare, 
or 114 man-hours per acre, in 1963; 56 
man-hours per hectare, or 138 man-hours 
per acre, in 1964), as annual fishing pres- 
sures below 125 man-hours per hectare 
per season (309 man-hours per acre) in- 
dicated the poor quality of the fishing. 
However, the eight boats available for 
angling were seldom, if ever, used to the 
maximum during August in any year. 
Annual fishing effort, 1963-1969, is 
shown in Table 3. 

Because the fish population was enu- 
merated in total at the fish census and 


Table 3.—Fishing effort, in man-hours per hec- 
tare and per acre, expended by fishermen during 
the seasons 1963-1969, inclusive, at Ridge Lake. 


Man-Hours Man-Hours 

Year Per Per 

Hectare Acre 
1963 46 114 
1964 56 138 
1965 98 242 
1966 103 254 
1967 107 265 
1968 107 264 
1969 102 252 


Vol. 31, Art. 1 


restocking in 1963 (at the beginning of 
the experiment) and in the fish census 
in 1970 (at the end of the experiment) 
and because the fishermen’s total catch 
was recorded each year, it was possible 
to show the population dynamics of each 
individual species during the 7-year 
period. 


Largemouth Bass 


Table 4 shows that the lake was re- 
stocked in the spring of 1963 with 2,270 | 
bass of less than 254 mm (10 inches) and 
116 that averaged more than 1.36 kg (3 
pounds) each. No bass were available 
in the 254-305-mm (10-12-inch) range, 
a situation that is inexplicable. In the 
following 7-year period, the catch con- 
sisted of 2,984 bass of less than 254 mm 
(10 inches), 962 bass of 254-305 mm 
(10-12 inches), and only 91 larger than 
305 mm (12 inches). The record shows 
that 59 bass ranging in weight from 1.4 
to 3.6 kg (3.0-8.0 pounds) were returned 
to the lake in 1963 and that 22 bass 
averaging 2.4 ke (5.35 pounds) and 47 
averaging 1.2 kg (2.65 pounds) were ex- 
posed in the 1970 census, 7 years later. 
Therefore, one must assume that Ridge 
Lake contained at least 10 bass weighing 
more than 2.25 kg (5 pounds) each and 
40 or more additional bass, each weigh- 
ing 1.1 kg (2.5 pounds) or more, through 


‘this period of years. In spite of this valid 


assumption, fishermen caught only 49 
bass as large as 1.1 kg (2.5 pounds) and 
only 2 larger bass, each weighing between 
2.7 and 3.2 kg (6 and 7 pounds). At 
the same time, they were catching and 
removing 2,984 bass smaller than 255 mm 
(10 inches) at rates between 300 and 600 
per season (fishermen were asked to bring 
in all bass regardless of size, but we know 
that some did not). 

In general, Ridge Lake bass popula- 
tions subjected to annual drawdowns over 
a period of years were composed of many 
small bass, a small number of very large 
ones, and relatively few of intermediate 
sizes. The thinning effect of the fall 
drawdown reduced the predation pres- 
sure on bass eggs and fry in the following 


EFFECTS OF FEEDING AND DRAWDOWNS ON FISHES 9 


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10 Inurnois NaturAL History SurvEY BULLETIN 


spawning season, and successive strong 
year classes were produced, some of which 
were later reduced by predation from a 
preceding year class. This cycle of pro- 
duction created by the drawdown caused 
severe competition and slow growth 
among the small bass and rapid growth 
among the few that survived the food 
competition, stranding, and predation of 
their first season and the fishing pressure 
of their second season. 

Severe predation upon small fishes oth- 
er than bass was indicated by changes in 
the population of lake chubsuckers in the 
7 years of this experiment. In 1963, 1,020 
lake chubsuckers were restocked, totaling 
110 kg (243 pounds) and averaging 108 
grams (0.24 pound) each. Only 232 
chubsuckers, weighing 44.7 kg (98.5 
pounds) and averaging 193 grams (0.42 
pound), appeared in the 1970 census. 


Vol. 31, Art. 1 


These fish were too large to be preyed 
upon by any but the very largest bass and 
catfish; none smaller had managed to 
survive. 


Bluegills 


A total of 4,492 bluegills, mostly within 
the 100- to 140-mm (4- to 5.5-inch) 
length range were returned to Ridge Lake 
after the spring census of 1963 (Table 
5). These bluegills constituted a popula- 
tion of 693 per hectare (281 per acre). 
With such a small population, very few 
were caught in 1963, but by 1964 enough 
bluegills were present to increase food 
competition and improve the catch. Large 
catches of bluegills exceeding 150 mm (6 
inches) in total length were made in each 
year from 1965 to 1969, inclusive, or 
throughout the f-d period (Table 5 and 
Fig. 2), and quite large numbers of small 


Fig. 2.—Fishermen returning to the laboratory pier with large catches of bluegills. 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWpDOWNSs ON FisHes_ 11 


Table 5.—Population dynamics of bluegills in Ridge Lake, 1963-1970, including the feeding- 


drawdown period (1965-1969). 


Smaller Than 152 mm (6 inches) 


Year 


Num- Weight in Average Weight 


152 mm or Larger 


Num- Weight in Average Weight 


ber Kilograms in Grams ber Kilograms in Grams 
1963 restocking 3,526 65.6 18 966 56.9 59 
Anglers’ catch 
1963 272 15.2 56 86 Uae 87 
1964 141 4.6 33 1,146 127.6 111 
1965 977 53.9 55 2,282 294.5 129 
1966 1,670 o3n9) 56 3,916 448.7 114 
1967 509 Pape. 53 4,007 554.0 138 
1968 1,880 79.8 42 2,285 336.0 147 
1969 2,126 100.2 47 2,754 331.7 120 
Total catch Vicsay fis) 374.8 : 16,476 2,100.0 : 
1970 census 7,967 306.2 38 1,579 197.5 125 


bluegills were caught by fishermen in 
1966, 1968, and 1969. Bluegills of de- 
sirable sizes averaged 127 grams (0.28 
pound) each. 

Of some interest is the fact that the 
fishing pressure was nearly the same dur- 
ing each of the f-d years (1965-1969, 
inclusive) (Table 3), in part a reflection 
of the goodness of the fishing. 

When the lake was drained in April 
1970, it contained about 9,500 bluegills 
larger than 75 mm (3 inches). Most of 
these fish were within the 100- to 140-mm 
(4.0- to 5.5-inch) length range; however, 


Table 6.—Population dynamics of warmouths 
drawdown period (1965-1969). 


Smaller Than 152 mm (6 inches) 


about 1,600 were larger than 150 mm (6 
inches), and many were more than 178 
mm (7 inches). 

In 7 years, fishermen had taken 16,476 
large bluegills (Table 5) and 7,575 small- 
er ones. The large bluegills averaged 178 
mm and 127 grams (7.0 inches and 0.28 
pound) each; the small ones, 49 grams 


(0.11 pound). 


Warmouths 


More than 4,500 warmouths were taken 
in the 1963 draining, and most of them 
were less than 150 mm (6 inches) in 


in Ridge Lake, 1963-1970, including the feeding- 


152 mm or Larger 


Year 


Num- Weight in Average Weight 


Num- Weight in Average Weight 


ber Kilograms in Grams ber Kilograms in Grams 
1963 restocking 1,330 28.3 21 5 Paik 220 
Anglers’ catch 
1963 49 2.4 49 are cee 
1964 21 1.4 67 87 11.3 130 
1965 211 15.8 75 279 34.0 122 
1966 30 1.9 63 74 18.6 251 
1967 8 0.4 50 110 14.9 135 
1968 65 3.0 46 138 18.6 135 
1969 34 1.8 53 44 Bra) 130 
Total catch 418 26.7 732 103.1 ys 
1970 census 422 21.9 52 134 17.5 130 


12 Intrvors NaTturAL History SurvEY BULLETIN 


length. Approximately 1,330 of the larg- 
er ones were restocked following the cen- 
sus (Table 6). The catch of both large 
and small warmouths was quite insig- 
nificant, 732 large warmouths and 418 
small ones being brought in by fishermen 
in 7 years. 

There were 556 warmouths in the cen- 
sus of April 1970, and only 134 of these 
were more than 150 mm (6 inches) in 
total length. 

That the total number of warmouths 
was reduced during this period suggests 
that the drawdown was not obviously ef- 
fective in stimulating an increase in the 
warmouth population. Warmouths have 
been observed to eat the pelleted food, 
and they presumably grow well on it. 
In 1966, for example, the 74 warmouths 
that comprised that part of the catch 
which exceeded 150 mm in total length 
averaged 251 grams (0.56 pound) each, 
or more than twice as much as the aver- 
age weight of large bluegills caught in 
that year. 

The warmouths, as is usually the case 
in Illinois (Larimore 1957:70), in com- 
petition with largemouth bass and blue- 
gills in Ridge Lake, have contributed very 
little to the fish population and to the 


Table 7.—Population dynamics of channel catfish in Ridge Lake, 


feeding-drawdown period (1965-1969). 
a few young in 1963 or 1964. 


Vol. 31, Art. 1 


anglers’ yield in every phase of experi- 
mental fish management that has been 
tested, including the f-d program. 


Channel Catfish 


Eleven large channel catfish appeared 
in the census of 1963 and were returned 
to Ridge Lake (Table 7). These were 
all very large fish, averaging 4 kg each 
(8.9 pounds). No catfish was caught by 
fishermen until 1965 when two small fish 
weighing 172 grams each (0.38 pound) 
were taken. These were believed to rep- 
resent survivals from a spawn produced 
in the lake in 1963. Others of this year 
class probably survived because some cat- 
fish were taken each year, 1966-1969, in- 
clusive (Table 7). These catches prob- 
ably represented this same year class, be- 
cause their average size moved progres- 
sively upward with each successive sea- 
son: 1966, 1 ke; 1967, 2.3 kg; 1968, 3 kg; 
1969, 3.4 ke (2.3, 5.1, 6.6, 7.5 pounds). 

It appears improbable that any of the 
11 catfish returned to the lake in 1963 
were caught. However, it is likely that 
the 32 large catfish caught by fishermen, 
1966-1969, and the 22 large fish taken 
in the census of 1970 were all survivors 
from a year class of fish produced in 


1963-1970, including the 


The large catfish restocked in 1963 apparently produced 


Smaller Than 304 mm (12 inches) 


Year 


Num- Weight in Average Weight 


304 mm or Larger 


Num- Weight in Average Weight 


ber Kilograms in Kilograms ber Kilograms in Kilograms 
1963 restocking 11 44.8 4.07 
Anglers? catch if 
1963 
1964 aoe sabes ae 
1965 2 0.3 0.15 me re ine 
1966 15 15.4 1.03 
1967 3 7.0 2.33 
1968 Beye Tame ai 1 3.0 3.00 
1969 110% 52.1 0.47 13 44.2 3.40 
Total catch 112 52.4 32 69.6 
1970 census 
Offspring of 
1963 restocking Byes ere ates 22 86.3 3.92 
1969 stocking 805 90.9 0.11 650 428.8 0.66 
Total 805 90.9 0.11 672 515.1 


® 1,000 channel catfish 203-330 mm (8-18 inches) long were stocked in Ridge Lake on May 1, 1969 from 
the Sam A. Parr Fisheries Research Center, Marion County, IIl., and 1,000 more of the same size range 
were stocked on October 21 and 22, 1969 after the fishing and feeding season. 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FisHEs_ 13 


Ridge Lake, probably in 1963. These 
channel catfish were able to survive in 
1963 because the fish population was well 
below the carrying capacity of the lake 
in that year, and predation pressure was 
probably low. 


THE EFFECTS OF EARLY FALL 
DRAWDOWNS IN REDUCING 
BLUEGILL NUMBERS IN RIDGE LAKE 


With stable water levels, the bluegill 
population of Ridge Lake increases rapid- 
ly in total number with each successive 
spawning season and with apparently little 
regard for the number of largemouth bass 
present. In 1949 and 1950 the bass pop- 
ulation of Ridge Lake was exceptionally 
large, and_-no bluegills other than those 
that remained in pockets of the old stream 
channel during the 1949 census were left 
in the lake. Yet the population found 
in the 1951 census amounted to 51,963 
bluegills larger than 65 mm (Table 8). 
The 66,600 bluegills that appeared in the 
1947 census originated from 61 large blue- 
gills returned to the lake following the 
1945 census. If the period between cen- 
suses is longer than 2 years, the numbers 
of bluegills become larger. For example, 


the 3-year period 1960-1962 started with 
zero bluegills and a dry lake basin over 
the winter of 1959-1960. A few bluegills 
appeared in 1960 from an unidentified 
source. These multiplied in competition 
with 4,500 hybrid sunfishes and a bass 
population that was building up to 6,000 
small fishes. In 3 years the bluegills 
numbered 85,500. 

Still larger numbers of bluegills were 
present after the four growing seasons 
with stable water levels, beginning in 
March 1956 and continuing until October 
1959. After the 1956 census, 1,008 blue- 
gills were restocked, and in the 1959 cen- 
sus the bluegill population was 92,700. 
It is impossible to suggest how much this 
population might increase numerically, 
but it seems apparent that in a relatively 
short period, say 7-10 years, the bluegills 
would become so dominant as to curtail 
the success of bass reproduction. Within 
the range of bluegill numbers (and time) 
shown in Table 8, there was no evidence 
of a reduction of bass numbers; in fact, 
the 6,200 bass exposed in the 1963 census, 
when 85,500 bluegills were present, was 
the largest population of small bass ever 
recorded for Ridge Lake. 

The effects of the drawdowns on the 


Table 8.—Total numbers and total weights in kilograms of fishes collected in several draining 
censuses at Ridge Lake when bluegills were present and when water levels were stable for two or 


more seasons prior to the census. 
fall drawdowns. 


Similar data are presented for censuses following 2—5 years of early 


All Fishes Largemouth Bass Bluegills Other Fishes 
Year of SE 
Census Num- Weightin Num- Weightin Num- Weightin Num- Weight in 
ber Kilograms ber Kilograms ber Kilograms ber Kilograms 
Stable water 
1947 69,801 2,092.5 2,509 257.0) (66,5629) 1577-8 663 20nd, 
1951 54,574 1,336.8 1,510 407.5 51,963 858.4 1,101 70.8 
1959 97,312 1,906.0 Ou 240.0 92,669 1,246.5 2,289 4194 
1963 995791) S17 8o6).5 6,218 39928) 289,928) 11,0439 8,045 453.2 
Average 80,370 1,798.0 S148: 316.1 74,197 1,181.6 3,024 300.3 
Fall drawdowns 
19538 10,377 901.0 1, 964 204.8 7,476 449.9 937 247.1 
1956» 20,308 1,538.7 2,242 289.3 17,180 924.6 886 324.8 
1970° 14,234 1,440.0 2,420 244.6 9,546 503.8 2,268 691.7 
Average TAS OTE M2932. 2,209 246.2 11,401 626.1 1,364 421.2 


® Drawdowns of 4.6 meters (15 feet) in 1951 and 1952. 
> Drawdowns of 3.0 meters (10 feet) in 1953, 1954, and 1955. 
© Drawdowns varying between 3.0 and 4.6 meters (10 and 15 feet) in 1965, 1966, 1967, 1968, and 1969. 


14 Intino1is NaTurAL History SuRvEY BULLETIN 


bluegill populations become evident when 
the bluegill numbers at the bottom of 
Table 8 are compared with those at the 
top. Also, a direct relationship apparent- 
ly exists between the severity of the draw- 
down and the extent of bluegill popula- 
tion reduction, as indicated by the 1953 
and 1956 census figures. 


During a drawdown, small bluegills are 
more vulnerable to stranding and preda- 
tion than are bluegills larger than a cer- 
tain minimum size (25-100 mm). The 
larger fishes may live through several 
drawdowns, while relatively few of the 
small ones survive. The reduced popula- 
tion remaining in Ridge Lake after a 
drawdown (10,000—20,000 fishes instead 
of 50,000—100,000) becomes an expand- 
ing population in the refilled lake, with 
plenty of available food and space for 
reproduction and growth in the growing 
season following a fall drawdown. 


Table 8 shows no large differences be- 
tween the numbers of largemouth bass 
with and without drawdowns. Draw- 
downs are associated with successful re- 
production of largemouth bass during the 
following spawning season, but when a 
drawdown is scheduled for every fall the 
young bass of each spawning season may 
be decimated by yearling bass from the 
previous year class. The discovery that 
drawdowns are almost always followed 
by successful bass fry production and the 
survival of these little fish beyond the 
size subject to predation by bluegills sug- 
gests a “surefire” method of producing a 
new year class of bass when a stunted 
bluegill population has been curtailing 
all bass reproduction. It is evident that 
annual drawdowns, with or without sup- 
plemental feeding, do not result in bass 
populations with superior potential for 
bass fishing although they probably should 
not be considered below average. 


A COMPARISON OF THE EFFECTS OF 
VARIOUS MANAGEMENT 
TECHNIQUES ON ANGLING YIELDS 


To make a comparison of management 
techniques, data from several fishing sea- 
sons directly affected by these techniques 


Vol. 31, Art. 1 


were selected and averaged (Tables 9 
and 10). For example, several years in 
the biennial draining-and-culling period, 
1941-1951, were characterized by large 
catches of largemouth bass. The year 
1948 was selected because of the alternate 
years in which the lake was not drained, 
it was the year of the largest catch of bass. 


In the first period of drawdown studies, 
1951-1956, catches for the years 1951 and 
1953 were omitted because they followed 
spring censusing operations in which the 
lake was completely drained and selected 
fish were returned. These operations also 
affected all other aquatic biota. While 
the fish returned to the lake after the 
1953 census must have been influenced 
by the drawdowns of the fall seasons of 
1951 and 1952, the fish population re- 
turned was probably more a reflection of 
the draining and censusing operation than 
of the drawdowns. The draining opera- 
tion of 1956 again upset the replacement 
fish population. It marked the beginning 
of the “steady-state” period which lasted 
from May 1956 to October 1959. Only 
the years 1957, 1958, and 1959 were used 
to represent this period of stable water 
levels. 


In the f-d period the years 1965-1969 
were included because, as mentioned pre- 
viously, feeding was begun during the 
summer of 1965 and the first drawdown 
was made in the fall of 1965. 


In Table 9, statistics are shown for a 
comparison of the yields of the four major 
species of fishes in Ridge Lake for the 
several periods mentioned above. The 
average yield of bass of 18 kg per hectare 
(16.1 pounds per acre) for the f-d period 
is lower than that of any of the other 
periods shown and was exceeded by nearly 
twice this average in 1948, one of the best 
bass fishing seasons. 


It seems safe to assume that annual fall 
drawdowns at Ridge Lake did not result 
in the production of large numbers of 
desirable-sized bass. Bass fishing was con- 
siderably better in the period when we 
drained the lake every 2 years and re- 
moved bass smaller than 200-255 mm 
(8-10 inches) along with large numbers 
of smaller bluegills. Under this culling 


Jan., 1973 BENNETT ET AL.: EFFects oF FEEDING AND DRAwpOWNS ON FisHEs 15 


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Average 


Number 
Per 
Acre 


Bluegills 
Weight 


in Grams 


Average 


Per 
Hectare 


Number 


Weight 
in Pounds 


emouth bass and bluegills taken by anglers in 1948, a very successful 
Average 


and by supplemental feeding and drawdowns. 


Largemouth Bass 
Average Number 
Per 


Per 
Hectare 


Number 


Average number per unit of lake surface and average weight of larg 


year for bass fishing, and in years affected by drawdowns only, by stable water levels, 


Year(s) 


Table 10. 


Inuinors NaturAL History Survey BULLETIN Vol. 31, Art. 1 


method, we selected for fast-growing bass 
and removed the slow-growing ones. 
ONO This method also stimulated the produc- 
oocso tion of large new year classes of bass at 
2-year intervals. 
It is also evident from the data in Table 
XSNS 9 that the f-d operation did not improve 
the catch of warmouths and channel cat- 
fish. This was due in part to the relative- 
ly small numbers of both species. We 
S made no direct observations on whether 
warmouths were eating the pelleted food 
although they readily learned to eat it 
in the laboratory. Channel catfish fed 
well on “Splash,” as was indicated in 
Table 7 by the large annual increases in 
the average weight of the catfish caught 
(except those stocked in 1969), 1966— 
1969, inclusive. However, so few were 
in the lake that their weight per hectare 
was small. 


Weight 
in Pounds 
1 
2 
1 
2 


801 
416 
797 
692 


0.38 


Table 10 shows average numbers of 
bass and bluegills caught per unit of lake 
surface and their average individual 
weight under the several methods of man- 
agement. With both largemouth bass 
and bluegills, apparently a negative rela- 
tionship exists between the average num- 
ber of fish caught and their average size. 
If one may assume a positive relationship 
between the number of fishes available 
in any season and the number caught by 
anglers, one may also assume that, in 
years when the fish population is relative- 
ly small, each individual fish may have 
plenty of food available and therefore 
may grow rapidly and attain a large size. 
Thus, because the population is relatively 
small, each individual is subjected to little 
competition for food and space. If re- 
cruitment does not greatly increase the 
population, it is reasonable to assume that 
the average size of the individuals in this 
population will be large. Conversely, 
overproduction, the survival of new year 
classes, and the consequent competition 
for food and space will result in fishes 
of small average size. One can therefore 
assume a relationship between the catch 
of fishes and the average size of those 
fishes although so many variables are in- 
volved that the relationship may be quite 
obscured. 


Acre 
42 


Weight 


in Grams 
361 
137 
148 
175 


89 
163 
138 
103 


Drawdowns only; 1952, 1954, 1955 
Stable water; 1957, 1958, 1959 
Feeding and drawdowns; 1965-1969 


Good bass season, 1948 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FisHES_ 17 


THE EFFECTS OF SUPPLEMENTAL 
FEEDING AND DRAWDOWNS ON THE 
CONDITION OF LARGEMOUTH BASS 

AND BLUEGILLS 

We believed that the effects of supple- 
mental feeding of the fishes of Ridge Lake 
would become evident through changes 
in the growth rate and in the relative 
plumpness of the fishes. The procedure 
of allowing two seasons to pass after the 
restocking of the lake before any experi- 
mental management or feeding was be- 
gun gave results from those 2 years that 
we could compare with results from 5 
years of f-d operations. Also, results from 
the 5-year f-d period could be compared 
with results from previous Ridge Lake 
studies (Bennett 1954a; Bennett et al. 
1969): 5 years of drawdowns without 
feeding, 10 years of biennial draining of 
the lake and culling of the fishes, and 4 
years of stable water levels. 

It is convenient to begin by comparing 
the condition of bass and bluegills col- 
lected in 1963 and 1964, when no supple- 
mental feeding or drawdowns were con- 
ducted, with collections made in 1965— 
1969, inclusive, when the fish were fed 
daily during the summer and the lake 
level was dropped 3 or more meters each 
fall and held down as long as the weather 
was warm. 

Fishermen’s catches at Ridge Lake were 
measured as total lengths in tenths of 
inches and weights in hundredths of 
pounds. Therefore, it was convenient to 
use the Index of Condition, C, formulated 
by Thompson & Bennett (1939:16-17) 

_ W 10,000 
c= Soe 
in which W is weight in pounds and L 
is length in inches. 

To interpret results from this formula, 
one must know that bluegills showing an 
index of condition, C, of 6.0-7.0 are in 
poor flesh; those showing a condition of 
7.18.0 are in the range of average plump- 
ness; and those showing a condition of 
8.1 or above are obese and usually show 
internal fat deposits. 

The largemouth bass, having more 
elongated shapes than the shapes of blue- 
gills, have a lower condition index range. 


In bass, condition indices of 3.5—4.5 are 
related to a thin body; bass in the range 
of 4.6-5.5 are about normal; and those 
within the range of 5.6 or higher are 
obese. Bluegills are known to have an 
annual cycle of condition (Bennett, 
Thompson, & Parr 1940:6), with condi- 
tion being lowest during winter, gradual- 
ly rising in March and April, and reach- 
ing a peak in late May or early June at 
the beginning of the spawning season. 
During the long spawning season extend- 
ing throughout the summer, bluegill con- 
dition usually drops. Sometimes it rises 
in late August and early September, drop- 
ping again to the winter low. No annual 
condition cycle has been reported for 
largemouth bass. 

Average indexes of condition are shown 
in Table 11. Data were taken from creel 
cards recorded for all fishes by the first 
junior author when fishermen returned 
to the laboratory pier with their catches. 
Length-weight data from fishes caught 
by anglers during early June, the first 


Table 11.—Average index of condition, C, of 
monthly samples of largemouth bass and bluegills 
taken by fishermen from Ridge Lake during the 
summer fishing periods, 1963-1969. 


Month Largemouth Bass Bluegills 
and SSS 
Year Num- Average Num- Average 
ber C Value ber C Value 
June 1963 32 4.82 96 6.17 
July 1963 17 4.70 128 6.81 
August 1963 26 4.08 118 6.37 
June 1964 244 4.18 211 7.38 
July 1964 134 4.53 245 don 
August 1964 61 4.46 268 7.30 
June 1965 84 4.60 147 7.54 
July 1965 74 4.55 246 7.43 
August 1965 114 4.44 237 7.44 
June 1966 138 4.25 224 8.12 
July 1966 93 4.44 270 7.84 
August 1966 67 4.20 219 7.65 
June 1967 174 4.49 287 8.43 
July 1967 104 4.92 246 8.29 
August 1967 144 4.77 282 8.11 
June 1968 185 4.77 318 8.51 
July 1968 92 4.58 337 7.57 
August 1968 119 4.53 224 vey 
June 1969 183 4.09 273 7.51 
July 1969 82 4.57 224 7.81 
August 1969 81 4.71 275 7.96 


18 


part of July, and the first part of August 
were used, and the numbers of fish rec- 
ords ranged between 17 and 337, depend- 
ing on the numbers available. 

Indexes of condition were calculated 
for bass within the length range of 173- 
399 mm (6.8-15.7 inches) and for blue- 
gills within the range of 147-246 mm 
(5.8-9.7 inches). Fishes were separated 
into 25-mm (l-inch) length groups (e.g., 
147-172 mm, 173-198 mm, etc.) so that 
any great variations in the relative plump- 
ness of these length groups would be ex- 
posed. Bluegills were fairly consistent in 
condition within the size groups recorded, 
but bass more than 12 inches long were 
heavier in proportion to their length than 
were shorter bass. Indices of condition 
for both bass and bluegills caught by an- 
glers at any one time (within a period of 
a few days) were quite uniform for their 
species, although occasionally a few in- 
dividuals varied widely. The bass con- 
dition data in Table 11 emphasizes the 
fact that in most months the bass aver- 
aged slightly below normal, or average, 
plumpness. In general, all bass were thin, 
but some were more so than others. 

During the summer of 1963 the blue- 


9.0 
Length Group 


Millimeters Inches 
147-170 5.8 -6.7 
171-196 ------ 6.8-7.7 
97> 22) ———--—— 77.8) — Sir 


Wtd. Avg. 


@ 
o 


INDEX OF CONDITION 


7.0 


Inurnois NaturAaL History SurveEY BULLETIN 


“FAT” CONDITION 


Vol. 31, Art. 1 


gills were thin (Fig. 3), but they improved 
to average plumpness in 1964 before any 
supplemental feeding was begun. Appar- 
ently, the feeding program in 1965 was 
not reflected in the condition of the blue- 
gills in that year although they were, on 
the average, somewhat plumper than they 
were in 1964. The effects of supplemen- 
tal feeding were evident in the collections 
for June in 1966, 1967, and 1968 (Fig. 
3) when the average C values for blue- 
gills were in the fat category. The con- 
dition cycle for this species was evident 
in most years. Bluegill plumpness in July 
and August was usually lower, on the 
average, than it was in June, with mid- 
and late-summer condition falling below 
the fat classification in all years except 
1967. In 1968, bluegills were in fat con- 
dition in June, and large bluegills again 
reached that level in August. In that 
same year, a high survival rate oc- 
curred among the small bluegills of a very 
large year class. This high rate of sur- 
vival became evident in 1969 when many 
bluegills were too small to interest anglers 
but were numerous enough to reduce the 
overall effect of the supplemental feeding. 
Fig. 4 shows the average condition for 


AVERAGE PLUMPNESS 


“THIN” CONDITION 


6.0 
| ama apol ael Fama Saal (fi=g-sileapa cl a aL Galion (= palin! 
J J A J J A J A J J A J J A J J A J J A 
1963 1964 1965 1966 1967 1968 1969 


Fig. 3.—Indices of Condition of three length groupings of bluegills and the weighted 
average for all groupings in summers of the f-d experiment. Supplemental feeding was begun 
in 1965, and the first drawdown of this program was conducted in the fall of 1965. 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FISHES 


Length Group 
9.0 


Millimeters Inches 
147-170 5.8 - 6.7 
171 -196 ————— 6.8 —-7.7 
(Ot S| SSS 
Wtd. Avg. —$—$$—$$$$_<$>_ 


INDEX OF CONDITION 


6.0 | | 
JUNE JULY 


1954 


AUGUST 


19 


“FAT” CONDITION 


AVERAGE PLUMPNESS 


“THIN” CONDITION 


| | | | 
JUNE JULY AUGUST 


1955 


Fig. 4.—Indices of Condition of three length groupings of bluegills and the weighted 
average for all groupings taken in 1954 and 1955 during a period of annual fall drawdowns 


but with no supplemental feeding. 


three length classes of bluegills taken 
from Ridge Lake in 1954 and 1955 when 
this fish population was being subjected 
to moderate annual fall drawdowns of 
about 3 meters (Bennett et al. 1969:16) 
but with no supplemental summer feed- 
ing. Bluegills showed an average condi- 
tion in the fat zone only in June 1954; 
during the rest of the 2-year span these 
fish were generally in high average condi- 
tion. As stated elsewhere, these fish did 
not appear to be plump. 


THE GROWTH OF BASS AND 
BLUEGILLS IN A FEEDING- 
DRAWDOWN PROGRAM 


Growth rates of largemouth bass and 
bluegills were estimated from scale analy- 
ses and from length, weight, and age 
data for fishes taken by anglers late in 
the summers of 1967 and 1968 when fish 
growth for the season was nearly com- 
plete. By averaging the total lengths of 
bass or bluegills separated into age classes 
on the basis of the number of annuli on 


selected scales, we were able to construct 
growth curves (Fig. 5, 6, and 7). 

The growth of largemouth bass was 
slow during the f-d period, particularly 
when compared with that of the period 
of biennial culling, 1941-1951 (Table 12 
and Fig. 5). In the 1941-1951 period 
the lake was completely drained five times 
at intervals of 2 years, and each time 
a census was made of the fish. The meth- 
od of culling the population after the 
census has been described (Bennett 
1954a:241). The data for the upper 
growth curve for bass (Fig. 5) were taken 
from Bennett (1954a:255). This curve 
shows that bass reached a useful size (254 
mm) in less than two growing seasons 
in the biennial culling period; in contrast, 
three complete growing seasons were re- 
quired to obtain 254-mm bass under the 
f-d program. Also, under this latter pro- 
gram there appeared to be a scarcity of 
305- to 380-mm (12- to 15-inch) bass 
(Table 12). Quite obviously, from the 
standpoint of growth rate, the f-d pro- 
gram cannot be recommended for bass. 


20 Intinois NATuRAL History SurRvEY BULLETIN Vol. 31, Art. 1 


400: 


o 
° 
co 


Useful Size 


‘Annual Drawdowns and Feeding 


8 
Total Length in Inches 


Total Length in Millimeters 


~4 


T 
4 


a4 


6 7 8 


5 
Year of Life 


Fig. 5.—Growth rates and annual length increments of largemouth bass under a system 
of biennial lake draining and culling of small bass and bluegills and under the f-d program. 


Table 13 and Fig. 6 and 7 show com- gram of drawdowns without feeding and 
parisons of the growth rates of bluegills under conditions brought about by stable 
under the f-d program and under a pro-_ water levels. 


Useful Size 


Total Length in Millimeters 
Total Length in Inches 


Year of Life 


Fig. 6.—Growth rates and annual length increments of bluegills under a system of fall 
drawdowns without feeding and under a system combining feeding with fall drawdowns. 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FisHES 21 


200: 


Fall Drawdowns with Feeding ye 


z 


YE 


__ 
—— 


A, , 
ee Useful Size 


Total Length in Millimeters 


Pee ae 


Dae 
a A stable Water Levels with No Feeding 


ed 


Total Length in Inches 


be 


3 
Year of Life 


Fig. 7—Growth rates and annual length increments of bluegills during the f-d program 
and under stable water levels (1956-1959, inclusive). 


Bluegills grew almost as fast when sub- 
jected to annual fall drawdowns with no 
supplemental feeding as they did under 
the program of supplemental feeding and 
fall drawdowns (Fig. 6). In both pro- 
grams useful sizes were attained early in 
the third summer of life. Under the f-d 
program, bluegills 4 and 5 years old aver- 
aged more than 200 mm (8 inches) in 
length; however, the average size of the 
4-year-old bluegills subjected to draw- 
downs alone was less than 200 mm, and 


there were too few 5-year-olds to give a 
significant average. Whether supplemen- 
tal feeding might have been a factor in 
slowing the mortality rate of the larger, 
older fish can only be conjecture at this 
time. 

With stable water levels at Ridge Lake 
in 1956-1959, bluegill numbers increased 
rapidly, and more than three growing 
seasons were required for these fish to 
average 150 mm (Fig. 7). None was 
able to attain a length greater than about 


Table 12.—Average total lengths in millimeters and inches of 405 largemouth bass taken at, 


or approximately at, the ends of the growing seasons in 1966, 1967, and 1968 during the feeding- 
drawdown period, and average total lengths of largemouth bass taken under similar circumstances 
in the 1941-1949 period of biennial draining and culling (from Bennett 1954a:255). 


Unit of 


Measurement 


Millimeters 
Inches 


Millimeters 
Inches 


Average Total Length At or Near End of Indicated Growing Season 


Ist 2nd 3rd 4th 5th 6th 7th 8th 


Feeding-Drawdown Period, 1965-1969 
201 244 287 318 358 
Tsk) 9.6 HES 12.5 Jorn 


Biennial Culling Period, 1941-1949 
206 262 320 348 363 424 442 478 
8.1 10.3 12.6 13.7 14.3 16.7 17.4 18.8 


22 Inurnois NatruraL History Survey BuLLeTIN Vol. 31, Art. 1 


Table 13.—Average total lengths in millimeters of bluegills captured and aged near the ends 
of the growing seasons of 1967 and 1968 during the period of supplemental feeding and annual 
fall drawdowns, of 1958 and 1959 during the period of stable water levels, and of 1955 during the 
period of annual fall drawdowns without feeding. 


Annual length increments are also shown. 


Period Number 
Supplemental feeding and fall 
drawdowns (1967 & 1968 collections) 316 
Length increment 
Stable water levels 
(1958 & 1959 collections) 326 


Length increment 


Annual fall drawdowns and no 
feeding (1955 collection) 
Length increment 


112 


Year of Life 
Ist 2nd 3rd 4th 5th 
83 147 185 205 221 
83 64 38 20 16 
51 119 146 161 169 
51 68 27 15 8 
48 137 170 188 
48 89 33 18 


Fig. 8.—A 254-mm (10-inch) bluegill from Ridge Lake weighing 499 grams (1.1 pounds). 
This bluegill was very fat and had an Index of Condition of 12. 


165 mm even though many reached the 
age of 5 years. Differences in the rates 
of growth of bluegills in Ridge Lake un- 
der differing systems of management ap- 
parently are related to the amount of 
available food and space per individual 
fish. 

If abundant food and space are avail- 
able, bluegill size must be limited by the 
length of life of this species and its maxi- 
mum genetic growth potential. Few blue- 
gills in central Illinois live longer than 5 
years and genetically the bluegill is a rela- 
tively small fish. Thus, in combining a 
supplemental feeding program with draw- 
downs, we are, in theory, projecting a 


management technique for producing 
bluegills of exceptional sizes (Fig. 8). 


DISCUSSION 


The decision to give warmwater fishes 
a supplemental source of food in a man- 
agement program to improve sport fishing 
depends, first of all, on cost as it is related 
to benefits. However, data about yield 
improvement may not be available to the 
individual fisherman, and he may judge 
the fishing quality by what he himself 
catches. Such judging was done by fish- 
ermen using Ridge Lake during our ex- 
periment. 


Jan., 1973 BenNetr er at.: Errects or FEEDING AND DRAWDOWNSs ON FisHES 23 


In waters open to public fishing, usu- 
ally no attempt is made to harvest more 
of the available crop of fishes than may 
be taken by angling; therefore, unless the 
benefits are readily observable through 
an improved rate of catch or improved 
sizes of the fishes caught, or both, fisher- 
men may consider the program a waste 
of money. As has been mentioned, the 
cost of the pelleted fish food used in this 
experiment was about 6 cents per pound, 
and the feeding rate was 2 pounds of 
food per acre per day. Thus, the daily 
cost was 12 cents per acre, or $1.92 per 
day for 16-acre Ridge Lake. The feed- 
ing program required 1.5 tons of food 
per year at about $120—-$130 per ton, 
representing a seasonal cost of $11.25— 
$12.19 per acre, or $27.80-$30.12 per 
hectare. 

Schmittou (1969:312-313) used 2,896 
pounds of food costing $159.28 in l-acre 
Pond T, over two growing seasons and 
8,766 pounds of food costing $482.13 in 
Pond T, (surface area, 3.5 acres) for five 
growing seasons. Thus, the food costs 
per acre per season were $79.64 for Pond 
T, and $27.55 for T,. These costs may 
be compared with $11.25—$12.19 per acre 
per season for the food used at Ridge 
Lake. 

The results of this f-d program were 
evident to most, if not all, bluegill fisher- 
men in furnishing them with (i) larger 
bluegills, (ii) fatter bluegills, and (iii) 
a larger total poundage of bluegills be- 
cause of the increased average weight 
of individuals. Greater numbers of blue- 
gills were taken by anglers in the period 
of stable water levels, 1956-1959, than 
were caught in the f-d period, but the 
fishes taken in the earlier period were 
hardly more than 152 mm (6 inches) in 
average total length and their average 
weight was less than 82 grams (0.18 
pound) each. Another benefit cited by 
fishermen was the improved flavor of the 
bluegills that fed on the prepared food. 

Schmittou (1969:318) fed “balanced” 
bass-bluegill populations in two “treat- 
ment ponds” while following the popu- 
lation changes in a control pond. In his 
feeding program without drawdowns, 
there appeared to be a gradual increase 


in the number of bluegills and a reduc- 
tion in the number of bass that eventually 
would have caused a severe slump in bass 
fishing. In the Ridge Lake experiment 
the annual fall drawdowns also upset the 
bass population dynamics by indirectly 
causing the production of excessive num- 
bers of small bass, of which only a small 
percentage attained attractive sizes. 

The fact that largemouth bass will not 
learn to eat pelleted food unless given 
special training when very small (Snow 
1965:193 and 1968:145) reveals the use- 
lessness of simply broadcasting pelleted 
food in the management of bass for sport 
fishing. 

It is our opinion, after studying the 
movements of marked bluegills in various 
parts of the lake, that these fish are fairly 
sedentary, Le., their normal range of 
movements would not insure that indi- 
viduals from all parts of Ridge Lake 
would find a feeding area. Therefore, 
if pelleted food is to be made available 
to all of the Ridge Lake bluegills, it must 
be well distributed in shallow water in 
all parts of the lake. 

Once the bluegills have learned to feed 
on the pelleted food, they will ingest the 
amount distributed in a relatively short 
time. There was no particular reason 
for setting the amount! to be fed at 2 
pounds per acre per day except that we 
wished to have the food used as a supple- 
ment to the bluegills’ natural diet rather 
than as a substitute for it. However, the 
weight of bluegills in the 1970 census (78 
kg per hectare, or 69 pounds per acre) 
was about one-third of the maximum 
standing crop found in any past census 
(223 ke per hectare, or 200 pounds per 
acre). Thus, the daily quota of pelleted 
food represents about 2.5 percent of the 
total weight of bluegills. The effect of 
the drawdown in thinning the bluegill 
population was to increase the amount 
of food available to each fish from about 
1.0 to 3.0 percent of body weight per day. 

Various companies in the business of 
preparing and marketing animal foods 
usually have one or more types of pelleted 
fish foods. Generally these preparations 
consist of one or two grades of “trout” 
food and at least one, and sometimes two, 


24 Ixnurnors NaturAL History SuRVEY BULLETIN 


grades of “catfish” food. The trout foods 
are usually more expensive than those for 
catfish and are more “complete” fish diets 
because trout have been fed on prepared 
diets for more years than have catfish and 
more research has been done on their 
specific food requirements. 

During the development of techniques 
for the culture of channel catfish in cages, 
many fish were confined in relatively 
small spaces with little chance of obtain- 
ing a significant amount of natural food 
from the ponds in which the cages were 
floated. In this situation foods that were 
quite adequate for free-swimming channel 
catfish lacked certain food elements that 
the free catfish were able to forage from 
pond sources. Very little is known about 
the nutrition requirements of bluegills, 
but it seems reasonable to assume that 
the food requirements of caged and un- 
caged bluegills might be similar to those 
of catfish, i.e., caged bluegills would also 
require a more nearly complete diet than 
would free-swimming bluegills. 

The drawdown, when combined with 
a feeding program, is a money-saving op- 
eration because it limits the survival of 
successive year classes of bluegills to the 
numbers that can be utilized in the fish- 
ing program. Thus, the bluegills that 
survived the drawdowns always had an 
adequate supply of food for rapid growth 
and were abundant enough to satisfy the 
needs of anglers. 

Diminishing the aquatic habitat selects 
against the survival of smaller fishes in 
two ways. First, it forces the small fishes 
away from the shore shallows into open 
water with little or no protective cover 
in an environment that may be entirely 
strange to them. Here they become prey 
to larger fishes and other aquatic verte- 
brates and invertebrates such as crayfish. 
Second, it strands the smaller fishes in 
mats of settling rooted aquatic plants or 
in pockets of water in an uneven lake 
bottom which dry up in a few hours or 
days. The relative importance of these 
two phenomena in reducing the numbers 
of smaller fishes is conjectural, however, 
as much depends on the normal behavior 
patterns of the species involved. Those 
that tend to avoid very shallow water are 


Vol. 31, Art. 1 


decimated more by direct predation than 
by becoming stranded, and vice versa. 
To some pond owners a fall drawdown 
may present a problem because many 
ponds are not equipped with controlled 
outlets; therefore, drawdowns are impos- 
sible except through pumping or siphon- 
ing. Often when a pond is being built, 
the owner is operating on a limited 
budget, and the elimination of the drain 
outlet appears to be one way to cut ex- 
penses. This view is unfortunate because 


the use of the drawdown as a fish man- — 


agement technique has become well es- 
tablished (Bennett 1971:209-219). In 
fact, it is considered the single most im- 
portant operational procedure for elimi- 
nating overpopulation and stunting 
among the fishes in artificial ponds and 
reservoirs. 

The procedure is simply to open the 
outlet valve in the dam and lower the 
water level until the surface area of the 
lake or pond is between one-fourth and 
three-fourths that of the full lake, de- 
pending on how severe an effect is de- 
sirable. It is usually unnecessary to build 
a weir in the outlet to prevent the larger 
fishes from leaving the lake because they 
will not go out of the outlet until the 
water level becomes much lower than the 
level which results in a 75-percent reduc- 
tion in the lake surface area. Presumably, 
the larger fishes will not leave because 
they do not immediately recognize the 
danger of becoming stranded in the lake 
basin. If the outlet valve in the dam is 
at the lowest level of the lake basin and 
the lake is of the eutrophic type, a draw- 
down in summer or early fall will release 
oxygen-deficient water which also may 
contain methane, hydrogen sulfide, car- 
bon dioxide, and other anaerobic decom- 
position products. This water may be 
toxic to fishes immediately below the out- 
let and is certainly uninhabitable for fishes 
attempting to enter or leave the lake 
through the outlet. 

It is always advisable that those re- 
sponsible for the operation of a draw- 
down inspect the surviving population of 
fishes to make certain that the expected 
results are occurring. In some instances 
it may be necessary to supplement a draw- 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FISHES 25 


down with seining, as Hulsey (1957:286) 
arranged for in Nimrod Lake, to remove 
a large population of carp or buffalo or 
suckers that cannot be stranded and are 
too large to become prey to fishes or 
other aquatic animals. Even excessively 
large populations of stunted sunfishes may 
require supplemental cropping, particu- 
larly when they are living with relatively 
small numbers of large bass, of which 
there are too few to make impressive in- 
roads on the hordes of sunfishes. In both 
cases boat-mounted electric shockers may 
be used effectively in thinning the stunted 
or undesirable fishes, because during a 
drawdown the fishes are concentrated in 
such small areas that large numbers may 
be stunned within a relatively short time. 
Where seine hauls have been planned for 
reservoir basins before the water has been 
impounded, small-meshed drag seines may 
be used to help reduce the numbers of 
undesirable fishes. 

In managing a lake or pond for sport 
fishing, it is desirable to manage all im- 
portant species that are present. This 
we were unable to do with our f-d pro- 
gram. The program was effective in pro- 
ducing superior bluegills, and probably 
superior catfish, but it was not so for 
largemouth bass. Perhaps a severe fall 
drawdown each year is unnecessary, and 
one in 2 or 3 years with annual summer 
feeding might improve the size of the 
bass caught without greatly reducing the 
average size of the bluegills. In another 
direction, the maintenance of a popula- 
tion of channel catfish of at least 100 
per hectare might add a new interest for 
fishermen. 


SUMMARY 


1.—After a draining census in 1963, 
Ridge Lake was restocked with 2,386 
largemouth bass, 4,492 bluegills, 1,335 
warmouths, 11 channel catfish, and 1,020 
lake chubsuckers, making a total of 9,244 
fishes weighing 510.6 kg (1,125.5 
pounds). This was 78.8 kg per hectare, 
or 70.3 pounds per acre. In the census 
preceding this restocking this lake was 
found to contain 287 ke per hectare, or 
256 pounds per acre, almost four times 


the weight of fish returned to the lake. 
In 1969, 2,000 additional channel catfish 
were stocked. 
2—The population of fishes was al- 
lowed to expand for two growing seasons 
(1963 and 1964) without drawdowns or 
supplemental feeding but with the usual 
controlled public fishing during the sum- 
mer months. The hook-and-line catch 
in 1963 and 1964 was below the average 
for the preceding 20 years. 
3.—Beginning in late May 1965, and 
continuing each year during the 3 sum- 
mer months, 1965 through 1969, the fish 
were fed daily on a commercial pelleted 
fish food (32 percent protein) at the rate 
of 2 pounds per acre per day. Food was 
spread in the shallows in all parts of the 
lake. The food cost was within the range 
of $27.80-$30.12 per hectare per season 
($11.25-$12.19 per acre per season). 
4.—Each year, beginning in September 
1965, the lake level was lowered: 
4.6 meters (15 feet) in 1965, leaving 
a surface area of 2.12 ha 
3.0 meters (10 feet) in 1966, leaving 
a surface area of 4.5 ha 
3.0 meters (10 feet) in 1967, leaving 
a surface area of 4.5 ha 
4.6 meters (15 feet) in 1968, leaving 
a surface area of 2.12 ha 
4.3 meters (14 feet) in 1969, leaving 
a surface area of 2.76 ha 
The level was maintained until the water 
temperature in the lake was about 13° C. 
(57° F.) in October, when the lake was 
allowed to refill. 
5—In March 1970, the lake was 
drained to make a census of the fishes. 
The lake contained 2,420 bass, 9,546 blue- 
gills, 556 warmouths, 1,477 channel cat- 
fish, 232 lake chubsuckers, and 3 fishes 
of other species, a total of 14,234 fishes 
weighing 1,440.0 kg (3,175.3 pounds). 
6.—The catch of largemouth bass dur- 
ing the seasons 1965-1969, inclusive, was 
composed mostly of small fish. The f-d 
program resulted in the production of ex- 
cessive numbers of small bass but gener- 
ally did nothing to improve bass fishing. 
7.—The fishermen’s catch included 
more than twice as many large bluegills 
(152 mm or longer) as it did smaller 
ones during the 1965-1969 period. Blue- 


26 Inuinois NaruraL History Survey BuLLETIN 


gills of desirable sizes averaged 127 grams 
(0.28 pound) each. 

8—Neither warmouths nor channel 
catfish produced large hook-and-line 
yields because their numbers were always 
small. Channel catfish produced a small 
year class in 1963 or 1964, and this year 
class appeared in the catch in 1966-1969, 
inclusive. ‘The catfish stocked in 1969 
were too small to appear in the 1969 
catch. 

9.—During years when the water level 
in Ridge Lake remained fairly constant, 
bluegill numbers increased to 50,000 in 
one 2-year period and to 66,000 in an- 
other, to 86,000 in one 3-year period, and 
to 93,000 in a 4-year period. Annual fall 
drawdowns of 4.6 meters reduced the 
bluegill population to 7,500, those of 3.0 
meters to 17,000 bluegills, and the 4.3- 
meter drawdown reduced the population 
to 9,500 bluegills. These drawdowns ap- 
parently had little effect on largemouth 
bass numbers. 

10.—The average hook-and-line yield 
of bass in the 5 f-d years was only 18.0 
kg per hectare (16.1 pounds per acre). 
This yield was below the average for 3 
drawdown years (1952, 1954, and 1955) 
and 3 stable water level years (1957, 1958, 
and 1959). The average bluegill yield un- 
der the f-d program was 71.6 kg per hec- 
tare (63.9 pounds per acre), higher than 
the catch in any other period. 

11.—The average index of condition of 
largemouth bass in the f-d period was 


Vol. 31, Art. 1 


slightly below normal. Average bluegill 
condition was “fat” in June of all f-d 
years except 1965 and 1969. Usually the 
average bluegill index of condition was 
lower in July and August, which followed 
a previously observed condition cycle for 
that species. The condition of bluegills 
in 1954-1955 with fall drawdowns, but 
without supplemental feeding, was “fat” 
in June of 1954 but only reached “high 
average” plumpness for July and August 
of 1954 and for all of the summer of 1955. 

12._Largemouth bass growth was — 
slower during the f-d period than during 
the period of biennial lake draining and 
culling of the fish population. Bluegills 
grew somewhat faster during the f-d peri- 
od than they did during the program of 
drawdowns without feeding. They ap- 
peared to live longer during the f-d peri- 
od and therefore attained larger sizes. 
They grew much faster under the f-d 
program than they did when water levels 
were stable. 

13.—The pelleted food for the f-d pro- 
gram cost about 12 cents per acre per 
day, or about $11.25—$12.19 per acre 
per season. Fishermen were enthusiastic 
about the program because they were 
able to catch larger and fatter bluegills, 
and they believed that the pelleted food 
improved the flavor of these fish. Feed- 
ing bluegills without fall drawdowns 
would probably be wasteful because the 
bluegill population would expand faster 
than the food supply. 


LITERATURE CITED 


BENNETT, GeorGE W. 1954a. Largemouth bass 
in Ridge Lake, Coles County, Illinois. IIli- 
nois Natural History Survey Bulletin 26(2) : 
217-276. 

-. 1954b. The effects of a late-summer 

drawdown on the fish population of Ridge 

Lake, Coles County, Illinois. North Ameri- 

can Wildlife Conference Transactions 19: 

259-270. 


1971. Management of lakes and 
ponds. Van Nostrand Reinhold Company, 
New York. 375 p. 


, H. Wickuirre ApkINs, and WILLIAM 
F. Cuitpers. 1969. Largemouth bass and 
other fishes in Ridge Lake, Illinois, 1941- 
1963. Illinois Natural History Survey Bul- 
letin 30(1) : 1-67. 

, and Leonarp DurHam. 1951. Cost 
of bass fishing at Ridge Lake, Coles County, 
Illinois. Illinois Natural History Survey 
Biological Notes 23. 16 p. 


, Davi H. Tuompson, and Sam A, 
Parr. 1940. Lake management reports 4. 
A second year of fisheries investigations at 
Fork Lake, 1939. Illinois Natural History 
Survey Biological Notes 14. 24 p. 

Curwpers, Wituiam F. 1967. Hybridization 
of four species of sunfishes (Centrarchidae) . 
Illinois Natural History Survey Bulletin 
29(3) :159-214. 

Duruam, LEONARD, and GeorcE W. BENNETT. 
1949. Bass baits at Ridge Lake. Illinois 
Wildlife 4(2):10-13. 

, and 1951. More about bass 
baits at Ridge Lake. Illinois Wildlife 6(2) : 
Baik 

Hutsty, ANDREW H. 1957. Effects of a fall 
and winter drawdown on a flood control 
lake. Southeastern Association of Game and 
Fish Commissioners Proceedings for 1956, 
10: 285-289. 

Larimore, R. WELDON. 


1957. Ecological life 


Jan., 1973 BENNETT ET AL.: EFFECTS OF FEEDING AND DRAWDOWNS ON FisHES 27 


history of the warmouth (Centrarchidae). 
Illinois Natural History Survey Bulletin 
PIL) 21-83. 

Scumitrou, H. R. 1969. Some effects of sup- 
plemental feeding and controlled fishing in 
largemouth bass-bluegill populations. South- 
eastern Association of Game and Fish Com- 
missioners Proceedings for 1968, 22: 311-320. 

Snow, J. R. 1965. Results of further experi- 
ments on rearing largemouth bass fingerlings 


under controlled conditions. Southeastern 

Association of Game and Fish Commission- 

ers Proceedings for 1963, 17:191-203. 

1968. Production of six- to eight- 
inch largemouth bass for special purposes. 
Progressive Fish-Culturist 30(3) :144—-152. 

Tuompson, Davin H., and Georce W. BEN- 
NETT. 1939. Lake management reports 3. 
Lincoln Lakes near Lincoln, Illinois. _ IIli- 
nois Natural History Survey Biological Notes 
11. 24 p. 


INDEX 


Biennial draining, 10 years of, 1 
Bluegills, 1 
condition under feeding-drawdown pro- 
gram, 17 
fat condition in 1966, 1967 and 1968, 18 
growth under feeding-drawndown _pro- 
gram, 20 
laboratory feeding of, 2 
original stock in Ridge Lake, 1 
population dynamics, 1963-1970, 10-11 
reductions in numbers caused by draw- 
downs, 13-14 
weight in 1970 census one-third of maxi- 
mum standing crop, 23 


Cc 
Channel catfish, 1 
evidence of reproduction in Ridge Lake, 
12-13 
origin of 1969 stock, 3 
population dynamics, 1963-1970, 12-13 
stocking in 1969, 3 
Condition index, 17 
Controlled public fishing, 1 
Cost of feeding fishes, 6, 23 


D 


Draining census, 1 
Drawdowns 
begun in 1951, 1 
combined with supplemental feeding, 2 
effects in 1951, 1952, 1953, 1954, 1955, 
2 
effects on bluegill sizes, 2 
effects on largemouth bass, 10 
extent of, 1, 2, 6, 24 
five years of, 1 
money-saving operation, 24 
reduction in numbers of bluegills caused 
by, 6, 13-14 
supplemented by seining, 25 
“surefire” method of producing new year 
class of bass, 14 
value in general fish management, 24 


E 


ee ee A FN ON 4 


F 
Feeding 
amounts fed, 6, 23 
composition of fish food, 3, 5 
cost, 6, 23 
effects on bluegills, 5-6 
method, 5 
program, 3, 5-6 
times, 5 
Feeding-drawdown program 
condition of bluegills under, 17-18 
condition of largemouth bass under, 17 
fishermen’s opinions of, 22-23 
growth of bass under, 19-20 
growth of bluegills under, 20-21 
Fishing effort 
range of, 1963-1969, 8 
varies with quality of fishing, 8 
Fish food as a supplement to natural diet, 23 
Fish yields from angling, 1963-1969, 7-8 
Food, pelleted, in relation to total weight of 
bluegills, 23 
Foods available, 23-24 
Fish population, concentration of during 
drawdowns, 2 


G 
Growth 
of bass and bluegills under various sys- 
tems of management, 20-22 
of bluegills under f-d program, 20-22 
of largemouth bass under f-d program, 
19-21 
H 
Hybrid sunfishes 
laboratory feeding of, 2 
testing value of, 1 


l 
Ictalurus punctatus (Rafinesque), 1 
Index of condition 
definition of, 17 
for bluegills, 17 
for largemouth bass, 17 


L 


Lake chubsuckers 
orioinal stock in Ridge Lake. 1 


28 Inuinors NatrurAL History SuRvEY BULLETIN 


Largemouth bass 
condition under feeding-drawdown pro- 
gram, 17-18 
effects of annual drawdowns on numbers, 
13-14 
growth in f-d period, 19-21 
training to eat pelleted food, 23 
original stock in Ridge Lake, 1 
population dynamics, 1963-1970, 8-10 
Lepomis macrochirus Rafinesque, 1 
Lepomis gulosus (Cuvier), 1 


M 
Management methods, effects on Ridge Lake 
fishes, 14-16 
Micropterus salmoides (Lacépéde), 1 
Movements of bluegills related to feeding, 23 


Vol. 31, Art. 1 


Pp 
Public fishing, 1—2 


R 
Restocking after 1963 census, 3-4 
Ridge Lake studies, 1-2 
S 
Sam A. Parr Cooperative Fisheries Research 


Center, 3 
Stable water levels, period of, 1956-1959, 1-2 


WwW 


Warmouths, | 


contribute little to angling yield, 11-12 ° 


original stock in Ridge Lake, 1 
population dynamics, 1963-1970, 11-12 


a 


Some Publications of the ILLINOIS NATURAL HISTORY bait 
a 


BULLETIN 


Volume 30, Article 2—Dynamics of One-Spe- 
cies Populations of Fishes in Ponds Subjected 
to Cropping and Additional Stocking. By D. 
Homer Buck and Charles F. Thoits III. 
‘March, 1970. 97 p., 10 fig., bibliogr., index. 

Volume 30, Article 3.—Migrational Behavior of 
Mallards and Black Ducks as Determined 
from Banding. By Frank C. Bellrose and 
Robert D. Crompton. September; 1970. 68 
p., frontis., 25 fig., bibliogr., index. 

Volume 30, Article 4.-Fertilization of Estab- 
lished Trees: A Report of Field Studies. By 
Dan Neely, E. B. Himelick, and Webster R. 
Crowley, Jr. September, 1970. 32 p., fron- 
tis., 8 fig., bibliogr., index. 

Volume 30, Article 5.—A Survey of the Mussels 
(Unionacea) of the Illinois River: A Pollut- 
ed Stream. By William C. Starrett. February, 
1971. 137 p., 17 fig., bibliogr., index. 

Volume 30, Article 6.—Comparative Uptake 
and Biodegradability of DDT and Methoxy- 
chlor by Aquatic Organisms. By Keturah A. 
Reinbold, Inder P. Kapoor, William F. 
Childers, Willis N. Bruce, and Robert L. 
Metcalf. June, 1971. 12 p., frontis., 5 fig., 
bibliogr., index. 

Volume 30, Article 7—A Comparative Study of 
Two Components of the Poinsettia Root Rot 
Complex. By Robert S. Perry. August, 1971. 
35 p., frontis., 10 fig., bibliogr., index. 

Volume 30, Article 8—Dynamics of Condition 
Parameters and Organ Measurements in 
Pheasants. By William L. Anderson. July, 
1972. 44 p., frontis., 6 fig., bibliogr., index. 


BIOLOGICAL NOTES 


70.-An Ecological Study of Four Darters of the © 


Genus Percina (Percidae) in the Kaskaskia 
River, Illinois” By David L. Thomas. De- 
cember, 1970. 18 p., 11 fig., bibliogr. 

71.-A Synopsis of Common and Economic 
Illinois Ants, with Keys to the Genera 
(Hymenoptera, Formicidae). By Herbert 
H. Ross, George L. Rotramel, and Wallace 
E. LaBerge. January, 1971. 22 p., 27 fig., 
bibliogr. 

72.-The Use of Factor Analysis in Modeling 
Natural Communities of. Plants and Ani- 
mals. By Robert W. Poole. February, 1971. 
14 p., 14 fig., bibliogr. 

73.-A Distributional Atlas of Upper Mississip- 
pi River Fishes. By Philip W. Smith, Alvin 
C. Lopinot, and William L. Pflieger. May, 
1971. 20 p., 2 fig., 107 maps, bibliogr. 


List of available publications mailed on request 2a 


2 


74.-The Life History of the Slenderhead Dart- — 
er, Percina phoxocephala, in the Embarras — 
River, Illinois. By Lawrence M. Page and 
Philip W. Smith. July, 1971. 14 p., 10 fig., 
bibliogr. 

75.—Illinois Birds: Turdidae. By Richard R. 
Graber, Jean W. Graber, and Ethelyn L. 
Kirk. November, 1971. 44 p., 40 fig., bib- 
liogr. 4 

76.-Illinois Streams: A Classification Based on ~ 
Their Fishes and an Analysis of Factors Re- 
sponsible for Disappearance of Native Spe- © 
cies. By Philip W. Smith. November, 1971. — 
14 p., 26 fig., bibliogr. j 

77-The Literature of Arthropods Associated 
with Soybeans. I. A Bibliography of the: 
Mexican Bean Beetle, Epilachna varivestis 
Mulsant (Coleoptera: Coccinellidae). By 
M. P. Nichols and M. Kogan. February, 
1972. 20 p., 1 fig., bibliogr. Beato: 

78.-The Literature of Arthropods Associated 
with Soybeans. II. A Bibliography of the 
Southern Green Stink Bug, Nezara viridula 
(Linneaus) (Hemiptera: Pentatomidae). 
By N. B. DeWitt and G. L. Godfrey. Macha 
1972. 23 p., 1 fies bibliogr. 


79.-Combined Culture of Channel Catfish analy 
Golden Shiners in Wading Pools. By DA 
Homer Buck, Richard J. Baur, Charles F._ 
Thoits III, and C. Russell Rose. April, 1972. 
12 p., 3 fig., bibliogr. 4 
80.-Illinois Birds: Hirundinidae. By Richard 
R. Graber, Jean W. Graber, and Ethelyn L. 
Kirk. August, 1972. 36 p., 30 fig., bibliogr. — 


CIRCULAR ‘ 


46.—Illinois Trees: Their Diseases. By J. Ced- 
ric Carter. June, 1964. (Third printing, 
with alterations.) 96 p., frontis., 89 fig. 


49-The Dunesland Heritage of Illinois. By 
Herbert H. Ross (in cooperation with Illinois _ 
Department of Conservation). August, 1963. — 
28 p., frontis., 16 fig., bibliogr. 


51—Illinois Trees: 
Care. By J. Cedric Carter. 
123 p., frentis., 108 fig. 

52.-Fertilizing and Watering Trees. By Dan 
Neely and E. B. Himelick. December, 1971. 
(Third printing.) 20 [p., 9 fig., bibliogr. 

53—Dutch Elm Disease in Illinois. By J. Cedric 
Carter. October, 1967. 19 p., frontis., 17 fig. 


Selection, Planting, and 
August, 1966. 


No charge is made for publications of the Ixt1No1s NaTuRAL History Survey. A single copy 
of most publications will be sent free to anyone requesting it until the supply becomes low. Costly 
publications, more than one copy of a publication, and publications in short supply are subjects 
for special correspondence. Such correspondence should-identify the writer and explain the use 


to be made of the publication or publications. 


Address orders and correspondence to the Chief, 
Illinois Natural History Survey 
Natural Resources Building, Urbana, Illinois 61801 


ILLINOIS 


History Survey 
BULLETIN 


_ 7 = 
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f 
i ey 
SES ESS 


The Reproductive Cycle 
of the Raccoon in Illinois 


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ae 


NATURAL KISTORY SURVEY 
NOV 14 1973 
LIBRARY 
DIS 
ENT OF REGISTRATION AND EDUCATION 
HISTORY SURVEY DIVISION 
THE LIBRARY OF THE 


NOV 7- 1973 wwe 


ILLINOIS 


tural History Survey 
BULLETIN 


The Reproductive Cycle 
of the Raccoon in Illinois 


® 


om 


Sanderson 
bandov 


LINOIS 
NT OF REGISTRATION AND EDUCATION 


HISTORY SURVEY DIVISION 
ILLINOIS 


VOLUME 31, ARTICLE 2 
JULY, 1973 


STATE OF ILLINOIS 


DEPARTMENT OF REGISTRATION AND EDUCATION 


BOARD OF NATURAL RESOURCES AND CONSERVATION 


DEAN BARRINGER, Ph.D., Chairman; THOMAS PaRK, Ph.D., Biology; L. L. Stoss, Ph.D., Geology; (VACANT), 
Chemistry; RosERT H. ANDERSON, B.S.C.E., Engineering; CHARLES E. OLMSTED, Ph.D., Forestry; W. L. 
Everitt, E.E., Ph.D., Representing the President of the University of Illinois; RoceR E. BEYLER, Ph.D., 
Representing the President of Southern Illinois University. 


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SCIENTIFIC AND TECHNICAL STAFF 


GEORGE SPRUGEL, JR., Ph.D., Chief 
Avice K. ADAMS, Secretary to the Chief 


Section of Economic Entomology 


umes H. LucKMANN, Ph.D., Entomologist and 

Hea 

Wiis N. Bruce, Ph.D., Entomologist 

WaynE L. Howe, Ph.D., Entomologist 

STEVENSON Moore, III, Ph.D., Entomologist, Exten- 
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Howarp B. Petty, Ph.D., Entomologist, Extension 

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Section of Aquatic Biology 


vey ist W. BENNETT, Ph.D., Aquatic Biologist and 

ea 

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ologist 

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ologist 

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Section of Faunistic Surveys and 
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Section of Wildlife Research 


GLEN < SANDERSON, Ph.D., Wildlife Specialist and 

ea 

FRANK C. BELLROSE, B.S., Wildlife Specialist 

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Harotp C. HANSON, Ph.D., Wildlife Specialist 

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Specialist 

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Specialist 

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Specialist 

Jack A. ELLIS, M.S., Associate Wildlife Specialist 

CHARLES M. NIXxon, M.S., Associate Wildlife 
Specialist 

KENNETH E. SMITH, Ph.D., Associate Chemist 

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Specialist 

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Specialist 

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Supporting Services 


VERNON F. BILLMAN, Maintenance Supervisor 
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Irwin, Chi- 


cago, Illinois; WILDLIFE RESEARCH, WILLARD D. KLIMSTRA, Ph.D., Professor of Zoology and Director of Co- 
operative Wildlife Research, Southern Illinois University; PARASITOLOGY, NORMAN D. LEVINE, Ph.D., Profes- 
sor of Veterinary Parasitology, Veterinary Research, and Zoology and Director of the Center for Human 
Ecology, University of Illinois; ENTOMOLOGY, RoBERT L. METCALF, Ph.D., Professor of Zoology and of En- 
tomology and Head of the Department of Zoology, University of Illinois; and GILBERT P. WALDBAUER, Ph.D., 
Professor of Entomology, University of Illinois; Statistics, HORACE W. NorToN, Ph.D., Professor of Sta- 
tistical Design and Analysis, University of Illinois. 


eS a ee ee a ee 


oe ep 


CONTENTS 


SEARED HDGACEN TS Iaye et hata ie or saci system aesee me cer- crisis iad agepkeces doe Wa eee eed wyeyareo ate ayers od ace 29 
 ETSRHOINS: 5 got egstovnepedt Oe Gee eon Ee HEC u URE ecco reli cn. Ea Ree Be Rear eRe Cee Ree 30 
SHE ARONA Ory GlesOfst Aen GONaGs x avast Hero i tart ses eieyeca oie drole case layer) one oceans 30 
MOLTEN CUR AC COODIS enter siepeiteta) 41.1ok aaNet Terese hayele ay sie se atergles ote duelonnla greparsa¥ene tele ole 30 
WHETIES 5°55 Sac ge och cpgecka CRN eral OS ROE RD CE RCAC RR a ec cca 31 
eric este eeverme erences eg a, pr ee are ne etn cccticbe gece) ces a rs “bare 8 ys. a mierecue Wey dois 31 
Manne bint hme Mater oti lea CCOOM@ I ntters more miete stationed oaees sia fo hip tn.cccohever ere: sasnote el aretene 32 
BeCOM Can yi SEX MR ALIOS fephye eects) oka, jaisilsl sie efoldclss ste hsv aiahal xsi V/éig. elelninia)2 Gis Wisua'e os areiarousi'e 32 
Agmawe (Chyalerinal O.nnenGn Syn noaan san sunenou ond HOemeneds Oooo none ogee ocMbkc 32 
lsgfmoD Ey (ORAKS : 3)4,0s'oid on. Do Se oe orino o OE oe oy Src Ron Cnr aitis AR minis s hey rnc 32 
Or ARL OTe ee Ree Ne osetia cr opars Mette sees Nota oie Sutejontls, wwe acelnvels Means 32 

PAL eUStit ia GM LISCH Emre ee Seeks mercy encarta ere ke. «eis ie) Sue cauey ais Ohentels wise Siena ke 33 
LETSGGIOPRT So 6, Gs of Bit eiplho a One ClO et IRC RET nc iy cic ati oe ey in ear Tara rae 33 
ACERT AS CATS meee seta eee Weir exe it ie caetey 2 cai arcane cancedsex y/o (eur ave" aah svdvereile ojopeea Super os 238) 
Morphologyof the Reproductive: Dracts\< 400.6 see 2 sere scsi mee ines cies eels 33 
INEWIERS sie coy db a Gr BGC 21 CIOS ean CHER IRC RC EROS CRO PR aera eee ar 33 

Fle rrtall Simei riya eres fone easy erctar a oye asco risen anndayn nysrehee wlene) Mourass SNe ewes SAE. wages 34 
Bere TSuO tim Cra SLUCiOl Memmi tranect Neon orcs cts coc ects a suelo terete ia arene dete ke eis. amare 34 
INIeS Pree eter ieee mere to cteicie nd cei dake iho atitnla ae sutsn ada aigk-epla kia s 34 
erarall eS mmenee gerne tegen tee Seen are yea TS AT SU ce SURRY cS eRe Sad See Shoe! Ste Kusiatel 34 
PATeCISFOLMPROP CUOUS! ELOLGIONES eae) nei fayeisrele) ain © Nees cis)Sey=-e eicis evi ithe sm ayes siatetr 34 
INTIS) Lucasol’ GES Road ole crepe 6? ean ceo Rs a eer ea ae 34 
Benita lesamerer merci niprmietncion cteiis ince oo isncladcreler caste Ciesiis ec rersceia chile ha aie aecckionsre 34 
SPelieeernnxresam Vn sega eee wpe seere pe ventae te oo corel cr care sone: aC ar enw ears aterah avanshissa, sats @heracecda,a ve basis 35 
SCS MAND REIS CUS SIO Nia mewsieh cient ace carseat aun SiG esas abel avatecsuel ates tvesduenaye Greve, arab 35 
Seasonal Cvclevorthen Gonads) ja.. aera siete sie cise bi cies area aiecaicleleic s/s were onl 35 
IMTNIER- crcl sib: heentla oo SIONS 4 ao Oza OTT ROR CICIC 1c CCIE IIE ENTE Seamartens 35 
ROTA GB G5 wicre earch bro ad Geexel Gu ea*| EROUE RCECRN Ty EAE Dar RoR SORE CR MC eee Or CTE 43 
arm bs ttle Aten tea Che RS ccteetetarserencta ce vey sree over iaeravinnd 1a (idl ci'e's alrevoveuoucl elveners 45 
Batnarin em birth Dates tOfmina CCOOUS metre lar areiats) aptelniole's clelainie.c © seller aiw, 2iaiedeya einer 46 
SHCORGHIBP IE INAS a5 o.5.00 oon botoh op Ose no) oon Eee ae ee naan Menu nme nr mna neo 47 
Hstrous G@ycle; Ovulation; andPseudopregnancy .... 6.00... 56..-26eeeen ss aree 48 
Histimonis ie Guy. clege cee mera stetave ates iota choc tomv er siaia ewe mie aye plele) uae wiletailel ve onal Mievelehaye 48 
Oirlaiti ore p arnt erie avoir eoiare tee hoe crovareie cuss easy alriiia, pebieia queso eco ens anevoverarene: ei 53 
PSericda pre CMAN Gy! mene ra van ttaehach cous epee seme Sie ee ee «aie Sey d ape S Saeuslg wilajaile eus'ee 55 
Percentage of Yearling Females That Were Sexually Mature .................. 56 


Pigmentation of Mamimae- si: caer: rel: 0) )e ey elelei selec oct ieleseehet stench eee 57 


Interstitial Tissue: 23. geo sive ds hal rie cn ene Geo 57 
Placental ‘Sears 2... 2. eee accs ociuis wep ene « bles se @ ain UanS Tees eee 61 
Morphology of the Reproductive Tracts ..........6.. 20.0 cece eee e cece ee eens 65 
Males: 52 2.5.5 sieve RRs coe dc aS Rea 65 

Bremmales: a. fojece.sosceisseuguocoactevell ve) Won cerenacascacora/seosteh oe Rae coe RPI, oo eel 66 

Effects of Castration (....)00.25..6- eae oe cae ss ote nde cdlone le ierel diene skeen en 68 
Males 5c 0.25 ssrtece fil aacaertaline,sel8 ayo Je;0. 4.000 oe ne eee ae eer 68 

Females, c.ciie isc acn avace Seaseorn cred teena 13 Gh ee 68 

Effects of Exogenous Hormones ........-.---0-- cece ce eee sects ee eene nee 70 
Mallese ci. ainlets sine aides a tetas ee Saree a eae ee 70 

Remales: thee 5 2 alt Acdece carspencnecauert ena S eae ee 71 

Weerine? Milks 355 50 Pe eee ee ee ct 78 
SUONGNGARY. 5 scd.c eter oie sieve. # (doe ae xe cgneunao wpa ashe cos oA RISERS SEAR 79 
PITERATURE: 'GHED. 12 3).04 dese aUs Sele aes pusne the ee ER ee 82 
ts ( 0) >. Career ee Ree ee ee Oe een ren ee tA SON am Bm ihinb.8od ac aca n+ 84 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It 
is a contribution from the Section of Wildlife Research of the Illinois Natural History Survey. 


Glen C. Sanderson is Wildlife Specialist and Head, Section of Widlife Research, Illinois 
Natural History Survey. A. V. Nalbandov is Professor of Animal Science, Physiology, and 
Zoology, University of Illinots. 


(50105—5M—7—73) 


S14 


The Reproductive Cycle of the 


Raccoon in Illinois 


ALTHOUGH THE RACCOON 
(Procyon lotor) is a commonly recog- 
nized, widely distributed, and abundant 
North American mammal, little has been 
known about its reproductive cycle except 
the season of birth, the number of young 
per litter, and the duration of the gesta- 
tion period. Basic information on the 
length of the estrous cycle, whether ovu- 
lation is spontaneous or induced, the 
period of sexual activity in the male, the 
occurrence of pseudopregnancy, the roles 
of the various hormones in reproduction, 
and the anatomy of the reproductive 
tracts has been either lacking or frag- 
mentary. 

The objectives of this study were to 
gather data on the reproductive cycle 
and the basic anatomy of the reproduc- 
tive system of the raccoon and to in- 
vestigate those aspects of the raccoon’s 
reproductive physiology that gave prom- 
ise of increasing our knowledge in the 
general field of mammalian reproductive 
physiology. This study was part of an 
effort to obtain a refined understanding 
of the population dynamics of the species. 
Other aspects of the study will be pub- 
lished elsewhere. 


ACKNOWLEDGMENTS 


Prior to 1961 this work was supported 
by Illinois Federal Aid Project W-56-R, 
the Illinois Department of Conservation, 
the U.S. Bureau of Sport Fisheries and 
Wildlife, and the Illinois Natural History 
Survey, cooperating. During 1961, 1962, 
and 1963 partial support for these studies 
was contributed by the National In- 
stitutes of Health under Research Grant 


Frontispiece.—Cages used to hold raccoons in Urbana, I'l. 


Glen C. Sanderson 
A. V. Nalbandov 


7849. The remainder of the support for 
this study was provided by the Illinois 
Natural History Survey. 

We thank Dr. T. G. Scott, former 
Head of the Section of Wildlife Research 
at the Survey, for his encouragement and 
advice throughout the study, and Dr. H. 
W. Norton, Professor of St-istical De- 
sign and Analysis in the Department of 
Animal Science at the University of Illi- 
nois College of Agriculture, for his help 
with the statistical analyses. We also 
thank Dr. Jean W. Graber, former Re- 
search Assistant Professor of Animal 
Science, University of Illinois, who pre- 
pared most of the histological sections 
and provided other valuable assistance, 
and the senior author’s wife, Beverley C. 
Sanderson, who drew the sketches of the 
male and female reproductive tracts and 
helped in many other ways. W. D. Zehr, 
Illinois Natural History Survey Tech- 
nical Photographer, and G. G. Mont- 
gomery, former Survey staff member, 
assisted with the photomicrographs. R. 
J. Ellis was employed as Research As- 
sociate on Project W-56-R from Feb- 
ruary 1, 1961 through June 30, 1962 and 
contributed specimens and other assist- 
ance to this study. G. G. Montgomery 
was employed as Research Associate on 
Research Grant 7849 and made many 
contributions to the study. C. L. Foley, 
Illinois Department of Conservation, 
Paris, Ill., supplied live raccoons for this 
study and was helpful to the project in 
other respects. Present and past em- 
ployees of the Natural History Survey 
who contributed specimens and informa- 
tion to the study include Dr. B. J. Verts, 
Dr. G. L. Storm, Dr. R. D. Andrews, Dr. 


Each double cage held either one 


pair, one female and her young, or 1-3 adult raccoons in each half. The outside dimensions of the 


cages were 3 feet (width) X 4 feet (height) X 6 feet (length). 
through the middle with wire and had a nest box on each end. 
The nest boxes had wire bottoms, 
A hinged wire top on each nest box fitted under the removable lid. 


14-gauge hexagonal netting. 
serted on top of the wire in winter. 


Each cage was divided crosswise 
The wire was 1.5-inch mesh, 
and wooden bottoms were in- 


30 Inurino1is NaruraL History SuRvVEY BULLETIN 


R. R. Graber, and others. Helen C. 
Schultz of the Survey staff and Robert 
M. Zewadski, Associate Technical Editor 
of the Survey, edited the manuscript. 

Dr. H. W. Norton and Dr. A. Sydney 
Johnson, Associate Director, Institute of 
Natural Resources at the University of 
Georgia, Athens, reviewed the man- 
uscript and made many valuable sugges- 
tions. 

We are especially grateful to Clifford, 
Albert, and Robert Perardi (Perardi 
Brothers Fur and Wool Company, Farm- 
ington, Ill.) for their active and en- 
thusiastic cooperation with our study. 


METHODS 


SEASONAL CYCLE OF THE GONADS 


Each year from 1955 through 1961 the 
senior author examined dead raccoons at 
a number of fur houses in central Illinois. 
The majority of the raccoons were exam- 
ined at Farmington in Fulton County and 
Colchester in McDonough County. Most 
or all of these animals came from within 
the range of Procyon lotor hirtus (Gold- 
man 1950:24). During the hunting and 
trapping season, which usually occurred 
during November through January (but 
occasionally included late October), 
large numbers of recently-killed raccoons 
were sold to fur-buying establishments 
and pelted. Often a majority of the ac- 
ceptable carcasses were dressed and 
frozen prior to being sold for human 
food. Thus, from the large number of 
raccoons examined, numerous data were 
recorded and many organs suitable for 
gross examination were collected as the 
animals were being skinned. 

The present report deals principally 
with the reproductive organs of the rac- 
coon. Before the animals were skinned, 
one testis and epididymis were removed 
from each male, and the condition of the 
nipples of each female was recorded. All 
pertinent information was recorded sep- 
arately for each animal. After the rac- 
coons were pelted, the complete repro- 
ductive tracts were removed from fe- 


Vol. 31, Art. 2 


males and were placed separately in 1- 
pint plastic bags to prevent the tissues 
from drying. Each plastic bag was 
placed in a small paper bag on which 
the data were recorded. 

The specimens were usually examined 
in the laboratory the day after collection 
but sometimes were examined on the day 
they were collected. The testes were 
weighed to the nearest 0.1 gram. A drop 
of fluid collected from the tail of th 
epididymis was diluted with a drop of 
normal saline solution and examined un- 
der the microscope for the presence of 
sperm. Both ovaries were examined vi 
sually and weighed to the nearest 0.1 mg. 

Raccoons found dead or collected by 
trapping and shooting specifically for au- 
topsy were processed in the same general 
manner as those examined in fur houses. 
A small number of raccoons, obtained 
from sources other than fur buyers, came 
from the southern and eastern sections of 
Illinois within the range of P. 1. loto 
(Goldman 1950:24). 

Gonads from both sexes were collecte 
from adult and juvenile raccoons eac 
month. Several gonads were removed 
immediately after the deaths of the 
animals and were preserved and pre- 
pared for histological study. The aver- 
age monthly weights of the gonads fro 
all of the raccoons studied, both those 
freshly killed and those dead for severa 
hours, were used in constructing graphs 
showing the seasonal gonadal weights fo 
juveniles and adults of both sexes. His- 
tological examinations of the testes, ep- 
ididymides, ovaries, and uteri contrib- 
uted information regarding the seasonal 
sexual cycle. 


CAPTIVE RACCOONS 


For many phases of the study captive — 
raccoons were kept in outdoor cages in 
Urbana, III. Most of these animals were 
trapped in the wild, both as adults and 
juveniles, and some as small young, most- | 
ly in Champaign, Piatt, Edgar, and Car- 
roll counties, Ill. We estimated the ages 
of wild raccoons at the times of their cap-_ 


4 
| 
i 


July, 1973 Sanperson & NALBANDov: REPRODUCTIVE CYCLE OF THE Raccoon 31 


ture (Sanderson 1961a). Some animals 
used for the study were born in captivity 
—some were conceived in captivity and 
others were born in captivity to females 
that were pregnant when captured. 

Captive raccoons were usually paired 
and held as one male and one female per 
cage. Pregnant females were isolated 
prior to parturition; the males were not 
returned while the young were with the 
females. Some females were isolated to 
determine whether ovulation in the rac- 
coon is induced or spontaneous. In some 
cases three or more animals—juveniles of 
both sexes and surplus males—were 
held in a single cage. 

The captives were given fresh food 
and water daily. The main diet was 
Dog Checkers or Laboratory Checkers, 
manufactured by the Ralston Purina 
Company. Occasionally the diet was 
supplemented by chickens, fish, eggs, and 
other available fresh foods. 

Captive raccoons that died or were 
killed were processed as described above, 
except that all of the gonads, after being 
weighed, were preserved for histological 
study. Usually a section of the uterus 
and occasionally accessory organs of the 
reproductive tract were also preserved 
for histological examination. 


Males 


The annual reproductive cycle in 
several captive male raccoons was deter- 
mined by restraining each male in a 
wire cone at irregular intervals through- 
out the year and collecting a drop of 
fluid from the tail of the epididymis. 
The tail of the epididymis was forced 
against the skin of the scrotum; then a 
pointed scalpel was used to prick through 
the skin, and a drop of fluid was collected 
on a glass slide. The drop was diluted 
with normal saline solution and _ ex- 
amined under the microscope for the 
presence of sperm. After the collection 
of the epididymal fluid, the animal was 
returned to its cage with no further treat- 
ment. No infection or other troubles 
resulted from this treatment. 

Occasionally, a captive male, or a wild 


male that had been livetrapped and was 
to be released at the point of capture for 
another phase of the study, was uni- 
laterally castrated to obtain a testis and 
epididymis for study. Captive males 
that fathered young were assumed to 
have had sperm in their epididymides at 
the time that they impregnated the 
females. 


Females 

The reproductive cycle of captive fe- 
male raccoons was studied by examining 
the ovaries and uteri during laparotomies 
of anesthetized animals. The anesthetic 
used was pentobarbital sodium admin- 
istered at the rate of 1 cc per 4 pounds of 
body weight. Given intraperitoneally, it 
usually produced surgical anesthesia in 
10-30 minutes; however, individual re- 
sponses to the anesthetic varied, and 
animals that required more anesthetic 
were given larger doses the second time 
laparotomies were performed. 

The raccoon is resistant to infection 
and withstands surgical incursions well. 
Instruments were washed in 70-percent 
alcohol but were not sterilized. As many 
as 12 laparotomies were performed on 
one female over a period of several 
months, sometimes on subsequent days, 
sometimes two or three times in 1 week, 
but usually from 2 weeks to several 
months apart. Animals were usually 
given penicillin after each operation al- 
though no infections developed when it 
was not used. Surgical silk or cat gut 
was used to close the peritoneal linings 
and muscle; these sutures were not re- 
moved until a subsequent laparotomy 
was performed. Wound clips, used to 
close the skin, were removed approx- 
imately 10 days after the operation. 

To examine ovaries for evidence of 
ovulation, it was usually necessary to slit 
the ovarian capsules. Because the cut 
edges of the capsules did not always grow 
together, this procedure was omitted 
when examining females that were being 
held to produce young. The uterus was 
gently withdrawn from the body cavity 
for examination and gross measurement. 


32 Ittinors NaturaL History SuRVEY BULLETIN 


In several cases one or both ovaries 
were removed for study. Uterine sec- 
tions were taken from living females for 
histological study of the development of 
the endometrium. 


MEAN BIRTH DATE OF 
RACCOON LITTERS 

The mean date of birth was deter- 
mined for 20 litters conceived in the 
wild in the northern half of Illinois. Of 
these 20 litters, 7 were born in captivity. 
The potential birth dates of the others, 
most of which were examined in female 
raccoons found dead along roadways, 
were estimated by measuring the uterine 
swellings in the manner described by 
Llewellyn (1953:321). Data obtained 
during the present investigation were 
also used in estimating the probable birth 
dates. Because Llewellyn (1953:321) 
recorded measurements of only three 
embryos in one litter at three different 
stages and at birth, several embryos were 
measured in captive females during this 
study. Although the dates of conception 
were not known, the maximum measure- 
ments of the uterine swellings were plot- 
ted in relation to the number of days 
prior to the known birth dates. Many 
wild females were examined throughout 
the year for pregnancy, lactation, and the 
presence of fresh placental scars and 
corpora lutea. This information helped 
to determine the limits of the breeding 
season in wild raccoons. 


SECONDARY SEX RATIOS 


Secondary sex ratios were obtained 
by examining 83 embryos and young at 
birth in 26 litters and by determining 
the sex of 54 wild raccoons less than 2 
months old from 23 litters. Chi-square 
tests were used to test whether the sex 
ratio of the wild young less than 2 
months of age was different from equal- 
ity and from the ratio of the embryos 
and young at birth. 


ESTROUS CYCLE AND OVULATION 
Estrous Cycle 


Estrous cycles were determined for 
individual captive female raccoons by 


Vol. 31, Art. 2 


examining the ovaries at or near ovula- 
tion and then reexamining the ovaries at 
intervals until the animals ovulated 
again. 

The raccoon’s main breeding season 
was interrupted throughout much of IIli- 
nois by colder -than-normal temper- 
atures and deep snows in 1960. Obser- 
vations of livetrapped raccoons and the 
body weights of young, wild raccoons 
weighed during the fall and winter of 
1960 indicated that some raccoons were 
born later than normal during that year. 


Lenses collected from several young rac- . 


coons during the hunting and trapping 
season of 1960-1961 were used to esti- 
mate the months of birth for these 
juveniles (Sanderson 1961b:482—-485). 
The time intervals between the peaks of 
estimated birth dates were assumed to 
represent the average interval between 
ovulations for wild raccoons in central 
Illinois. 

Cotton swabs were used to take daily 
vaginal smears from several captives in 
an attempt to delineate the estrous cycle. 
Observations of vulval swelling, size and 
pigmentation of the nipples, and general 
disposition of the animals were made 
each time the animals were handled. 
Vaginal tissues were removed from sev- 
eral females for histological study. 


Ovulation 


Each of two females was placed alone 
in a small cage in the fall of 1960 to 
obtain information on the mechanism of 
ovulation and on _ pseudopregnancy. 
These females could see other raccoons 
but could not come into physical contact 
with them. Also, one pet female, re- 
ported by the owner to have had no con- 
tact with other raccoons, was observed. 
Individual corpora lutea were studied in 
these females during a series of laparot- 
omies. 

Some of the corpora lutea in the ova- 


ries of three females were marked with ~ 


India ink—and the locations of all 
corpora lutea were mapped. By follow- 
ing the fate of the marked and mapped 
corpora until they disappeared, we found 
that mapping the corpora lutea was as 


July, 1973 Sanperson & NaLBanpov: REPRODUCTIVE CycLE OF THE Raccoon 33 


reliable a method of determining their 
life-spans as was marking them with ink. 
Mapping was used in subsequent studies. 
At each initial observation the ovary was 
forced through the slit ovarian capsule, 
the corpora were examined for color and 
measured grossly, and their locations in 
the ovary were mapped. 


INTERSTITIAL TISSUE 


Ovarian interstitial tissue was studied 
in wild raccoons on which observations 
as to pregnancy and lactation had been 
made, in several captive females treated 
with various hormones prior to the re- 
moval of the ovaries, and in untreated 
captives whose breeding histories were 
known. A uterine section was usually 
obtained when ovaries were collected, 
and the condition of the endometrium 
was studied in relation to the degree of 
development of the interstitial tissue. 
Representative sections selected from 
each ovary and uterus were photo- 
graphed by mounting the slide in the 
carrier of a photographic enlarger and 
projecting the image directly onto 4- X 
5-inch contrast process ortho sheet film. 
Prints 8 X 10 inches were made on F5 
Kodabromide paper. By examining the 
photographs, we determined the abun- 
dance and distribution of cells of each 
type in the interstitial tissue in relation to 
the development of the endometrial 
glands, the time of year, the age of the 
animal, and the stage of the reproductive 
cycle. 


HISTOLOGY 

Tissues were preserved in Bouin’s 
solution or in 10-percent formalin 
neutralized with either MgCO, or 


CaCO,. The organs preserved in Bou- 
In’s solution were left for an indefinite 
period, but those preserved in 10-percent 
formalin were transferred to 70-percent 
alcohol after 48-72 hours. With a few 
exceptions, all tissues prepared for his- 
tologcial examination were stained with 
hematoxylin and eosin. The ovaries of 
a few females that had died some time 
prior to the preservation of the organs 


were sectioned at 15-20 microns; the 
number of corpora lutea was our main 
interest in these ovaries. In all other 
cases the sections were cut 6 microns 
thick. The preserved organs were em- 
bedded in paraffin and sectioned and 
mounted by routine methods. 


PLACENTAL SCARS 


In dead female raccoons placental 
scars were counted, using transillumina- 
tion. The uterus was then slit and the 
inside surfaces were examined for scars. 

In captive pregnant females the uter- 
ine swellings were measured and the 
locations of the embryos were mapped 
during laparotomies. After parturition 
the presence and persistence of placental 
scars at the sites of known placental 
attachment were studied during a series 
of laparotomies. The scars were ex- 
amined in living animals by gently pull- 
ing the uterus far enough out of the body 
cavity to allow it to be transilluminated. 
Uterine sections containing scars at 
various stages were removed from living 
females at intervals for histological study. 


MORPHOLOGY OF THE 
REPRODUCTIVE TRACTS 


Males 


A few complete male reproductive 
tracts were removed and preserved for 
histological study. The entire tract from 
one male, and individual accessory 
organs from a few additional males, were 
sectioned. India ink was injected into 
one vas deferens of a fresh specimen until 
the ink ran out the urethral opening of 
the penis. The tract was then preserved 
and sectioned for histological study to 
trace the duct system, containing parti- 
cles of India ink, through the prostate 
gland. 

A schematic diagram of the male re- 
productive system was sketched from a 
fresh specimen that had been partially 
dissected but was sufficiently undisturbed 
to show its relationships to adjacent 
structures. A complete reproductive 
tract that had been dissected and pre- 
served was used for reference. 


34 Intinors NATURAL History SURVEY BULLETIN 


Females 


A schematic diagram of the reproduc- 
tive tract (frcm one female) was prepar- 
ed from a fresh tract that had been sufh- 
ciently dissected to reveal its conforma- 
tion but that maintained its position 
relative to adjacent structures. One en- 
tire tract that had been removed and 
preserved was used for reference. 


EFFECTS OF CASTRATION 
Males 


Four captive male raccoons were 
castrated at ages ranging from 72 days 
to approximately 9 months to study the 
effects of castration on the development 
of the penis bone, the opening of the 
preputial orifice, and the age at which 
the epiphyses close in the radius and 
ulna. These studies were not completed 
because the four animals died of various 
causes at different ages; the one that 
lived the longest attained an age of 
approximately 22 months. 


Females 


One female raccoon, born in captivity, 
was 3 months of age when castrated; the 
second, born in the wild, was estimated to 
be 4 months old when castrated. Several 
adult females were also castrated to study 
the effects of castration on vaginal 
smears, the vaginal epithelium, the 
uterus, and the closure of the epiphyses 
in the radius and ulna. 

Vaginal tissues and uterine sections 
were taken from castrated females at 
intervals. These tissues were prepared 
for histological study and used for com- 
parison with similar tissues from females 
believed to be anestrus. The females 
castrated as adults were also used to 
study the effects of various exogenous 
hormones on vaginal smears, the vaginal 
epithelium, and the development of the 
endometrium. 

Two pregnant females were castrated 
as the first phase of a study of the effect 
of castration on pregnancy. The first fe- 
male, with four embryos, was castrated 
38 days (estimated time) after concep- 
tion. The second female, with five em- 


Vol. 31, Art. 2 


bryos, was castrated approximately 11 
days after conception. These females 
were observed daily after castration for 
signs of abortion. A second laparotomy 
was performed on the first female 21 days 
after castration and on the second female 
19 days after removal of the ovaries. 


EFFECTS OF EXOGENOUS 
HORMONES 


Males 


Two captive adult male raccoons were 
used for preliminary studies of the effects 
of androgen on spermatogenesis. Begin- 
ning in August, near the midpoint of 
sexual inactivity, injections of  testos- 
terone cyclopentylpropionate (Res. No. 
8961-1, Upjohn) were administered to 
both of these males. The first male re- 
ceived seven subcutaneous injections of 
30 mg each at 3-day intervals. 

Immediately before the first injection 
of the hormone the left testis and ep- 
ididymis were removed from each an- 
imal. The testis was weighed and a 
smear from the tail of the epididymis 
was examined for the presence of sperm. 
Each testis and epididymis was prepared 
for histological study. 

The first male was killed 21 days after 
receiving the first androgen injection, 
and the right testis, right epididymis, and 
the prostate were removed. The testis 
was weighed and a smear from the tail 
of the epididymis was examined for 
sperm. The second male was similarly 
treated but received four injections of 
12 mg each and was killed 15 days after 
the first injection was administered. 


Females _ 


Several attempts were made to cause 
the growth and development of Graafian 
follicles: and to cause ovulation by in- 
jecting various hormones into female 
raccoons. The hormones used were 
pregnant mare’s serum (PMS, Upjohn), 
the pituitary gonadotropins (FSH and 
LH, Armour), estradiol cyclopentylpro- 
pionate (ECP, Upjohn), estradiol valer- 
ate (estradiol, Squibb), hydroxyproges- 
terone caproate (progesterone, Squibb), 
chorionic gonadotropin (CGH, Up- 


July, 1973 Sanperson & NatBanpov: RepropuctTivE CycLe oF THE Raccoon 35 


john), and human menopausal gonado- 
tropin (HMG-J5, Statens Seruminstitut, 
Copenhagen). Because these hormones 
were administered by many different 
routes and at many different dosage 
levels and time intervals, the methods 
used are discussed in connection with 
the particular animals involved or are 
given in the tables where the results 
from the individual animals are sum- 
marized. 


UTERINE MILK 


Studies were made to determine the 
hormone or hormones responsible for the 
secretion of uterine milk by the endo- 
metrial glands and to learn the nature of 
this secretory material. Ovaries and 
uteri were sectioned and stained from 18 
raccoons—all were collected during the 
breeding season and some of them were 
pregnant—in which corpora lutea were 
present and from 89 raccoons—collected 
throughout the year—whose ovaries con- 
tained no corpora lutea. None of these 
107 raccoons had been injected with 
hormones. In all cases the endometrial 
glands were examined for the presence of 
secretory materials. 

Various hormones were administered 
to castrate females, uterine sections were 
removed at varying time intervals, and 
the endometrial glands were examined 
by histological methods for the presence 
of secretory material. The hormones 
used on individual castrate and intact 
females to study hormonal control of the 
secretion of uterine milk were progester- 
one and ECP, ECP alone, and progester- 
one alone; however, progesterone alone 
was not given to any castrate animal for 
a sufficient time to determine whether 
it would cause the uterine glands to se- 
crete. Also studied were the direct and 
secondary effects of PMS, FSH, and LH, 
used primarily in attempts to cause the 
growth of Graafian follicles and to cause 
ovulation, and the production of secre- 
tory material by endometrial glands in 
intact females. 

Methods described by Pearse (1960: 
265-271) and Lillie (1954: 274-299) 


were used to demonstrate the nature of 
the material observed in the lumina of 
the endometrial glands. Uterine sections 
from three female raccoons that had 
material present in the endometrial 
glands were used. The uterine section 
from one was fixed in 10-percent for- 
malin neutralized with CaCO,. The 
uterine section from another was fixed in 
Bouin’s solution, and the section from a 
third was fixed in 10-percent formalin 
neutralized with MgCO,. All of these 
tissues were imbedded in paraffin for 
sectioning, and control slides were used 
in each case. 


RESULTS AND DISCUSSION 


SEASONAL CYCLE OF THE GONADS 
Males 


The age at which male raccoons reach 
sexual maturity may vary from one re- 
gion to another. In Michigan, on the 
basis of meager circumstantial evidence, 
Stuewer (1943b: 72) concluded thai 
males ‘“‘are probably not sexually mature 
by the first breeding season after their 
birth.” In Illinois Pope (1944: 91) had 
two captive males—of parent stock sup- 
posedly “from northern Illinois or some 
adjacent region’’—that , mated success- 
fully before they were 1 year of age. 

Stuewer (1943b:63) reported that the 
testes of juveniles and yearlings were in 
an abdominal position; those of adults 
were usually descended during the breed- 
ing season and, though variable in posi- 
tion, during the remainder of the year 
were most often in the coelom. Stuewer’s 
evidence suggested that testis size might 
reflect the capacity to breed. He measur- 
ed the lengths of testes in the scrotum 
with an accuracy of approximately 5 mm. 
Stuewer (1943b: 64) concluded that if 
“testis size is significant, males are prob- 
ably capable of breeding at all times of 
year after reaching maturity.” Asdell 
(1946: 136), on the basis of Stuewer’s 
work but omitting his qualifications, stat- 
ed that the male raccoon was capable 
of mating at any time. Nalbandov 
(1958: 162) cited the male raccoon as 


36 


a species in which spermatogenesis is 
continuous although the breeding season 
of females is restricted to late winter and 
early, spring. 

The data in Table 1 and Fig. 1 show 
that raccoon testes grew at a rather uni- 
form -rate from birth until about 10 
months of ‘age (through the February 
after birth), when the average weight of 
one testis was 5.6 grams. The testes of 
juvenile males showed the most rapid 
gains in weight between December and 
February. The average weight of a 
testis from a juvenile male in November 
was only 30 percent of the average 
weight in February. The sample sizes 
for February, March, and April were 
small, but there was an indication that 
the weights of testes in juveniles declined 
after February. After April testicular 
weights of juveniles were included with 
those of adults because a majority of 
the juvenile males were sexually active 
by April. 

In our experience raccoon testes were 
nearly always found in the scrotum, 
even at birth, Stuewer’s (1943b: 63) 
statements to the contrary notwithstand- 
ing. They were more prominent in 
adults than in juveniles, and most prom- 
inent in adults during the breeding sea- 
son. Even in immature animals the 
testes were rarely withdrawn into the 
body cavity. 


WEIGHT IN GRAMS 


BS Poaweinteg 


MAY JUNE JULY AUG. SEPT. 


Intino1is NaTurRAL History SurRvEY BULLETIN 


2 ADULTS 


ocT. 


Vol. 31, Art. 2 


In Illinois a majority of the male rac- 
coons reached sexual maturity as year- 
lings. Although the presence of sperm 
in the epididymis does not necessarily 
indicate sexual potency, it does indicate 
that an animal is in or approaching the 
period of sexual activity. No juvenile 
male had sperm in its epididymis prior 
to October (Table 1). In October the 
epididymides of about 9 percent of the 
juveniles contained sperm; by February 
this figure had increased to 87 percent. 
An extrusible penis was another indica- 


tion of a juvenile’s stage of sexual de-: 


velopment (Sanderson 1961a: 14). Oc- 
casionally, a male was found with a non- 
extrusible penis but with sperm in its 
epididymides. Among juvenile males, 5 
percent had extrusible penes in Septem- 
ber. This figure had increased to about 
67 percent by February and March but 
declined slightly in April. These data in- 
dicated that, in Illinois, from one-half 
to two-thirds of the juvenile male rac- 
coons are sexually mature by the time 
they are 1 year old (Table 1). By sex- 
ually mature we mean that the male 
has an extrusible penis and a relatively 
high concentration of sperm in the epi- 
didymides. 

Comparison of data from juvenile and 
adult male raccoons shows that juveniles 
became sexually mature 3-4 months later 
in the year than did adults. Several 


HX X. 
CxKXS 
SOR 


rd 
S505 


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CORK 

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2 


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ves 
<5 

5 
veraterens 


396/ 
ee, 


Se 
SOO 
QO 
QOL? 
QOOQ? 
SSC 


NOV. DEG. 


Fig. 1.—Seasonal variations in the average weight of one testis in adult and juvenile raccoons. 
With each mean are the number of observations and a vertical line representing the mean plus or 
minus one standard error. All animals were taken in Illinois from November 1955 through April 1961. 


The data are given in Table 1. 


a ee ee a See ae ee eee 


REPRODUCTIVE CYCLE OF THE RACCOON 37 


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July, 1973 Sanperson & NALBANDOV 


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38 Intinois NaturaL History SuRvEY BULLETIN 


juvenile males had no sperm in their 
epididymides during the peak of the 
breeding season but became sexually 
mature after most of the breeding had 
been accomplished. At least some adult 
males were incapable of breeding when 
the second and third ovulations occurred. 
Hence, we believe that a majority of 
the second litters born to raccoons are 
sired by yearling males. 

This study is the first to establish that 
seasonal variations occur in the testis 
weights of raccoons (Fig. 1). The aver- 
age weights of the testes of adult males 
were minimal in June, July, and August, 
began increasing during September, and 
reached their peak in December. Among 
adults the maximum average weight of 
one testis was nearly three times the 
average minimum weight. A decline in 
testis weight appeared to occur prior to 
the peak of the breeding season in Feb- 


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JAN. FEB. 


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. MAY JUNE JULY AUG. 


Vol. 31, Art. 2 


ruary; however, our sample sizes for Feb- 
ruary, March, and April were small. 
Testes continued to decline from their 
peak weight in December to a low point 
in July. In adults there was a positive 
correlation between testis weight and the 
presence of sperm in the epididymis (Fig. 
Ne 

Four males were unilaterally castrated 
on different dates. The second testis 
was removed from each at a later date. 
The weights of these testes are shown 
in Table 2. In many species the removal 


of one gonad causes the second one to : 


hypertrophy, but our observations on the 
effects of unilateral castration in the rac- 
coon on the weight of the remaining 
testis are inconclusive. The remaining 
testes in the two adult males castrated 
unilaterally in July showed greater-than- 
average increases in weight from July 
to December. A _ greater-than-average 


SR RRRRNG 


CULO GA 
PES 


1 
LO 


“5 


SK 


ae 
SS 


DEC. 
Bess]: PERCENTAGE WITH SPERM 


Fig. 2.—Seasonal variations in the average weight of one testis taken from adult raccoons and 
the percentage of adults with sperm in the epididymis. The numeral at the top of each bar in- 


dicates the number of observations. 


REPRODUCTIVE CYCLE OF THE Raccoon 39 


July, 1973 SanpEerson & NaLBANDoy: 


Table 2.—Testis weights of four raccoons, showing seasonal changes in individual animals. 


Estimated 
Age 
in Months 


2 
BRO 
ew ON 
ono 
AL< 
na 

fo) 
nan 
roush 
” & 
ae 8 
ENS) 
Po 
Sg 
=oO-5 
oO 
2 
3 
(= 


Raccoon 
Number 


and Type 


ae War lar Wl 


NeooNnorn 
AONDMOMA 


~nrennrwowoo 
Sipe n e 
AamnwuomMmnn 
TTA we 
ee et anes 
Aol “ere 
moNnoOnatn 
mana Seales! 


1783 (captive) 
1803 (captive) 
2121 (captive) 


1771 (wild) 


® The plus sign indicates that sperm was present, and the minus sign that it was absent. 


were €xX- 


raccoons 


captive 


Several 
amined repeatedly for sperm in their 


increase in weight occurred from Novem- 
ber to January (the period of maximum 


epididymides after they became sexually 


mature (Fig. 3). 


growth rate in juvenile males) in one 


From July through 


October the greatest percentage of adult 


After unilateral 


juvenile male after the removal of one 
in November. 


testis 


inactive. The 


histories of several captives show that 


sexually 
individual males had periods that aver- 


were 


males 


castration in May the second testis in 


OD. @.@.9.2.@.@: 


aged 3-4 months when they were incap- 
able of breeding although males with 


IIIT ST I OT OT Oe 


Yee 


“Gn Gb 4b > Oe ea eee aa: 


).O.2.@.@.@.@.2@.@ 2.0.3.3 .2@.38.2.@.8.2.@.2.O.2.2.2@.e.2o.e: 


JAN. FEB. MAR. APR. MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC. 


the remaining adult showed a decrease in 
weight from May to September, a period 
when the average weights of testes from 
wild adults did not differ significantly. 


JAILDV ATIVNXAS JOVWLNIIDY3d 


The 


Fig. 3.—Percentage of sexually mature captive male raccoons with sperm in the epididymis. 


numeral at the top of each bar indicates the number of observations for each month. 


40 Ituino1is NaturaL History Survey BULLETIN 


sperm in their epididymides were found 
in all months (Fig. 3 and 4). Lower 
concentrations of sperm were found at 
the beginning and end of the period of 
sexual activity than were found during 


[IMM SPERM PRESENT 


3347, MI2 


1783, MI2 


3337, MI2 
mt 


SS SN 11 1111110110001 Wt 
TT 
HN! a a Ee 
aaa 


RACCOON NUMBER AND AGE AT FIRST OBSERVATION 


2123, Mi2 
2154B, A7 
2298, AT 
2299, A7 
2752, Al7 
2825, AlO 
2968, Al2 
3087, AB 


JAN. 


MEEENNB NO._- SPERM PRESENT 


Vol. 31, Art. 2 


the peak, but how the concentration 
of sperm is related to a male’s fertilizing 
ability is not known. 

Histological examinations of the testes 
and epididymides of 85 wild and 37 cap- 


[-__] NO DATA 


FEB. MAR. APR. MAY JUNE. JULY AUG. SEPT. OCT. NOV. DEC. 


Fig. 4.—Presence or absence of sperm in the epididymides of captive male raccoons. These 
raccoons were held in outdoor cages throughout the year in Urbana, III. The animal's identification 
number is given first, and the estimated age in months when the first observation was recorded is 
given next. A = approximate age, M=minimum age. The different lines for each animal indicate 


different years. 


July, 1973 Sanperson & NaLBANDov: REPRODUCTIVE CYCLE OF THE Raccoon 41 


tive male raccoons (Table 3) confirmed 
the gross observations, reported above, 
made on captives. In all but five males 
if sperm were present in the testes, they 
were also present in the epididymides, 
and vice versa. Data from these five 
cases indicate that sperm may be stored 
in the epididymis for some time after 
spermatogenesis ceases and that sperm 
may be found in the testis prior to being 
stored in the epididymis. Males of some 
species are able to ejaculate fertile sperm 
for as long as 4 weeks after castration 
(Nalbandov 1958: 176). Testes and 
epididymides removed from two adult 
raccoons in August (Table 3) are repre- 
sentative of the conditions found. One 
male had sperm in the seminiferous 
tubules but none in the epididymides. 
The second male had low concentrations 
of sperm in both the testes and the epi- 
didymides. 

These data show that the male rac- 
coon has a seasonal sexual cycle. The 
general correlation between the size of 
the testis and the presence of sperm in 
the epididymis did not hold in individual 
cases. Three hundred eighty-four testes 
with sperm in the corresponding epi- 
didymides, taken from adults from Oc- 
tober through April, averaged 7.2 grams 
and ranged from 2.6 to 11.3 grams. 


Fifteen testes with no sperm in the cor- 
responding epididymides, taken from 
adults during the same months, averaged 
4.6 grams and ranged from 1.2 to 9.5 
grams, 

A substantial number of testes were 
weighed during November, December, 
and January from the 1950-1951 fur sea- 
son through the 1960-1961 fur season 
in Iowa and Illinois. Raccoons in Iowa 
were examined at a fur house in Bloom- 
field, Davis County. Most specimens 
from Illinois were collected at a fur 
house in Farmington, Fulton County. 
Farmington is approximately 130 miles, 
almost due east, from Bloomfield, and 
the reproductive cycles of the animals 
collected at these two locations probably 
were similar. A few Illinois specimens 
were collected from other fur houses 
located in the central (north-south) 
third of Illinois. Each fur buyer bought 
dead raccoons from hunters and trap- 
pers living within a radius of about 100 
miles around his location. 

These data were collected to study 
seasonal and annual trends in the weights 
of testes (Table 4) and the timing of 
spermatogenesis in relation to age among 
male raccoons. The testes of juveniles 
gained weight significantly (P<0.02) 
from November to January. No signifi- 


Table 3.—Occurrence of sperm in the testes and epididymides of adu!t and juvenile raccoons as 
determined by histological examination of 85 wild and 37 captive males.” 


Adults” Juveniles” 
Month Percent Number of Percent Number of 
with Sperm Observations with Sperm Observations 

January 67 a Aero 0 
February 100 3 0 1 
March 100 1 25 4 
April 75 4° 57 7 
May 25 4 0 2 
June 0 1 0 6 
July 17 12 0 8 
August 25 4° 0 10 
September 33 9 0 as 
October 80 10 0 gt 
November 86 Ue 14 7 
December 100 1 0 32 


8 All observations were made in Illinois from 1957 through 1960. 

> An adult was more than 12 months of age; a juvenile was 0-12 months of age. 

© One adult male in April and one in August had sperm in the epididymides but not in the testes. 

4 One adult male in November and one juvenile male in October and one in December had sperm in the 


testes but not in the epididymides. 


Vol. 31, Art. 2 


Intrnois NaTurRAL History SuRVEY BULLETIN 


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July, 1973 Sanperson & NatBanpov: REPRODUCTIVE CycLE OF THE Raccoon 43 


cant differences in the average weights of 
adult testes occurred from November to 
January. This finding was not unex- 
pected, because virtually all adult males 
were capable of breeding by November 
but only 8 percent of the juveniles had 
sperm in the epididymides during No- 
vember (Table 1). There were some 
statistically significant annual differences 
in the weights of testes, but the meanings 
of these differences were not clear. 


Females 


The ovaries of raccoons showed a 
nearly steady rate of growth from birth 
in April through the following November 
(Table 5 and Fig. 5). In contrast, the 
testes of juveniles showed their most 
rapid increases in weight between De- 
cember and February (Fig. 1). 

The ovaries of juveniles reached their 
maximum average weight in November, 
approximately 3 months prior to the peak 


of the breeding season. The heaviest 
normal ovaries encountered were found 
during November in juveniles; the 
average weights are shown in Table 5 
and Fig. 5 and 6. In October, Novem- 
ber, and December the ovaries of juve- 
niles weighed more than the ovaries of 
parous raccoons. The average weights 
of ovaries for the two groups of females 
in January were practically identical 
(Fig. 5). 

The ovaries of juvenile (nulliparous) 
females showed a significant decline in 
average weight from November through 
January, and perhaps through March, 
but the sample sizes, for February, March, 
and April were too small to be definitive. 
The small sample of juveniles for these 
latter 3 months resulted partly from classi- 
fying raccoons as nulliparous (juveniles) 
or as parous or pregnant. During those 
3 months many females approximately 1 
year of age were either pregnant or par- 
ous, and hence their ovaries were placed 


Table 5.—Average weights of ovaries by month in the raccoon.” 


Nulliparous Parous or Pregnant 
Average Average 
Ovaries Error Ovaries Ergor 
in Milligrams in Milligrams 
April (at birth) 4.2 (2) 0.0 4.2- 4.2 282 (3) 31 249-344 
May* 10.9 (3) 0.4 10.4— 11.8 217 (4) 50 164-259 
June 51 (3) 16 20 ease 224 (2) 43 181-266 
July 121 (6) 24 45 —222 Ale C7)) 30 93-327 
August 118 (7) 14 82 -180 198 (4) 42 76-260 
September 184 (14) 23 66 —386 217 (3) 27 164-256 
October 253 (14) 25 147 -524 227 (4) 25 163-295 
November 312(195) 9 59 —970 271 (80) 12 102-613 
December 295 (186) 13 78 -699 229 (81) 10 82-555 
January 239 (45) 15 100 -452 244 (25) 23 92-655 
February 189 (3) 54 112 —294 275 (5) 41 132-381 
March 124 (1) : scnc 246 (5) 57 74-382 
April 260 (1) ood Bree S000 
May* 167 (1) 
August 270 (1) roar So0.e 
September 149 (2) 70 78 -219 
November 367 (2) 70 225 -—410 
December 233 (12) 27 119 -313 
January 261 (3) 43 204 —346 
March 136 (1) ants 


* All raccoons were collected in Illinois, July 18, 1958 through April 18, 1961. 


> The numbers of observations are in parentheses. 


© Most nulliparous juvenile raccoons are approximately 1 month old in May. 
4 Most nulliparous adult raccoons are approximately 13 months old in May. 


300 


200 


is 


MILLIGRAMS 


100 


MAY JUNE JULY AUG. SEPT. 


Fig. 5.—Seasonal variations in the average 
nant and nulliparous raccoons. With each mean 


representing the mean plus or minus one standard error. 
The data are given in Table 5. 


July 18, 1958 through May 30, 1961. 


with the adult group. Hence, nulliparous 
females probably represented only females 
that did not reach sexual maturity at 
approximately 1 year of age. All females 
less than 1 year of age in the January 
sample were counted as juveniles, as no 
pregnant female was found during that 
month. 


50 


Inuinois NaturaL History SurvEY BULLETIN 


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Vol. 31, Art. 2 


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total weight of both ovaries from parous or preg- 
are the number of observations and a vertical line 
All animals were taken in Illinois from 


The seasonal weights of the ovaries in 
parous raccoons (Table 5) followed a 
pattern similar to that found for the 
gonads of adult males (Table 1). The 
minimum average weight was reached 
during July, with a slow but consistent 
increase in weight during August, Sep- 
tember, and October. The fall peak 


81 


APR. 


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as By WEIGHT : 

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123456789) 12345678910 12345678 910 
NOVEMBER DECEMBER JANUARY 


WEIGHT CLASSES 


Fig. 6.—The average total weights of both ovaries from parous or pregnant and nulliparous 
raccoons. All animals were taken in Illinois from November 1958 through January 1961. The numerals 
at the tops of the bars indicate the numbers of animals in the Parous or pregnant and nulliparous 
groups. 


{ 
/ 


oh 


July, 1973 Sanperson & Natsanpov: RepropucTivE CycLe OF THE Raccoon 45 


weight was reached in November. The 


o 
cS elaaws a ovaries of parous raccoons declined sig- 
2 £ Aas = nificantly (P<0.01) in average weight 
= 2/3g8s 2 from November to December but again 
4 as me increased in weight during January. By 
= April the ovaries of parous raccoons had 
e reached their peak average weight for 
= a Se the year, slightly heavier than in Novem- 
ra nS mae ber. The average weight of adults’ 
° ailaan a ovaries in April was a little more than 1.6 
2 Ey times their average weight in July, in 
= 3]. contrast to the approximately 2.8-fold 
= .|7|a\e@ee 2 increase in average weight reported for 
z || & § La~ 2 the testis in the adult between the low 
is bb 8 = 3 2 oS ge of July and the high of Decem- 
= 3 er. 
- . From the study of ovaries collected 
= 2 Bille a during all months and seasons, we gained 
g U5 gleee 8 the impression that differences in weight 
Ee Io z\noa a existed from month to month and year 
3g | 4 Z|SNS 5 to year. For example, the total weight of 
Fi | both ovaries of parous females averaged 
3 : near! i 
y 350 mg in November 1958 but only 

: &| )» &|oo5 S 219 mg in November 1959 (Table 6). 
= RES g{/Le& < Ovaries from nulliparous females killed 
8 a < 3 a5 < in November 1958 also weighed consid- 
8 i erably more on the average than did 
3 e 3 ovaries es peas cra killed 
5 g Vv in November . Less striking varia- 
: g g aa a ; tions were noted for other months and 
é a|naonm + = years. There were also annual differences 
E g | |Saa a = in the average weights of ovaries from 
3 F 3 parous females but no significant differ- 
2 Bult — ences in those from nulliparous females. 
ia B/s--> 5 shes 
3 : sats) < SS2 MEAN BIRTH DATE OF LITTERS 
5 Ajaas st XMS Wood (1955:409-410) concluded that 
£ eeee 7 of the 16 females he examined in 
2 7 2 5 53 Texas had mated by the end of February, 
3 2 asa Ss fone but the earliest pregnancy he recorded 
= 5 Sot £ ibe was March 18. George & Stitt (1951: 
: Z|saa a $8225 218) found three litters that were born 
3 225.2 during March 1950 in Michigan after an 
3 Eeees unseasonably warm January. Berard 
> Egeey (1952:248) observed a lactating female 
3 scees in West Virginia that he estimated had 
T 2>>8 given birth no earlier than August 15, 
oe SEaa¢ whereas normal births in that area usu- 
23 2Ss.y& y|f8se"F ally occur before May 15. Dorney (1953: 
23 § SSASE ~ £26222 123) weighed young raccoons taken in 

© a BoaS58 8 8|/***** Wisconsin from November 25 through 

= a SS oS 


December 22, 1950 and concluded that 


46 Intino1s NaTuRAL History SURVEY BULLETIN 


Vol. 31, Art. 2 


Table 7.—Months of birth of raccoons in the northern half of Illinois as determined by actual 
births or as estimated from examination of embryos.” 


Number of Litters 
Conceived in 
Captivity That 
Were Born in Month 
Designated 


Month 


January 
February 
March 
April 
May 
June 
July 
August 


COoOrFWUNOCO 


Number of Adult Wild 
Females Examined 
for Pregnancy? 


Number of Litters 
Conceived in the 
Wild That Had 

Actual or Potential 

Birth Date in Month 


Designated* 

202 0 
6 0 
15 5 
18 12 
9 1 

6 2 

11 0 
4 0 


8 All embryos were examined between April 2, 1957 and June 24, 1961. 
»b Many nonpregnant, adult females examined from April to August were lactating. 


¢ Potential birth dates were estimated (Fig. 8). 


“a sizable percentage” of the young had 
been born later than usual in that year. 
He suggested that the cold spring weather 
in 1950 had decreased raccoon mobility 
and thus had decreased the normal num- 
ber of early conceptions. A similar situ- 
ation, discussed later, apparently occurred 
in Illinois during the breeding season in 
1960. 

The reports cited emphasize the varia- 
tion in birth dates that is normal in the 
raccoon. Most raccoons in the northern 
half of Illinois are born during April 
(Table 7). The mean date of birth for 
20 litters conceived in the wild, 7 of 
which were born in captivity, was April 
18; the earliest date of birth was March 
9, and the latest, June 24. The potential 
birth dates of embryos measured in dead 


Table 8.— Estimated number of days prior to 
birth based on the measurement of uterine swell- 
ings in captive raccoons. 


Largest Measurement Days Prior 
of Uterine Swellings to Birth 
in Millimeters 
8 55 
20 39 
25 35 
30 33 
35 33 
40 30 
45 23 
55 17 
110 1 


females were estimated by measuring 
uterine swellings (Table 8 and Fig. 7). 

The mean date of birth for 11 litters 
conceived and born in captivity in Ur- 
bana, Ill., was April 24. The monthly 
distribution of these births is shown in 
Table 7. The earliest date of birth for a 
litter conceived in captivity was March 
16; the latest was June 3. 

Although 202 wild females were ex- 
amined for pregancy during January and 
21 were examined during February, July, 
and August (Table 7), only one pregnant 
female was observed during these 4 
months. When examined on February 25, 
she appeared to be due to give birth in 
about 3 weeks. 


ESTIMATING BIRTH DATES 
OF RACCOONS 


Uterine swellings were measured to the 
nearest millimeter and the dates of birth 
were recorded for eight litters believed 
to have been born at full term (Table 8). 
(One litter was measured on two occa- 
sions.) Because it was difficult to mea- 
sure accurately the crown-rump length of 
embryos, especially during the early stages 
of pregnancy, the measurement recorded 
was the greatest measurement of the ex- 
ternal uterine swelling. In the early 
stages the swellings were essentially 
round, and the measurement was greater 
than the crown-rump measurement of 


July, 1973 Sanperson & NaLBaNnpov: REPRODUCTIVE CYCLE OF THE Raccoon 47 


LARGEST UTERINE SWELLINGS (MM) 


O 10 


20 


30 40 50 


DAYS PRIOR TO BIRTH 


Fig. 7.—Sizes of uterine swellings in raccoons at various numbers of days prepartum. 
was fitted by least squares, not including Llewellyn's data. 
to conception 63 days prepartum, is not based on data. 
conception was 5 mm, the approximate average diameter of the uterus during estrus. 


given in Tab!e 8. 


the embryo. In the later stages the swell- 
ings were elongate and the measurement 
approximated that of the crown-rump 
measurement. 

The dates of mating were not known, 
but it was possible to graph the size of 
the uterine swellings in relation to the 
number of days prior to parturition (Fig. 
7). The line was fitted by least squares 
and gave a good fit for uterine swellings 
between 20 and 60 mm in size. When 
we used this line to estimate the dates 
of birth for eight litters, the maximum 
error was 4 days when the uterine 
swellings were between 20 and 60 mm. 
In one litter uterine swellings larger 
than 60 mm were measured and in 
another litter uterine swellings smaller 
than 20 mm were measured. Measure- 


The line 

The dash line, an extension of the line 
The size used for the uterine swelling at 
The data are 


ments of the swellings in these two litters 
appear to indicate slower-than-average 
growth from conception to the 20-mm 
size and faster-than-average growth from 
the 60-mm size to birth. Measurements 
of uterine swellings made during this 
study were similar to those reported by 
Llewellyn (1953:321). Our data and 
Llewellyn’s make it possible to estimate 
the date of birth (Fig. 7). If we assume 
a gestation period of 63 days, which many 
authors agree is average for the raccoon, 
it is possible also to estimate the date of 
conception. 


SECONDARY SEX RATIOS 


Incidental information collected during 
the present study indicated that the sex 


48 Inuinois NaTurAL History SuRvEY BULLETIN Vol. 31, Art. 2 ; 


Table 9.—Secondary sex ratios in raccoons.” 


Sex of Embryos 
and of 
Young at Birth” 


Sex of Wild 
Litters Less Than 
2 Months of Age 


Percent Percent 
Females Males Males Females Males Males 
47 36 43° 21 33 61° 
x°=1.46 x°=2.67 
(P<0.25) (P<0.10) 


a All were from Illinois, taken during the breeding seasons of 1957 through 1961. : : 
> Includes young conceived in the wild and born in captivity as well as young conceived in captivity. 
¢ Neither group is significantly different from 50:50 (P > 0.05); however, the ratios of the two groups may 


be different from each other (P < 0.06). 


ratio of raccoon embryos and of young 
at birth, combined, and the sex ratio of 
young raccoons less than 2 months old 
were not significantly different from 50: 
50 (Table 9). There were more males 
(P<0.06) among the young less than 
2 months old than among the embryos 
and young at birth, indicating some 
differential mortality of females be- 
tween birth and 2 months of age. Other 
investigators have examined limited num- 
bers of young raccoons at birth or prior 
to 2 months of age to determine second- 
ary sex ratios. Stains (1956:31) reported 
that the sex ratio was approximately 1:1 
at birth in the raccoon, as shown by 
counts of litters. Stuewer (1943a:213; 
19436:68) counted all of the young in 
eight litters ranging in age from 7 to 
60 days and found 14 males and 19 
females (42 percent males). 


ESTROUS CYCLE, OVULATION, 
AND PSEUDOPREGNANCY 


Estrous Cycle 


Published reports regarding the estrous 
cycle in the raccoon do not agree. The 
raccoon has been reported to have one 
heat period and one breeding season 
each year (U.S. Department of Agricul- 
ture 1936). Stuewer (1943a:212) found 
that occasionally an adult female failed 
to mate successfully in spring and then 
bred later, but that yearling females ei- 
ther mated during the regular breeding 
season or did not mate until the next 
breeding season. Asdell (1946:135) re- 
ported: “In New England mating begins 
in the last week of January and there 


may be a later season for young females | 
....” Whitney & Underwood (1952:83) 
stated that, on the basis of actual obser- 
vations under normal conditions, the 
period of mating in the raccoon was from 
January until March; if this period was 
missed, another normal period of 2 
months’ duration occurred 4 months later. 
During his studies of raccoons in Texas, 
Wood (1955:409) found Graafian folli- 
cles in the ovaries of one female examined 
in April and thus concluded that breeding 
can occur as late as April. Because ovu- 
Jation normally occurred early in the 
year, this finding suggested to him that 
raccoons were possibly polyestrous. 
Observations made on captive raccoons 
held in outdoor cages in Urbana, IIL, 
confirmed Stuewer’s (1943a:212) obser- 
vations on the absence of delayed breed- 
ing in yearling females. General obser- 
vations made on several captive, yearling 
females showed that either they became 
pregnant or pseudopregnant at the time 
when adults became pregnant, or they did 
not breed until the next breeding season. 
These observations were confirmed by 
examining the ovaries of two yearling fe- 
males several times from March through 
August. Their ovaries and uteri remained 
small and inactive during the entire peri- 
od. Thus, we believe that delayed breed- 
ing in yearling females does not contrib- 
ute substantially to the number of litters 
born later in the year than usual. 
Dates of ovulation were estimated by 
observing birth dates and by direct ex- 
amination of Graafian follicles, corpora 
lutea, and embryos. Heat and ovulation 
probably occurred at about the same time. 


July, 1973 SanpErson & NaLBanDov: REPRODUCTIVE CYCLE OF THE Raccoon 49 


Tab'e 10.—Approximate number of days between ovulations in five captive raccoons. 


Estimated Estimated 


Remarks 


Date of Date of Days 
5 Between 

First Second eae 

Ovulation Ovulation 2GOns 
2-10 5-10 89 
Before 3-14 5-26 70° 
Before 3-10 Pali! 62" 
52 (==2) 6-23(+2) 84 
1-29 6-16 141° 


During first pregnancy, carried embryos half 
way or more to term but resorbed them. 
Pseudopregnant 

Pseudopregnant 

Pseudopregnant 

Carried embryos to term each time. Young 
of first pregnancy all dead 4 days postpartum. 


® Minimum. 


» The interval between the births of two litters in one season. 


On the basis of our observations of five 
captive raccoons for which the approxi- 
mate dates of the first and second ovula- 
tions were known (Table 10), we found 
that the interval between ovulations in 
captive raccoons in Urbana, IIl., varied 
approximately from 80 to 140 days—and 
not invariably 4 months, as reported by 
Whitney & Underwood (1952:83). The 
shorter intervals that we observed agree 
with Millard’s (1939:28-29) data. He 
obtained two litters in one breeding sea- 
son from 6 of 10 captive raccoons in Wis- 
consin whose young were removed on 
the day of birth and whose mates were 
returned 3 days later. Seven of the fe- 
males were observed to mate 10-16 days 
after the young were born. If we assume 
a gestation period of 63 days and that 
the female raccoon ovulates on the day of 
mating, ovulations in Millard’s animals 
occurred 73-79 days apart. 

Under normal circumstances wild. 
adult female raccoons in Illinois rarely 
skip a breeding season. Special circum- 
stances may interfere with the regular 
breeding cycle, causing a_higher-than- 
normal percentage of the litters to be born 
late (Dorney 1953:123). Such interfer- 
ence occurred in some sections during the 
1960 breeding season in Illinois. Temper- 
atures at the Urbana and Peoria weather 
stations (U.S. Weather Bureau 1960) 
were average for January 1960, but mean 
temperatures in February were 15.7° F 
[9.0° C] below normal. Snowfall at 
Urbana and Peoria for February and 
March 1960 ranged from 8 to 16 inches 
[20.3-40.6 cm] per month higher than 


the average for the preceding 10 years. 
On the Allerton Park Study Area (Piatt 
County, east-central Illinois) young rac- 
coons were caught in live traps beginning 
in early June of each year from 1957 
through 1961, with the exception of 1960. 
In 1960 the first young were livetrapped 
after September 1 even though trapping 
was conducted during the entire summer. 

Eyes were collected from 257 juvenile 
raccoons killed by hunters and trappers 
over a wide area centered around Farm- 
ington in west-central Illinois during the 
1960-1961 hunting season. The lens tech- 
nique (Sanderson 1961b:482-485) was 
used to estimate birth dates. The lenses 
indicated that the peak of births in 1960 
occurred in mid-April, the usual time, 
but that a second, smaller peak occurred 
at the first of July, about 11 weeks later. 
These two peaks were separated by about 
the length of one estrous cycle, as it 
was estimated from our observations of 
captive raccoons. According to the lens 
data, approximately 16 percent of the 
young were born during August, Septem- 
ber, and October in 1960—later than the 
latest date of birth reported in Table 7— 
indicating that under some circumstances 
a substantial number of wild raccoons 
have had more than one estrous cycle in 
a year. 

In view of Millard’s (1939:28-29) 
success in getting two litters in one season 
from captive raccoons and because the 
present study demonstrated that some of 
our captive pseudopregnant and pregnant 
females had second heat periods in cap- 
tivity, it was, at first, surprising that so 


50 Ittinors NaturaL History SuRVEY BULLETIN 


few second litters were conceived in cap- 
tivity during our study. Millard’s (1939) 
objective was to rear a large number of 
young raccoons for restocking purposes, 
and no doubt he disturbed his animals 
as little as possible. Our study, on the 
other hand, required frequent handling 
of the animals and their subjection to 
laparotomies. Only two pregnancies are 
known to have resulted from second ovu- 
lations during our study. The first female 
became pseudopregnant after her first 
ovulation, and the single embryo from 
her second ovluation was resorbed; the 
second female give birth to her second 
litter in August, 141 days after the first 
litter was born. She had killed the last 
surviving young of her first litter 4 days 
postpartum. 

Female raccoons will not ovulate and 
come into estrus so long as they are nurs- 
ing young. Young were removed at birth 
from four female raccoons and 5 days 
after birth from one female. All of these 
females were returned to their mates 
when the young were removed, but no 
second matings were observed and no 
second pregnancies resulted. Young were 
removed from six females at periods vary- 
ing from 17 days to 6 weeks after birth, 
and the males were returned to the fe- 
males. One female was given a drug that 
caused her to abort or resorb her young. 
Her mate remained with her at all times. 
However, no second pregnancies resulted 
in any of these animals. In addition to 
these females several others underwent 
periods of pseudopregnancy during the 
normal breeding season while remaining 
with their mates through the summer. 
No late pregnancies resulted. Possibly 
some of the males were no longer capable 
of fertilization (Fig. 3 and 4) by the time 
their mates experienced their second es- 
trous cycles. 

Our data make it clear that in Illinois 
it is possible for raccoons to ovulate two 
times during one season and even to give 
birth to two litters. However, to give 
birth to the second litter, the female must 
lose her first litter on or shortly after the 
day of birth. We have no evidence that 


Vol. 31, Art. 2 


ovulations occur after lactation ceases in 
the raccoon. In any case, it appears that 
raccoons must nurse for 2-3 months in 
the wild and that probably they usually 
nurse for 3-5 months (Stuewer 1943a: 
213; Montgomery 1969: 155-158). 

The vaginal smear is frequently used 
to determine the stage of the reproductive 
cycle in the laboratory mouse, rat, and 
guinea pig. Stockard (1932:1612-1627) 
gives a general review. Although +this 
technique can theoretically be applied to 
other species, many problems occur with 


species that have relatively long periods \ 


of proestrus and estrus. Nalbandov 
(1958:103-104) pointed out that all 
mammalian females show changes in 
their vaginal histology during the estrous 
cycle. He further reports: 
“The vaginal-smear technique is 
most useful, however, with animals 
having short estrous cycles . . .; in 
animals with longer cycles . . . vagi- 
nal changes lag from one to several 
days behind ovarian changes, and 
vaginal smears are therefore less re- 
liable indicators of ovarian events.” 

Stuewer (1943b:64) observed that 
from 1 to 2 weeks elapsed from the onset 
of vaginal swelling in the raccoon until 
the female would receive the male. After 
a receptive period of about 3 days, 3 or 
4 weeks elapsed before the vulva returned 
to normal appearance. Whitney & Un- 
derwood (1952:83) reported that the 
onset of the mating cycle could be recog- 
nized by a thickening or swelling of the 
vagina and vulva and traces of bloody 
fluid (absent in some females), and that 
the female would accept the male at the 
onset of-the mating cycle and was recep- 
tive for a period of 3-6 days. 

We obtained estrous-type vaginal 
smears for a period of several weeks in 
our raccoons; examples of these smears 
are shown in Fig. 8. An estrous-type 
smear was obtained from a castrated fe- 
male 36 days after the end of treatment 
with estradiol and progesterone (Fig. 
8B). One captive female must have mat- 
ed during the 7 days between the taking 
of two vaginal smears; she gave birth 


i ees | 


July, 1973. Sanperson & NaLBanpov: ReEpRopucTIVE CycLe oF THE Raccoon 51 


Fig. 8.—Vaginal smears from captive raccoons representing various stages of the estrous cycle. 
A, female 1292; ovaries removed May 14, 1958; smear taken July 23, 1959. B, castrated female 
1297; second ovary removed August 13, 1957; smear taken May 16, 1958, 36 days after treatment 
with estradiol and progesterone ended. C, nulliparous female 2525; smear taken December 4, 1958. 
D, female 2959; smear taken January 11, 1958. E, female 2959; smear taken March 7, 1958. 
F, female 2959; smear taken March 14, 1958; female 2959 must have mated between these two 
dates, because she gave birth to young on May 13, 1958. G, adult female 1786; smear taken April 
12, 1957. H, female 2114; smear taken February 21, 1958; uterine swellings were 8 mm in diameter. 
The smears were stained with Wright's blood stain and are shown 62 times actual size. 


52 InLino1is NaturAL History Survey BULLETIN 


60 days after the second smear was taken 
(Fig. 8E and F). On the basis of the 
results obtained from this study, we 
conclude that the days when a female 
raccoon will receive a male can not be 
identified by examination of vaginal 


ii 
) 


. 


Vol. 31, Art. 2 


smears. The vaginal smear appeared to 
be no more specific than gross vulval 
swelling—-which can be observed much 
more readily. Leucocytes (Fig. 8C and 
G) were seen in vaginal smears from rac- 
coons only infrequently. The paucity of 


July, 1973 Sanperson & NaLsanpov: ReEpRopucTIVE CycLE OF THE Raccoon 53 


vaginal smears containing leucocytes sug- 
gested that the raccoon may pass through 
metestrus in a relatively short time. 

Many of the difficulties inherent in 
using vaginal smears may be avoided by 
taking vaginal tissue for biopsies—a 
simple procedure in the raccoon. Sam- 
ples of vaginal tissues were removed 
from both anestnetized and unanesthetiz- 
ed animals (Fig. 9). However, vaginal 
tissues from castrated females (Fig. 9A 
and B) showed that the histology of the 
vaginal epithelium is not a reliable indi- 
cator of estrus in the raccoon. 


Ovulation 


Whitney & Underwood (1952:84), 
without citing evidence, reported that in 
the raccoon “sufficient stimulation is 
produced during copulation to insure 
ovulation.” Llewellyn & Enders (1954a: 
440) removed one ovary from each of 
four sexually mature raccoons that had 
been isolated from males before and 
during the normal breeding season. They 
found “well developed follicles” in each 
ovary but no corpora lutea and, on the 
basis of this evidence, suggested that 
ovulation in the raccoon is not spontane- 
ous but is induced by copulation. 

In our study one female was approxi- 
mately 5 months old when captured about 
5 months before the breeding season. 
She was isolated for 3 months prior to 
the breeding season. Her nipples were 
moderately stimulated but unpigmented 
when a laparotomy was first performed 
on her 3 months after she was isolated. 
The ovaries were each 8 X 5 mm—small 
for ovaries with corpora lutea—yet each 


ovary had two corpora lutea, each 5 mm 
in diameter. Thirty-five days later the 
corpora lutea were essentially unchanged 
in size and appearance. Sixty days after 
the first examination the ovaries were 
10 X 5 mm, and the corpora lutea were 
slightly paler and were between 3 and 
4 mm in diameter. Ninety-three days after 
the initial observation no traces of the 
corpora lutea were visible. 

A second female that was isolated was 
approximately 40 days old when cap- 
tured. She was isolated 2 months prior 
to the breeding season, and the first lap- 
arotomy was performed on her 2 months 
after the isolation began. Her nipples 
were Only slightly stimulated, and her 
ovaries, measuring 8 X 4 mm, contained 
no corpora lutea. The left ovary had one 
clear follicle and the right ovary had 
two, each follicle measuring 2 mm in 
diameter. 

Nine days later the right ovary had 
four freshly ovulated follicles, each 3 mm 
in diameter. The left ovary had a single 
follicle of the same size with a tiny 
hole in its highest point. It was believed 
that this female had ovulated no more 
than 2 days earlier. 

Twenty-six days after the freshly ovu- 
lated follicles were observed, the right 
ovary was 11 X 5 mm and had three 
corpora lutea, each 4 mm in diameter. 
Either the fourth follicle in the ovary 
did not form a corpus luteum, or it was 
obscured by one of the other corpora. 
The left ovary was 10 X 5 mm and had 
one corpus luteum of the same size as 
those in the right ovary. 

Eighty-one days after the freshly ovu- 


Fig. 9 (Page 52).—Photomicrographs of vaginal biopsies from captive raccoons representing various 


stages of the estrous cycle. 


A, female 1292; ovaries removed May 14, 1958; biopsy performed July 23, 


1959. B, castrated female 1297; second ovary removed August 13, 1957; biopsy performed May 
16, 1958, 36 days after treatment with estradiol and progesterone ended. C, female 2114; biopsy 
performed February 21, 1958; uterine swellings 8 mm in diameter. D, nulliparous female 2525; 
biopsy performed December 4, 1958. E, female 1297; biopsy performed July 23, 1959; no treatment 
with estradiol and progesterone after April 3, 1958. F, female 1298; biopsy performed October 5, 
1959; ovulated about September 24, 1959 as a result of injections of pregnant mare's serum. G, 
female 1782; biopsy performed October 29, 1957; treated with estradiol beginning October 17, 
1957; ovaries removed June 10, 1957. H, female 2959; biopsy performed January 27, 1960; ovulated 
after December 18, 1959 as a result of treatment with follicle-stimulating hormone and luteinizing 
‘hormone; corpora lutea present. I, female 2805; biopsy performed February 24, 1960; fresh corpora 
lutea present. The sections were stained with hematoxylin and eosin and are shown 122 times 
actual size. 


54 Intino1is NaturaL History SurvEY BULLETIN 


lated follicles were observed, the ovaries 
were each 8 X 3 mm;; the four corpora 
lutea, each now 3 mm in diameter, were 
still present. By 102 days after the cor- 
pora were first seen, four whitish corpora 
albicantia (not examined histologically) , 
each measuring 1 mm in diameter, had 
formed at the sites of the preceding cor- 
pora lutea. At this time there were also 
two follicles, each 2 mm in diameter, 
in the right ovary and one of similar-size 
in the left ovary. The female was judged 
ready to ovulate a second time. 

Thirty-one days later one corpus lute- 
um was found in the left ovary and five 
or more were found in the right ovary; 
each corpus luteum was approximately 
5 mm in diameter. The left ovary was 
removed (133 mg) and sectioned, but 
no ovum was found in the corpus luteum. 
Thus, this animal had probably ovulated. 
Also, the secretory material in the endo- 
metrial glands indicated that progesterone 
had been secreted. Seventy-three days 
after the follicles were examined, the 
corpora lutea were still present but mea- 
sured only 2 mm in diameter. 

A female found when approximately 
3 weeks old was kept as a house pet until 
the middle of April, when she was about 
1 year of age. At that time she suddenly 
became vicious, severely biting both own- 
ers. She remained the most vicious rac- 
coon we have seen among the many 
dozens of wild, captive, and pet raccoons 
that we have handled. According to her 
owners, she had never come into contact 
with other raccoons. At the time her 
behavior changed, her nipples were mod- 
erately stimulated and moderately pig- 
mented, indicating that she was either 
pregnant or pseudopregnant. When first 
examined, her ovaries were 9 X 6 mm 
and 7 X 5 mm, respectively, and con- 
tained a total of five corpora lutea, each 
3 mm in diameter. The size of the cor- 
pora indicated that they were regressing 
when examined, because newly formed 
corpora lutea in the ovaries of raccoons 
are approximately 5 mm in diameter. 
Thirty-five days after the initial examina- 
tion all five corpora lutea were plainly 
visible but were regressing and were 


Vol. 31, Art. 2 


slightly smaller than when first examined. 
Forty-eight days later (83 days after 
the first examination) no traces of the 
corpora lutea could be seen by gross 
examination. 

The data on these three isolated fe- 
males, one of which ovulated twice in 
one season, show that the raccoon is a 
spontaneous ovulator, and refute Llewel- 
lyn & Enders’ (1954a:440) interpretation 
of their observations. The statement of 
Whitney & Underwood (1952:84) that 
ovulation in the raccoon is dependent 
upon copulation is not true for captive . 
raccoons in Illinois. 

In many captive raccoons, especially 
those reared as pets, the onset of estrus 
and pseudopregnancy was apparent from 
changes in behavior. A docile house pet 
sometimes suddenly became vicious and 
unmanageable. In all such cases that 
we examined, corpora lutea were present 
in the ovaries. The formation of corpora 
lutea was invariably accompanied by 
changes in the uteri and nipples whether 
the animal was pregnant or only pseudo- 
pregnant. The nipples always enlarged, 
and some became heavily pigmented, 
some became only slightly pigmented, and 
still others remained unpigmented. With 
the onset of pseudopregnancy the uteri 
became turgid and opaque and were 
considerably enlarged from their size dur- 
ing anestrus; however, they were not flu- 
id filled and somewhat rubbery, as they 
were during estrus. 

A female raccoon born in April was 
reared as a pet until the following Janu- 
ary, when she became too unruly for 
the owners to handle and was donated to 
our project. She is described here as 
representative of the females, housed with 
other raccoons, that ovulated but did not 
become pregnant. According to her own- 
ers, she had not come into contact with 
other raccoons before she was donated 
to our project. Two days after we re- 
ceived her, she was placed in a cage 
with four yearling males. Forty-nine days 
later her nipples were tiny and white, 
but after 21 more days (March 28) they 
were elongated and black, indicating that 
she was either pregnant or pseudopreg- 


July, 1973. SanpeRson & NatBanpov: RepRropuctivE CycLe or THE Raccoon 55 


nant. Five days later her left ovary was 
removed, and histological examination 
showed four freshly formed corpora 
lutea. 

Histological examinations of the ova- 
ries from this nonisolated female, from 
one isolated female, from three nonpreg- 
nant wild females, and from two addi- 
tional nonisolated, nonpregnant, captive 
females revealed no ova in the corpora 
lutea. No substantial difference was not- 
ed between the corpora lutea of the iso- 
lated nonpregnant and of the nonisolated 
nonpregnant females. Ovaries from sev- 
eral nonpregnant females housed with 
other females, or with males, were ex- 
amined during and after the breeding 
season. In several cases these ovaries had 
corpora lutea, which were grossly identi- 
cal to those seen in females isolated prior 
to the breeding season and to corpora lu- 
tea in pregnant females. Thus, we con- 
cluded that corpora lutea in both isolated 
pseudopregnant and nonisolated pseudo- 
pregnant females formed from ovulated 
follicles and not from luteinization of fol- 
licles. Normal-appearing corpora lutea 
were also formed in the ovaries of a fe- 
male in which ovulation was induced by 
exogenous hormones. 

Data gathered from examination of 13 
captive raccoons indicated that corpora 
lutea persist in pregnant females until 
parturition. Observations on four of these 
captives indicated that corpora lutea dis- 
appeared 14-16 days after parturition if 
the young were taken from the mother 
within 5 days after birth. In one of 
these four the corpora lutea were not 
present 16 days after parturition; in an- 
other they were present 14 days after 
parturition. 

One female ovulated, apparently for 
the second time in the season, about 
May 11. She mated, and one embryo 
was implanted; approximately 20 days 
after ovulation the embryo was dead. 
Traces of one corpus luteum were still 
present in each ovary approximately 52 
days after ovulation, and, on the basis 
of size and appearance, we concluded 
that they undoubtedly persisted for a 
maximum of 60 days. 


In five nursing females the corpora 
lutea disappeared before the ovaries were 
examined from 11 to 35 days postpartum. 
A sixth female examined 11 days after 
parturition had four regressing corpora 
lutea, each 3 mm in diameter, in her left 
ovary and none in the right ovary. The 
corpora were those observed when she 
was first examined 34 days before the 
birth of her young. She was examined 
again 20 days after giving birth, when 
only four corpora albicantia were pres- 
ent in her left ovary. Thus, in this nurs- 
ing female, the corpora lutea disappeared 
between 11 and 20 days after parturition. 
Corpora lutea were not found in histo- 
logical preparations of ovaries from two 
wild, lactating females, nor by gross ex- 
amination of the ovaries from six other 
wild, lactating females. 


Pseudopregnancy 


Our data indicate that corpora lutea 
persisted for about the same length of 
time in captive pseudopregnant raccoons 
as they did in those that give birth to 
young. Corpora were present 61 days but 
not 82 days after the estimated date of 
ovulation in one pseudopregnant captive. 
Three other pseudopregnant females 
showed similar periods of pseudopreg- 
nancy, although the data for these fe- 
males were less precise than were the 
data for the first. The persistence of 
corpora in females that went at least half- 
way to term did not appear to differ sig- 
nificantly whether the young were abort- 
ed, were resorbed, or were born and 
were removed at birth or nursed until 
weaned. In one female, discussed in the 
preceding section, the young were re- 
sorbed at an early stage and the corpora 
lutea disappeared no more than 60 days 
after ovulation. 

In some species pseudopregnancy may 
equal normal pregnancy in duration, but 
in most animals it lasts about half as 
long (Nalbandoy 1958:218). Our obser- 
vations indicated that all captive rac- 
coons that ovulated, but did not become 
pregnant, underwent a period of pseu- 
dopregnancy much as does the dog. In 
the raccoon pseudopregnancy lasted ap- 


56 Inuino1s NaTurAL History SuRvEY BULLETIN 


proximately the same length of time as 
does normal pregnancy and followed 
ovulation. 

Our observations of wild female rac- 
coons during the breeding season indi- 
cated the relative incidence of pregnan- 
cies and pseudopregnancies, and supplied 
substantiating evidence that corpora 
lutea disappear in wild, lactating females, 
as in captives, shortly after they have 
given birth. Histological sections were 
made of ovaries collected from March 
through June (1957 through 1961) from 
15 wild females 2 years of age or older. 
Six were pregnant, four were pseudo- 
pregnant, and five had recently given 
birth. Corpora albicantia were present in 
the ovaries of four of the five parous 
females, and corpora lutea were present 
in all of the pregnant and pseudopreg- 
nant animals. The fifth parous female, 
collected March 1, had recently given 
birth or aborted, as indicated by the 
fresh placental scars in her enlarged uter- 
us and the four corpora lutea in her 
ovaries; however, she was not lactating. 
Corpora lutea were not found in histo- 
logical sections of the ovaries from 39 
young-of-the-year, 14 yearling (12-20 
months old), and 10 adult wild raccoons 
collected from July through January. 

Of 15 wild, parous female raccoons 
collected from Februray through Sep- 
tember, only 1 had freshly ovulated folli- 
cles in February, and another had cor- 
pora lutea in March. None of the re- 
maining 13 females, including 6 that 
were lactating, had corpora lutea. As 
mentioned earlier, histological examina- 
tions of ovaries from lactating, captive 
females indicated that corpora lutea dis- 
appear between 11 and 20 days after 
parturition, regardless of whether the fe- 
males nurse their young. 

From February through June (1957 
through 1961) we made 30 observations 
on 24 captive female raccoons 2 years 
of age or older. Five were caught only 
a few days prior to examination. Of the 
30 observations, 18 were of pregnant ani- 
mals, 8 were of pseudopregnant females, 
and 4 were of animals neither pregnant 
nor pseudopregnant when examined. The 


Vol. 31, Art. 2 


one animal that accounted for two of the 
four latter observations had an abnor- 
mally large uterus but inactive ovaries in 
1959. Her uterus was enlarged but her 
ovaries were small when she was exam- 
ined in May of 1957. Thus, she did not 
represent the norm. The other two ob- 
servations of females that were neither 
pregnant nor pseudopregant were of adult 
females that had given birth to litters in 
previous years; each was examined once 
during subsequent mating seasons. Be- 
cause each was examined only once dur- 
ing the breeding season of the year in: 
which corpora lutea were not found, it 
is conceivable that they had undergone 
pseudopregnancy but that the corpora 
had regressed before they were examined. 
Thus, evidence from both captive and 
wild females indicated that a majority of 
the females 2 years of age or older were 
either pregnant or pseudopregnant each 
year. 

Every year during the hunting and 
trapping season a small percentage of 
females, judged to have ovulated on the 
basis of the stimulated or pigmented nip- 
ples, or both, were without uterine pla- 
cental scars. During the fur seasons in 
Illinois from 1956-1957 through 1960- 
1961, uteri were examined from 284 fe- 
males that appeared, on this basis, to have 
ovulated, and 7 (2.5 percent) had no 
placental scars. The evidence indicated 
that these animals had been only pseudo- 
pregnant. Some annual variation occurs 
in this characteristic. During the 1960- 
1961 fur season, all 77 females judged, 
upon examination of their nipples, to 
have ovulated had placental scars in 
their uteri. 


PERCENTAGE OF YEARLING 
FEMALES THAT WERE 
SEXUALLY MATURE 


Of 21 captive female raccoons approxi- 
mately 1 year of age examined from Feb- 
ruary through June, 11 were either preg- 
nant or pseudopregnant, but 10 were sex- 
ually immature. Histological sections of 
the ovaries from nine wild yearlings col- 
lected from February through August 
showed no corpora lutea in the five non- 


July, 1973 Sanperson & NaLBanpov: RepropucTive Cycle or THE Raccoon 57 


pregnant females nor in the two lactating 
females, but corpora were present in the 
ovaries of the two pregnant yearlings. 
Gross examination of the ovaries from 
five wild, nulliparous yearlings collected 
from March through August showed that 
the ovaries of four contained no corpora 
lutea, but that three corpora lutea were 
present in one female collected in May. 
Thus, 10 of 21 captive yearlings and 9 
of 14 wild yearlings were sexually im- 
mature. 

During two fur seasons in Illinois 
(1959-1960 and 1960-1961) nulliparous 
adults with tiny unpigmented nipples ac- 
counted for 15 of 164 (9.2 percent) adult 
female raccoons examined. These nulli- 
parous adults, with tiny unpigmented 
mammae, probably did not ovulate dur- 
ing the first breeding season after their 
birth. 


SECRETION OF PROGESTERONE 
BY CORPORA LUTEA 


The period of the production of pro- 
gesterone by corpora lutea in the raccoon 
is unknown, but circumstantial evidence 
indicates that corpora lutea probably 
secrete progesterone as long as they are 
present (discussed later in connection 
with the production of uterine milk). 
One female had five corpora lutea, each 
5 mm in diameter, when first examined 
on April 10. At that time we traumatized 
her left uterine horn by inserting a nee- 
dle into the uterine lumen two times, 
each time scratching the entire length of 
the inside of the uterine horn- with the 
point of the needle as it was withdrawn. 
Eight days later the ovaries and corpora 
were unchanged in gross size and appear- 
ance. The left uterine horn showed no 
evidence of trauma, but there is no direct 
evidence that the uterus of the raccoon 
will respond to traumatization with a 
decidual reaction in the presence of pro- 
gesterone. 


PIGMENTATION OF MAMMAE 


Several female raccoons were studied 
to establish a possible physiological cause 
for the pigmentation or » -npigmentation 


of nipples. Some pseudopregnant yearling 
females developed heavily pigmented nip- 
ples, whereas others did not. The pres- 
ence or absence of pigment was not cor- 
related with nursing, abortion, resorp- 
tion of embryos, age at first estrus, or any 
other factors we could*recognize. Unpig- 
mented nipples remained so throughout 
life, but lightly pigmented nipples some- 
times became darker with age. The pig- 
ment was not sloughed after nursing as 
Snyder & Christian (1960:650) found 
in the woodchuck (Marmota monax). 


INTERSTITIAL TISSUE 


Many studies were conducted before 
1920 on the interstitial tissue in mam- 
malian ovaries. Interstitial tissue is 
present in greater or lesser amounts in 
the ovaries of some species and is ap- 
parently absent in others. Little is known 
about its function. His (1865) was ap- 
parently the first to describe interstitial 
tissue cells in mammalian ovaries and to 
discuss their importance. Allen (1904: 
120, 141) concluded that interstitial cells 
were formed from connective tissue dur- 
ing a process of degeneration in both the 
testis and ovary, and noted many points 
of similarity between the cells of .the 
interstitial tissue and the lutein cells of 
corpora lutea. Kingsbury (1914:86) dis- 
cussed the interstitial cells in the do- 
mestic cat (Felis catus) and recognized 
the lipoid nature of the granules in these 
cells, but he found no evidence that the 
cells constitute morphologically an intra- 
ovarian gland. He also reported their 
presence in immature, newly born, and 
fetal kittens. 

Rasmussen (1918:395) believed that 
in the woodchuck the interstitial cells 
proliferated from t rmuinal epithelium 
during adult life. 1: found a marked 
seasonal variation in the number of in- 
terstitial cells and in the amount of lipoid 
present in them in the woodchuck. These 
cells gradually increased in number dur- 
ing hibernation and hypertrophied rapid- 
ly immediately after hibernation (Ras- 
mussen 1918:371-372). Maximum num- 
bers were seen in females that did not 


58 Inurno1s NaTursL History SuRvEY BULLETIN 


become pregnant until late in the breed- 
ing season. Retrogression began with 
pregnancy and the growth of corpora 
lutea and continued until July. The ovar- 
ian interstitial cells were minimal in size 
in late summer and early autumn but 
then began to enlarge. After an exten- 
sive review of the literature, Rasmussen 
concluded, in accord with the, vast ma- 
jority of the investigators, that the inter- 
stitial cells come either directly from the 
connective tissue (stroma) of the ovary, 
or indirectly from the theca interna of 
atretic follicles. 

According to Corner (1932:1597), the 
stroma of the rabbit ovary consists so 
largely of epithelioid cells heavily laden 
with lipoid granules that the entire organ 
is a solid mass of interstitial cells in which 
the follicles and corpora lutea are em- 
bedded. This finding led to the concept 
that the ovarian stroma in this and simi- 
lar species was a gland of internal secre- 
tion, the so-called interstitial gland. Em- 
bryological study showed that interstitial 
cells were largely derived from the theca 
interna of atretic follicles and that inter- 
stitial cells were found in many species 
at a very early stage of embryonic differ- 
entiation, in which case they seemed to 
be produced by the modification of the 
cells of the stroma and of the various 
epithelial proliferations. The pig ovary 


(Corner 1932:1597) contains epithelioid. 


cells only in follicles and corpora lutea, 
the stroma cells being simply fibroblasts. 
Corner (1932: 1597) reported that the 
cat ovary was between the extremes rep- 
resented by the rabbit and pig ovaries. 
In the adult human ovary there appeared 
to be epithelioid cells only in follicles and 
corpora lutea. 

It is conceivable that interstitial cells, 
whether found in great numbers in the 
stroma of the rabbit or in thin layers in 
atretic follicles in humans, are function- 
ally the same, but proof is lacking (Cor- 
ner 1932:1597). Corner (1932:1598) 
further reported that in all of the species 
he studied the interstitial cells contained 
granules of neutral fat or, at least, of 
lipoids, which reduce osmic acid and 
stain with Sudan III. Some workers are 


Vol. 31, Art. 2 


ready to assume that the lipoids found 
in the interstitial cells represent a true 
internal secretion. 

Much of the older work, mentioned 
by Stafford & Mossman (1945:97), 
showed that in some mammals the de- 
velopment of ovarian interstitial tissue 
is at its maximum during proestrus and 
estrus and that all of the animals in- 
cluded in this group, most of which breed 
annually or semiannually, have long re- 
productive cycles. The literature reports 
no evidence of ovarian interstitial tissue 
in laboratory rodents, which have short: 
estrous cycles, and there is no easily dis- 
cernible cycle in the amount or state of 
interstitial tissue that could be correlated 
with pregnancy in the guinea pig. There 
is a trend toward a maximum amount of 
interstitial tissue in the cortex near estrus 
and into early pregnancy and a minimum 
in midpregnancy. The high and low in 
the medulla seemed to occur a week or 
two later than in the cortex, suggesting 
that in the guinea pig medullary inter- 
stitial tissue originates from that of the 
cortex. 

Patzelt (1955) studied the interstitial 
tissue in several carnivores and empha- 
sized that age, time of year, and stage of 
the reproductive cycle greatly affected 
the interstitial cells. He also pointed out 
that other investigators considered thecal 
cells, which are traced back to the par- 
ticularly active atresia of follicles during 
pregnancy, to be closely associated with 
the cells of the corpora lutea. Thus, Alt- 
mann (1927) thought it conceivable that 
only a topographical contrast existed be- 
tween thecal granulosa and lutein cells. 

Patzelt (1955) regarded the intersti- 
tial tissue cells as producers of hormones 
and as a storage place for the substance 
necessary for the formation of new folli- 
cles and for propagation in general. The 
basis for his ideas was the fact that the 
lipoid-containing cells are variously de- 
rived within the rudimentary ovary from 
germ layers, thecal cells, and cells of 
the surrounding stroma, and that it is not 
possible to demarcate the source of the 
interstitial cells. He usually found that 
ovarian interstitial cells were filled with 


July, 1973 Sanperson & NaLsanpov: ReEpropuctTiIve CycLe or THE Raccoon 59 


stored lipoids after a heat period and 
during pregnancy. After parturition a 
decrease in stored lipoids occurred that 
led to a functional dimorphism simultan- 
eously with the formation and maturation 
of new follicles. 

Hansson (1947) concluded that the 
abundance of interstitial tissue in the 
mink (Mustela vison) indicated that the 
tissue performed a special task. Because 
anestrus in the mink lasts from May to 
January, when no follicular growth be- 
yond the vesicular stage takes place, 
the interstitial tissue may serve as a regu- 
lator during this time, governing sexual 
differentiation. 

A preliminary study of the abundance 
of interstitial tissue in histological sec- 
tions of the ovaries of 119 raccoons taken 
in all months indicated that interstitial 
tissue cells were abundant at some stages 
of the reproductive cycle, often occupying 
as much as 50-90 percent of the space 
in the ovary. However, interstitial tissue 
cells were seldom abundant when corpora 
lutea were present. Of 21 pairs of ovaries 
with corpora lutea, only 3 had significant 
amounts of interstitial tissue. One of 
these is shown in Fig. 10G. 

Females less than about 2 months of 
age did not have large amounts of inter- 
stitial tissue in their ovaries. With this 
exception the ovaries of females less than 
12 months old contained more, both 
relatively and absolutely, of this tissue, 
on the average, than did the ovaries of 
older females. 

Seasonal trends in the abundance of 
interstitial tissue were apparent in ovaries 
with no corpora lutea. The ovaries re- 
moved from 16 adults from January 
through June contained little interstitial 
tissue. Ovaries removed from 26 adults 
killed from July through December con- 
tained more interstitial tissue than did 
those collected earlier in the year. No 
trend was apparent in the amount of 
interstitial tissue within the July-Decem- 
ber period. During this interval ovaries 
from adults did not contain as much 
interstitial tissue as did ovaries from fe- 
males less than 12 months old. 

Ovaries from raccoons less than 12 


months of age showed less seasonal vari- 
ation in the abundance of interstitial 
tissue than did the ovaries from older 
animals. Small amounts of interstitial 
tissue were present in ovaries removed 
from seven juveniles in May and June, 
when most young were less than 2 months 
old. The ovaries excised from 24 juveniles 
in July, August, and September contained 
more interstitial tissue than did the 
ovaries examined in May and June, but 
the differences among the amounts of 
interstitial tissue found in July, August, 
and September were slight. The greatest 
abundance of interstitial tissue was dis- 
covered in 16 pairs of ovaries taken from 
juveniles during October and November. 
The maximum ovary weights recorded 
during this study were those of juvenile 
females in November (Table 5). Nine 
pairs of ovaries were examined from fe- 
males not yet 1 year old killed during 
the period December through April. The 
abundance of interstitial tissue in these 
ovaries did not appear to differ from. 
that in the ovaries of juveniles examined 
from July through September. 

In spite of marked seasonal and age 
differences in the abundance of intersti- 
tial tissue, there was no apparent correla- 
tion between its abundance and the size 
or amount of coiling of the uterine 
glands. The development of the uterine 
glands and the presence of secretory ma- 
terial in these glands were largely de- 
pendent upon the presence of corpora 
lutea. 

Three sources of interstitial tissue have 
been suggested, germ layers, thecal cells, 
and cells of the surrounding stroma, and 
all three may be present in the raccoon. 
Small amounts of interstitial tissue were 
present in some raccoon ovaries at birth. 
Judging from appearance alone, we be- 
lieve it probable that some interstitial 
tissue in the raccoon is formed from 
degenerating follicles. Several cases simi- 
lar to the one shown in Fig. 10A were 
seen during this study. In other ovaries 
there appeared to be a streaming of the 
cells as the interstitial tissue formed, 
presumably from the germinal epitheli- 
um 


60 Inuinois Natura History SURVEY BULLETIN Vol. 31, Art. 2 


One striking feature of interstitial cells although the luteal cells were generally 
was their resemblance to luteal cells, larger (Fig. 10). Under the microscope 


July, 1973 Sanperson & NALBANDOv: REPRODUCTIVE CycLE oF THE Raccoon 61 


these two kinds of cells appeared more 
alike than the photographs in Fig. 10 
indicate. 


PLACENTAL SCARS 


Deanesly (1935:464) first reported 
that she could recognize parous uteri in 
the stoat (Mustela erminea) by the pres- 
ence of pigment granules that later work- 
ers called placental scars. Deno (1937: 
433, 445) found that placental scars 
were produced in the mouse by accumu- 
lations of hemosiderin in the cells of the 
reticulo-endothelial system and that the 
placental scars were associated with the 
involuting metrial gland. Deno (1941) 
later reported that placental scars were 
visible in both the rat and mouse for a 
year or longer. Conaway (1955:516—517) 
stated: 

“The placental scars of the rat ap- 
pear as yellow to black pigmented 
areas along the utero-mesometrial 
border. Their origin seems identi- 
cal with that of the scars in the 
mouse .... In both the rat and 
mouse, the metrial gland is a promi- 
nent structure at the base of the 
placenta .... Presumably it is formed 
by an extension of the decidual re- 
sponse into the connective tissue of 
the myometrium. The pigment-lad- 
en cells are concentrated in this area 
between the longitudinal and circu- 
lar muscle layers although some are 
found in the deeper stroma of the 
endometrium. As the age of the 
scar increases the pigmented area 
may decrease in size and appear 
darker in color.” 

Sooter (1946:69-70) counted placen- 
tal scars to determine the numbers of 
young produced by muskrats (Ondatra 


zibethicus) although no critical work has 
been done to determine whether the num- 
ber of placental scars corresponds to the 
number of young born. Elder (1952) 
reported the failure of placental scars 
to reveal breeding history in captive mink. 
Brambell & Mills (1948:241), working 
with the European rabbit (Oryctolagus 
cuniculus), again pointed out 
“that although there is little likeli- 
hood of failure to detect implanta- 
tion sites containing living embryos 
the possibility remains of the disap- 
pearance before full term of sites 
in which the embryos had died and 
were reabsorbed soon after implan- 
tation or, more probably, that such 
sites might be overlooked, through 
becoming less conspicuous, and hence 
omitted from the counts.” 

In laboratory rats and wild brown rats 
placental scars were only a crude indica- 
tion of the number of young produced 
(Davis & Emlen 1948: 166), with errors 
as high as 100 percent in either direction. 
Conaway (1955:531) found that pla- 
cental scars in the laboratory rat were 
always formed if all embryos were re- 
sorbed after the 11th day of pregnancy, 
whereas total resorption prior to this time 
never caused the formation of scars. If 
some of the embryos were resorbed, death 
on the seventh day or later resulted in 
scar formation at all resorption and term 
sites. If some embryos were resorbed be- 
tween the 8th and 11th days and the 
remainer after that, scars were formed 
at all sites. The size and appearance of 
resorption scars were similar to those of 
term scars. Momberg & Conaway (1956: 
379) found that 32 of 312 placental 
scars from previous pregnancies were 
overlapped by scars of second pregnan- 


Fig. 10 (Page 60).—Photomicrographs of luteal and interstitial cells of raccoons, showing similari- 


ties in the two. A, female 2137; luteal cells (X 94); ovary removed March 19, 1958; pregnant. B, fe- 
male 1292; luteal cells (X 94); ovary removed May 14, 1958; ovulation caused by injections of preg- 
nant mare's serum. C, female 2805; luteal cells (X 94); ovary removed February 24, 1960; fresh corpora 
lutea resulted from natural ovulations; pseudopregnant. D, female 2232; interstitial cells (X 94); 
ovary removed August 7, 1958; wild animal approximately 3 months old. E, female 2403; interstitial 
cells (X 94); ovary removed November 6, 1958; wild animal 7 months old. F, femalé 2234; inter- 
stitial cells (X 94); ovary removed August 8, 1958; wild animal 3 months old. G, female 1292; luteal 
cells (X 375); ovary removed May 14, 1958; ovulation caused by injections of pregnant mare's serum. 
H, female 2242; interstitial cells (X 375); ovary removed August 27, 1958; wild animal 4 months old. 
The sections were stained with hematoxylin and eosin. 


c 
62 Inurnors NaturAL History SurvEY BULLETIN 


cies in the white rat. They could not 
always recognize the superposed scars 
by gross examination, but microscopic 
recognition was possible. 

The placenta of the raccoon was first 
described by Watson (1881:280-296). 
The zonary placenta of the raccoon is 
similar to that of other carnivorous mam- 
mals. Watson (1881:279) noted: 

“The placenta formed a complete 
ring, but at the centre of its widest 
part, 7.e., opposite the back of the 
foetus, there was a spot similar to 
that figured by Daubenton in the 
placenta of Martes domestica, and 
described by Bischoff in that of Lutra 
vulgaris, Mustela foina and Mustela 
martes, where the substance of the 
placenta was deficient. This defi- 
ciency involved the entire thickness 
of the placenta, so that a probe 
could be passed from the uterine to 
the chorionic surface of the organ 
without injury to its substance.” 
The placenta of Procyon is truly decidu- 
ous in character, as it is in the dog, cat, 
fox, and seal. According to the classifi- 
cation of Mossman (1937:224), the rac- 
coon placenta is endotheliochorial. Pla- 
cental scars in the raccoon were apparent- 
ly first noted by Stuewer (1943b:68), 
who autopsied a female raccoon in May 
and found four placental scars in the 
uterus; he believed they indicated that 
four young had been born. Sanderson 
(1950:399) examined uteri from six cap- 
tive females and concluded that “pla- 
cental scars may be an accurate measure 
of litter size in raccoons.” 

If placental scars are to be useful in 
estimating the reproductive performance 
of a species, several facts about them 
must first be known. Pertinent questions 
are: (1) Is one placental scar formed 
for each implantation site regardless of 
the fate of the developing embryo? (2) 
If the answer to the first question is no, 
then what stages of embryonic develop- 
ment result in the formation of placental 
scars? (3) Is it possible to differentiate 
placental scars formed from embryos that 
go to term from those formed from em- 
bryos that are aborted or resorbed? (4) 


Vol. 31, Art. 2 


How long do the placental scars persist, 
and is the length of time they persist 
affected by the female’s subsequent breed- 
ing history? (5) Are the placental scars 
recognizable at all seasons of the year? 
(6) If the placental scars persist beyond 
a subsequent pregnancy, is it possible to 
recognize scars representing litters from 
different years? Some preliminary infor- 
mation on all of these questions has been 
obtained. 

In only 2 of 27 litters with a total of 
98 embryos in 2 of 20 captive female 
raccoons that we examined did we find 
discrepancies between the number of em- 
bryos observed and the number of pla- 
cental scars identified later. One female 
(No. 2960) had four embryos, estimated 
to be 30 days of age when examined on 
May 26, but five grossly identical pla- 
cental scars when the uterus was re- 
moved 6 months later. The additional 
scar may have represented a litter of one 
from a previous year. If so, the scar was 
overlooked when. this same uterus was 
examined during the fall before the four 
embryos were observed. The extra scar 
may have also represented an additional 
embryo that was aborted or resorbed 
prior to the time the four embryos were 
examined. The second female (No. 
4022) had two live and one dead em- 
bryo when first examined on February 
19. She gave birth to two live young 
29 days later, but when her uterus 
was examined 11 days after parturition, 
there were two grossly identical scars in 
each horn. There were four corpora 
lutea in her left ovary and none in the 
right. Thus, the additional scar observed 
at the second laparotomy was probably 
from an embryo that was aborted or re- 
sorbed prior to the first examination. 

A captive female raccoon (No. 2960) 
had four embryos, estimated to be 30 
days of age, when examined. She was 
given a drug, Malucidin, that caused_ 
either abortion or resorption. The em- 
bryos were gone 13 days later, and the 
sites of attachment were indicated by 
large bumps. When the uterus was re- 
moved 6 months later, five placental scars 
were identified by slight bumps. We 


July, 1973 Sanperson & NaLBanpov: REPRODUCTIVE CycLE oF THE Raccoon 63 


split the uterine horns and identified the 
five placental scars as typical for captive 
females. (Possible differences in placental 
scars of captive and wild animals are dis- 
cussed below.) Thus, in this female, 
one placental scar was formed for each 
of the four embryos even though all four 
embryos were either aborted or resorbed 
at midterm. As has been discussed, the 
fifth scar either persisted from the previ- 
ous year or resulted from an embryo 
resorbed prior to the first examination 
when the four embryos were about 30 
days of age. 

Another captive female raccoon (No. 
3333) had four live embryos and one 
that was being resorbed in her uterus 
on March 21. It was estimated that the 
embryo being resorbed had died 30 days 
after conception. The young were born 
33 days after the initial examination. 
Sixteen days after the birth of the litter 
the placental scar representing the re- 
sorbed embryo was smaller than the 
others, but 47 days after parturition no 
gross difference could be detected among 
the five scars. 

One pregnant captive female raccoon 
(No. 2824) was castrated approximately 
50 days prepartum, but her embryos con- 
tinued to grow for about 20 days before 
they were aborted and resorbed. A 
second captive pregnant female (No. 
2151) was castrated approximately 30 
days prepartum, and her young were 
aborted about 1 week prepartum. Two 
months after abortion or resorption the 
placental scars in these females could 
not be differentiated grossly from those 
formed by normal embryos born at term. 
Female No. 2824 was killed 4.5 months 
after she was castrated. When she was 
killed, only one placental scar was found, 
both before and after the uterus was 
split, even though the exact locations 
of the embryos were known. The one 
scar was dark and broad, and appeared 
to be typical of those formed from young 
born during the current breeding season. 
The scar was formed at the site of one of 
three embryos present 19 days after cas- 
tration. All three of the embryos were 
aborted prior to 26 days after castration. 


The data from the two castrated fe- 
males (No. 2824 and 2151) that lost 
their young and from six intact captive 
females that resorbed or aborted some 
or all of their embryos indicated that 
one placental scar was formed for each 
embryo that existed for approximately 
30 days, whether or not any embryo 
went to term. 

We have made some observations on 
the persistence of placental scars in the 
raccoon (Table 11). Placental scars 
were present, although indistinct, in one 
female (No. 2959) when her uterus was 
removed nearly 19 months after her 
young were born. Scars were visible 12 
and 17 months after parturition in 
another female (No. 1786), but could 
not be seen in her enlarged uterus stimu- 
lated by hormones 14 and 24 months 
after parturition. Her ovaries were re- 
moved approximately 1 year after the 
birth of her young, and, after castration, 
she was treated with estradiol and pro- 
gesterone at various intervals. These 
treatments may have affected the rate of 
disappearance of her scars. A third fe- 
male (No. 2779) had placental scars for 
14 months, but not 23 months, after 


Table 11.— Persistence of placental scars in 
captive raccoons. 


Number of Months Scars 


Raccoon Persisted After Parturition 
Number : 
Minimum Maximum 

19 4.0 
1782 5.0 5c 
1786 17.0° 30.0 
2120 6.0 Stele 
2124 4.5 18.5 
2124 8.0 we 
2125 16.5° 
2151 3.0 AoC 
2184B 15.0 27.0 
2230 6.0 ane 
2779 14.0 23.0 
2959 18.5 aac 
2960 19.0 30.0 
3333 14.0 5 
3350 3.5 

a Not visible macroscopially — approximately 14 
months after parturition — in the uterus stimulated 


by hormones. 

> Not visible macroscopically during the subsequent 
estrus, approximately 10 months after parturition, but 
the same scars were again visible macroscopically 16.5 
months after parturition. 


64 Intino1s NatursL History SurvEY BULLETIN 


parturition, and a fourth female (No. 
2125) retained placental scars nearly 17 
months after parturition. None of these 
females gave birth in the second year. 

There was no macroscopic evidence of 
placental scars from a 1958 litter (Fe- 
male 2124) 18.5 months postpartum, 
but histological examination revealed a 
few scattered pigment granules, and scars 
from a 1959 litter were prominent. Thus, 
the 1958 scars disappeared, for practical 
purposes, prior to 18.5 months after 
parturition, when she had a litter the 
following year. All female raccoons had 
placental scars when examined from 2 
to 10 months after the birth of their 
young. The evidence indicated that if 
a female failed to give birth to a litter 
in the next year, placental scars persisted 
for approximately 19 months in captives, 
but not as long as 24 months. If a cap- 
tive gave birth to a litter the next year, 
scars from the first litter persisted for 10 
or more months but not as long as 19 
months. 

One captive raccoon became pregnant 
at the second ovulation during one sea- 
son. The single embryo, in the process 
ot being resorbed when it was first ob- 
served, was estimated to be 20 days old. 
Twenty-one days later the site of placen- 
tal attachment was readily identified as 
a bump 8 X 7 mm in size; 61 days after 
the initial observation no trace of the 
scar could be seen. Thus, this scar dis- 
appeared between 21 and 61 days after 
the resorbing, 20-day embryo was ob- 
served. The absence of living embryos 
in this captive may have been an im- 
portant factor in the rapid disappear- 
ance of the scar. 

The variability in the length of time 
that placental scars were visible in the 
raccoon after parturition is shown in 
Table 11. In two females scars were 
not grossly visible in their stimulated 
uteri during or near estrous cycles of 
the ensuing years, because their enlarged 
uteri caused a diffusion of the pigment 
granules of the scars, making them in- 
visible. Scars in these females were 
again visible macroscopically when the 
uteri regressed. 


Vol. 31, Art. 2 


Scars from a litter born in May 1958 — 
(discussed above) could not be seen (No. — 
2124, Table 11) 18.5 months later (De- 
cember 1959) even though their exact 
locations were known and the uterus was 
removed and split. After we sectioned © 
the site of one scar, we were able to 
identify a few scattered pigment granules _ 
in the endometrium. Two scars from a — 
litter born in April 1959 were easily — 
identified macroscopically in this same ~ 
uterus 8 months (December 1959) after — 
the birth. In a second female (No. 2959, 
Table 11) scars from young born in May 
were not visible with translucent light 
after the uterus, which was stimulated, — 
was removed 18.5 months later. 
scars from this litter were located and © 
were identified by the slight bumps vis- © 
ible at the placental sites. After the © 
uterus was opened, all four scars were ~ 
visible as pale, brownish areas, but they ~ 
might have been overlooked had not ~ 
their exact locations been known. When 
one of these scars was examined his-_ 
tologically, moderate numbers of pigment 
granules were seen in clumps and scat- 
tered in the endometrium and in the ad-— 
jacent myometrium. | 

The distribution of pigment granules — 
in the uteri of two wild females was stud- — 
ied in histological sections. Each of © 
these females had four scars at autopsy. — 
Pigment granules in the uterus of one fe- 
male were somewhat scattered but seem- _ 
ed to concentrate in a ring deep in the — 
endometrium near the myometrium. — 
Many pigment granules were scattered 
throughout the endometrium of the 
uterus of the other female. 

Pale placental scars were often diffi- 
cult to see in situ in a live animal, and” 
early in the study some scars may have 
been overlooked. We believe that, after 
we became experienced in looking for 
scars, no visible placental scar was over-— 
looked, but they could not be seen in- 
pregnant females and females at or near 
estrus. When the scars had practically 
disappeared, they could be observed only 
by splitting the uterus. Thus, these pale 
scars would be overlooked when examin- 
ing live females by laparotomy. If the 


All four — 


July, 1973 SanpEerson & NatBanpov: ReEpRopucTIVE CycLe oF THE Raccoon’ 65 


uterus of a live female was stimulated, 
many of the placental sites could best be 
identified by slight, opaque bumps rather 
than by the pigmentation. Identification 
of the location of scars by the presence of 
bumps was possible for several weeks 
after parturition, when the uterus was 
still stimulated, as well as in the stimu- 
lated uterus at or near estrus. After the 
uterus regressed, scars were usually read- 
ily visible as bumps or could be identi- 
fied by using translucent light to observe 
the pigmented areas. The pigmented 
areas could also be located when the 
uterus was opened or by histological ex- 
amination. 

Placental scars seem to persist longer 
in wild raccoons than they do in captives. 
The placental scars of captives that we 
examined from October through January 
after the births of their litters were gen- 
erally pale brown, small, and _ slightly 
opaque. A majority of the wild females 
examined during these same months had 
larger, more opaque scars, often black. 
Many (55.2 percent in 1959 and 41.0 
percent in 1960) uteri of wild, parous 
females had more than one group of 
scars, which differed in size and density 
(Fig. 11 ). Presumably these scars were 
from different years; however, some 
might have been from different litters 
born in 1 year. There was no evidence 
that as many as 40 or 50 percent of the 
wild females gave birth to second litters 
during a single season. Thus, placental 
scars probably persist for 20 months or 


longer in many — perhaps in all — wild 
females. In the few wild females with 
three groups of scars, the first group may 
have persisted for as long as 32 months. 

The placental scars of raccoons are 
useful for estimating litter size and rate 
of productivity. However, these scars 
must be used with caution, and care must 
be taken to separate properly the groups 
of scars. We do not know for certain 
the significance of multiple groups of 
scars. We can say with reasonable con- 
fidence that each embryo that reaches 
1 month of age is represented by one 
scar for 10 or more months. Scars in wild 
females with only one group of scars 
probably reflect implantation rates for 
the preceding breeding season. Most 
single groups of placental scars occur in 
females that have mated successfully only 
once. 


MORPHOLOGY OF THE 
REPRODUCTIVE TRACTS 


Males 

The duct system and accessory glands 
in the reproductive system of the male 
raccoon (Fig. 12) are similar to those 
found in the dog, as described and shown 
by Nalbandov (1958: 42-44). Seminal 
vesicles are lacking, as they are in the 
dog, fox (Vulpes fulva) , and wolf (Canis 
lupus). The Cowper’s glands (bulbo- 
urethral glands) are also absent. The 
walls of the vasa deferentia thicken prior 
to entering the prostate and form the 
ampullae. The ampullae and the urethra 


Fig. 11.—Raccoon uterus (X 0.75) split to show two groups of placental scars. 


was killed on January 23. 
visible in the fresh specimen. 
by the light stippling (arrows). 


This female 


Two light scars were only barely visible in the photograph but were readily 
Their locations ard densities relative to the three dark scars are indicated 


66 Inuino1s NaTurAL History SurvEY BULLETIN 


Ampulla 


Prostate 


Vas Deferens 


Vol. 31, Art. 2 


Fig. 12.—Schematic drawing (side view) of the reproductive system (X 0.85) of an adult male 


raccoon, 


unite inside the prostate to form a com- 
mon duct. The many compartments of 
the prostate gland open into this duct 
system. 

The os penis or os baculum (bone of 
the penis) is well developed in the rac- 
coon. Its stage of development has been 
used to separate males into two age 
groups (Sanderson 1950: 395-396; 
1961a: 11-14). The os baculum was once 
used by tailors as a ripping tool for taking 
out basting threads (Jaeger 1947: 297). 

We found several raccoon bacula that 
had been broken and then healed. Sand- 
erson (1950: Plate 11) showed a photo- 
graph of some of these bones. Our data 
from wild males shed some light on pos- 
sible causes for these broken bones. Dur- 
ing four hunting and trapping seasons 
in Illinois (1957-1958 through 1960- 
1961), 7,233 bacula from juvenile rac- 
coons were examined. Forty-three (0.6 
percent) of these had been broken but 
were healed or healing, and 238 (3.3 per- 
cent) were freshly broken. At the same 
time, 4,152 bacula from adults were ex- 
amined. Eighty-six (2.1 percent) of 
these had been broken but were healed, 
whereas 41 (1.0 percent) were freshly 
broken. 

These data indicate that most of the 
breaks in the os baculum of the raccoon 
occur in juveniles. The bacula of juven- 
iles are much softer and more easily 
broken than are those of adults. Hunters 
often shake a raccoon out of a tree and 


let their dogs fight it. Fighting with 
dogs could account for the freshly broken 
bones found in both adults and juveniles, 
and the more durable bones of adults 
would explain the smaller percentage of 
freshly broken bacula found in older 
raccoons. 


Females 


The raccoon uterus (Fig. 13) is some- 
what intermediate between the bicornuate 
uterus found in the pig and insectivores, 
and the bipartite uterus found in the cat 
and dog. There is a single cervix and 
the horns are distinct, but after the horns 
join externally to form the single, small 
uterine body, the uterine lumina remain 
separate — even though this separation 
is not apparent from the outside — to a 
point near the cervix. 

Llewellyn & Enders (1954b: 439) re- 
moved one ovary, ovarian capsule, ovi- 
duct, and proximal end of the uterine 
horn in each of two raccoons. After 
closing the cut ends of the uteri with 
sutures, they released the females. When 
retrapped the next year, each female was 
carrying three embryos, two each in the 
normal horns and one each in the ovar- 
iectomized horns. Thus, even though 
the internal separation of the uterus ex- 
tends nearly to the cervix, ova can pass 
from one uterine horn to the other. In 
our study some indirect evidence of 
transuterine migration of ova was noted. 
In a few cases more embryos were found 


° 
July, 1973 Sanperson & NaLBaNDov: 


Fimbria 


Bursa ovarii 


Septum 


Oviduct 
Bladder 
pk Colon 23 = 


Cervix 


Uterine Body \ 


Placental scar 


REPRODUCTIVE CYCLE OF THE Raccoon’ 67 


Uterine horn oo” 
Oviduct Ovary 


Fig. 13.—Schematic drawing (ventral view) of the reproductive tract (X 0.5) of a parous female 


raccoon. 


in a uterine horn than there were corpora 
lutea in the corresponding ovary, but 
the total number of corpora lutea present 
in both ovaries was usually the same as 
the number of embryos or placental scars 
present in both uterine horns. 

The ovary in the raccoon, ovoid in 
shape, is completely surrounded by the 
bursa ovarii (Fig. 13). This sac is in- 
tact except for a small slit on one side, 
not large enough to permit passage of the 
ovary as in the mink (Mustela vison), 
dog, and fox. One of our captive fe- 
males had a congenital deficiency of the 
bursa that was large enough to permit 
passage of the right ovary. This opening 
was slightly dorsal to the normal slit in 
the bursa but was not connected with it. 
The left ovarian bursa was normal. This 
captive was the only such animal among 
several hundred examined. Watson 
(1881: 273-274) observed one raccoon 
and reported that the ovary was destitute 
of any peritoneal pouch or pavilion such 
as formed an almost complete sac in 
many animals. 

The fimbria is extensive, and in the 
estrous female the edge of the fimbria 
is bright red and protrudes through the 


slit in the capsule. This bit of fimbria 
grossly resembles the gills of a fish. The 
fimbria joins with the end of the oviduct. 
The oviduct is highly convoluted and 
makes an almost complete circle around 
the ovary before entering the uterus (Fig. 
13). 

Two, three, and sometimes four ova 
were observed in a single follicle. When 
an ovary contained one follicle with 
multiple ova, several other follicles with 
multiple ova were usually present. 

Approximately 25 female raccoons 
from Iowa and approximately 25 from 
Illinois were examined for the presence 
of the os clitoridis. A bone — 11 mm in 
length — was found in only one clitoris. 
Rinker (1944: 91) found four ossa cli- 
toridae in four female raccoons examined 
in Kansas, but found no bones in the 
clitoria of four other females from a 
“distant locality,’ apparently in Kansas. 
Burt (1960: 8) used Rinker’s observa- 
tion as the basis for stating that the os 
clitoridis is present in the raccoon. 
Sanderson (1950: 398) found only one 
os clitoridis among 100 female raccoons 
in Missouri. Because only a small per- 
centage of females examined from Mis- 


68 


souri, Iowa, and Illinois had ossa clitor- 
idae, there may be geographic variation 
in the presence of this bone. Its presence 
is not of general occurrence in raccoons 
in all localities. 


EFFECTS OF CASTRATION 
Males 


Some effects of castration on the de- 
velopment of the os baculum in the rac- 
coon have been discussed by Sanderson 
(1961a: 13-14). The information in 
that report, with additional observations, 
is presented here. The lack of sex hor- 
mones in males was reflected by the 
much shorter and thinner bacula in 
castrated animals in comparison with 
bacula from intact animals of similar ages 
(Sanderson 196la: Fig. 4). The lack 
of sex hormones became apparent at 8- 
11° months of age in castrated males. 
Sanderson (1950: 396) showed that in 
intact males the penis normally became 
extrusible at about 10 months of age, but 
a castrate male (Sanderson 196la: Fig. 
6, No. 59) had a nonextrusible penis and 
a small baculum at 22 months of age. 
This baculum was only slightly longer 
and heavier than one from a castrate 
raccoon only 10 months of age (Sander- 
son 196la: Fig- 5, No. 209), but both 
were much shorter and thinner than were 
the bacula from intact males 18-23 
months of age (Sanderson 196la: Fig. 
6). The baculum from the castrated rac- 
coon 22 months of age was dense like an 
adult bone and not spongy at the base 
as were bacula of similar size from rac- 
coons 12 months of age and younger. 

Thus, we concluded that the level of 
sex hormones affected the enlargement 
of the preputial orifice and maturation of 
the penis bone but had little or no effect 
on the development of the baculum prior 
to 7 months of age. 

Castration in males also apparently 
caused a slight delay in the closure of 
the epiphyseal cartilage in the radius 
and ulna, but because most of the castrat- 
ed males in this study died of disease at 
early ages, not enough information was 
available to demonstrate this relation- 
ship conclusively. 


Intrinois NaTurAL History SurRvEY BULLETIN 


Vol. 31, Art. 2 


Epiphyseal plates were classified as 
closed (without cartilage), thin (inter- 
mediate condition), or broad (with a 
thick plate of cartilage) (Sanderson 
1961a: 7). One castrated male had 
broad epiphyses at 17 months of age and 
thin epiphyses at 20 months of age. His 
epiphyses were still thin when he died at 
22 months of age. When examined, 35 
intact males with broad epiphyses were 
15 months of age or less, whereas 14 of 


17 males (82 percent) with thin epi-. 


physes were 13-19 months of age. Epi- 
physeal plates in 11 of 13 intact males 
closed between 16 and 21 months of age 
(Sanderson 196la: 16). The effects 
of castration on epiphyseal closure merit 
further study. 


Females 


The time of closure of epiphyses in fe- 
males was much like that in males, but 
the greater variation in the upper ages 
of females with thin epiphyses indicated 
that epiphyseal closure was delayed in 
some females or occurred later in some 
than in others. 

One factor that perhaps influences age 
at epiphyseal closure is the level of cir- 
culating hormones. Two females were 
castrated to study the effects of the ab- 
sence of ovarian hormones on epiphy- 
seal closure. One female, born in the 
wild, was castrated at an estimated age 
of 4 months, and one, born in captivity, 
was 3 months old when castrated. The 
first had broad epiphyses at 14 months of 
age, thin epiphyses at 20 months of age, 
and thin epiphyses when she died at 23 
months of age. The second had broad 
epiphyses at 22 months of age, thin epi- 
physes_at 25 months, and nearly closed 
epiphyses at 27 months of age. Epiphy- 
seal development and closure in these 
two castrated females were delayed in 
comparison with the rate of development 
and closure found in the average intact 
female. Four additional females were 
castrated at estimated ages ranging from 
13 to 24 months. Our observations sug- 
gested that the removal of the ovaries, 
even after raccoons had reached sexual 
maturity but before the epiphyses closed, 
delayed the rate of epiphyseal closure. 


r 


July, 1973 Sanperson & NaLBanpov: REpRopuUCTIVE CycLE OF THE Raccoon 69 


Major factors that may have contrib- 
uted to the variations we observed in 
age at epiphyseal closure in female rac- 
coons were (1) age at first mating, (2) 
hormone secretion level, and (3) quality 
and quantity of nutrition. Factors influ- 
encing age at epiphyseal closure should 
be studied further because the available 
data are somewhat contradictory. 

Thus, our data and those of Sanderson 
(1961a: 10-11) suggest that epiphyseal 
closure in the castrated female raccoon 
is delayed in comparison with that found 
in the average intact female but falls 
within the limits of variability for intact 
females. 

In mammals castration after implanta- 
tion and during the first third or first 
half of pregnancy usually leads to abor- 
tion or resorption of the fetuses (Nalban- 
dov 1958:221). In some mammals the 
ovaries are required throughout gestation, 
but other mammals do not lose their 
young after castration, once the crucial 
period is past. 

Two pregnant females were castrated 
to learn whether the raccoon is a species 
in which pregnancy is maintained after 
castration. To establish limits after which 
castration is tolerated, these two animals 
were castrated approximately 11 and 38 
days after conception, respectively. (It 
had been established earlier that perform- 
ing laparotomies on pregnant raccoons 
did not interfere with pregnancy.) 

The first of these two pregnant female 
raccoons that we castrated was born in 
1959 and reared as a pet. She gave birth 
to a litter in 1960. On February 5, 1961 
she forcibly repelled the approaches of 
her mate. Twenty-five days later she had 
three embryos in her left uterine horn 
and two in the right but only four cor- 
pora lutea. Each uterine swelling was 
10 mm in diameter. We estimated the 
embryos to be 11 days old, suggesting 
that mating had occurred about February 
17. Both ovaries were removed on March 
2, and 19 days later two embryos in the 
left uterine horn were being resorbed. 
These two swellings were almost as large 
as the other three, but the surfaces were 
collapsed and flaccid, not turgid like 
those of a normal swelling. The embryos 


were still present at both sites. The 
remaining three embryos, 45 X 20 mm, 
looked almost normal, except that the 
swellings appeared less round and turgid 
than normal swellings are. We could not 
discern whether the embryos were alive 
or dead. We estimated that, if they were 
alive, they would be born in 25 days 
(Fig. 7). Thus, these embryos had a 
normal rate of growth for 19 days after 
the castration of the female. No embryo 
was found 26 days after castration. From 
gross appearances we concluded that the 
last three embryos were aborted and the 
first two were resorbed. 

Twenty-eight days after this female 
raccoon was castrated, her mate was re- 
turned to her cage. The next day, only 
3-10 days after her young were aborted, 
this pair was observed in copulation. 
This activity suggests the possibility that 
postpartum heats, which occur in several 
species such as the sow and mare, may 
not be dependent upon the presence of 
the ovaries. 

This female escaped 3 months after 
she was castrated and was taken in a steel 
trap 52 days later. After she was killed, 
it was discovered that she was lactating 
profusely. When the mammary gland was 
sliced with a scalpel, the entire cut area 
immediately filled with milk. She was 
lactating more than 4.5 months after her 
ovaries had been removed and 4 months 
after her young had been resorbed and 
aborted. However, the second pregnant 
female that was castrated showed no 
indication of lactation 5 months after re- 
moval of her ovaries. 

No traces of ovarian tissue were found 
during the autopsy performed on the 
first of these females. The uterus, measur- 
ing 7 X 4 mm, was turgid and appeared 
similar to uteri of animals at estrus, but 
sectioning showed the endometrium to 
be devoid of even traces of glands. Other 
female raccoons, months after being cas- 
trated, had thick epithelia lining their 
vaginas (Fig. 9A), suggesting the possi- 
bility of an extraovarian source of estro- 
gen in the castrated female. 

The second of the pregnant raccoons 
that was castrated was placed in captivity 
in 1958 when she was about 2 months 


70 Intino1is NaturAL History Survey BULLETIN 


old. She had two embryos in each uterine 
horn on February 28, 1961, when both 
ovaries were removed (approximately 38 
days after conception and 25 days before 
parturition), but the left ovary had three 
corpora lutea and the right ovary only 
one corpus luteum. Nineteen days after 
castration and 6 days prior to expected 
parturition, one dead embryo weighing 
43 grams was found in her nest box. The 
average birth weight of eight newly born 
raccoons that we weighed was 61.8 grams. 
The embryo was well developed but did 
not have much hair, and its hair was 
shorter than in most young at birth. 

Twenty-one days after castration the 
uterus contained enlarged areas where the 
young had been attached. One of these 
sites was opened and examined. Detritus 
was present, but there was no other evi- 
dence of resorption, which was occasion- 
ally seen in both wild and captive fe- 
males. Thus, all four embryos were prob- 
ably aborted about 1 week prepartum. 

Although the two females were cas- 
trated at different stages of pregnancy, 
the embryos apparently persisted for 
about the same length of time in each— 
19 days after castration—33 days short 
of term for the first female and approxi- 
mately 7 days short of term for the 
second. 


EFFECTS OF EXOGENOUS 
HORMONES 


Males 


Two male raccoons were studied to 
learn whether injections of androgen 
would initiate or prolong spermatogenesis 
during the male’s period of summer steril- 
ity. The first animal chosen was an 
adult male at least 20 months old when 
he was captured. In August (during the 
period of sexual inactivity) the left testis 
was removed, weighed (1.6 grams), and 
preserved for histological study. The 
average weight of one testis from an adult 
in August was 2.6 grams (Table 1). 
Sperm could not be found in the epididy- 
mis, but spermatogenesis was occurring 
in a few seminiferous tubules. This male 
was given six testosterone doses of 30 


Vol. 31, Art. 2 


mg each subcutaneously over a period of 
18 days. He was killed 21 days after the 
removal of his left testis and the first 
injection of testosterone. The right testis 
weighed 1.6 grams, and no sperm were 
present in either the seminiferous tubules 
or epididymis. 

Histological comparison of the two 
testes and the epididymides showed 
slight changes that we attributed to the 
testosterone injections. Both before and 
after the hormone treatment most sper-, 
matogenic cells were approximately 8 mi- 
crons, and the nuclei 3 microns, in di- 
ameter. However, after testosterone in- 
jections a few of the cells were as large 
as 11 microns in diameter. The lumina 
of the seminiferous tubules remained 
about the same size after the treatment as 
they were before, but after the injections 
the cells of the seminiferous tubules were 
more scattered than they were before 
treatment. Sperm were present in the 
seminiferous tubules prior to treatment, 
but not afterwards. Sperm could not be 
found in either epididymis, one of which 
was removed and examined before and 
the other after the homone treatment. 
The epithelial lining of the tubules was 
46 microns tall prior to treatment and 
30 microns after treatment (each height 
is an average of five measurements) , indi- 
cating degenerative changes, perhaps 
caused by the hormone. The average 
outside diameter of the tubules was 140 
microns prior to treatment and 65 mi- 
crons afterwards. 

A second male raccoon, captured when 
approximately 3 months of age, was 
reared as a pet. He was approximately 
16 months old when his left testis, weigh- 
ing 3.3 grams, was removed in August. 
The epididymis contained many motile 
sperm. 

He was treated with four doses of 12 
mg cach of testosterone over a 13-day— 
period. He was killed 15 days after the 
removal of the left testis and the first 
injection of the hormone. At that time his 
right testis weighed 2.5 grams, and many 
motile sperm were in the epididymis. His- 
tological examination revealed few chang- 
es in the cells of the seminiferous tubules. 


July, 1973 SanpERson & NaLBanpov: ReEpRopucTIVE Cycle oF THE Raccoon 71 


A few sperm were present in the semini- 
ferous tubules both before and after treat- 
ment. After treatment sperm were not 
found in a section of the epididymis, but 
a few were observed in a drop of fluid 
collected from the tail of the epididymis. 


Females 


Ovulation can be induced during anes- 
trus im several species of domestic and 
laboratory animals by the injection of 
gonadotrophic hormones. Hammond 
(1952:218) used pregnant mare’s serum 
(PMS) as a follicle-stimulating agent and 
chorionic gonadotropin to cause ovula- 
tion in ranch mink. 

We made several attempts, using 19 
individuals, to cause the growth and de- 
velopment of follicles and to cause ovu- 
lation in the raccoon by injecting hor- 
mones. Only four individuals ovulated, 
and three of these cases involved the use 
of PMS (Table 12). Only the four cases 
in which ovulation occurred are discussed. 

In one series of experiments various 
dosages of follicle-stimulating hormone 
(FSH) given subcutaneously were fol- 
lowed by luteinizing hormone (LH) 
given intravenously. Later FSH and LH 
were mixed and given subcutaneously, 
followed by intravenous injection of LH. 
With one exception all attempts using 
FSH and LH were unsuccessful in caus- 
ing ovulation. In some cases normal-ap- 
pearing follicles were numerous in the 
ovaries after injections of FSH and mix- 
tures of FSH and LH, but attempts with 
LH and with a mixture of LH and FSH 
to cause the follicles to ovulate were un- 
successful. The ovaries generally were 
overstimulated; that is, they were larger 
than normal and contained more follicles 
than normal. 

The successful ovulation that did not 
involve injections of PMS occurred in a 
female raccoon (the first female in Table 
12) approximately 44 months old, weigh- 
ing 6.7 kg. Each ovary was 11 X 6 mm, 
with no follicles or corpora lutea ap- 
proximately 2 months prior to the breed- 
ing season. A section of her uterus was 
removed when the ovaries were measured 
(approximately 2 months prior to the 


breeding season) so that we could study 
the pigment granules. The next day sub- 
cutaneous injections of a mixture of 10 
Armour units (AU) each of FSH and 
LH were begun. These injections were 
given for 10 days, and on the 12th day 
a mixture of 80 units each of FSH and 
LH was injected intravenously. At that 
time each ovary was 12 X 8 mm and con- 
tained 10-20 clear follicles, each about 
1 mm in diameter. On the 13th day 100 
units each of FSH and LH were injected 
intraperitoneally as a mixture. On the 
16th day the left ovary was 18 X 9 mm 
and contained approximately 20 follicles, 
each about 2 mm in diameter, but ovu- 
lation had not occurred. The left ovary 
weighed 760 mg, compared with an aver- 
age weight of about 137 mg for one ovary 
of parous or pregnant females auring the 
mating season (Table 5). When the rac- 
coon was killed 45 days after the first 
injection, her right ovary weighed 290 mg 
and contained 11 corpora lutea. 

In 2 second series of experiments PMS 
was injected into eight females in at- 
tempts to cause the development of folli- 
cles and to cause ovulation. Three of 
these attempts were successful. The first 
female was approximately 2 years of age 
and had been in captivity for more than 
a year when hormone treatments were 
begun. We injected 100 international 
units (IU) of PMS subcutaneously each 
day for 12 days and 500 IU each on the 
13th and 16th days. Thirteen days later, 
28 days after the treatment was begun, 
the uterus and both ovaries were removed. 
The contents of the oviducts and uterine 
horns were flushed out, but no ova or 
blastocysts were found. Each ovary con- 
tained approximately 30 corpora lutea. 
Even though no ovum was recovered, 
the abnormally large number of corpora 
lutea containing no ova indicates that this 
female probably ovulated. The secretory 
material found in the lumina of the uter- 
ine glands indicated that progesterone 
had probably been secreted. 

The second female was captured when 
she was at least 18 months of age; how- 
ever, she was not injected with hormones 
until she was about 53 months old. Sub- 


Vol. 31, Art. 2 


Iutinois NaturaL History SuRVEY BULLETIN 


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July, 1973 SanpErson & NaLBANDov: REPRODUCTIVE CYCLE OF THE Raccoon 75 


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pea? HE) aviv REDS ESSAI NI LY Sl A VA ARE AN OS 
cutaneous injections of PMS at the rate 
of 100 IU daily for 12 days were begun 
in September. Many large follicles were 
found on the 13th day of treatment. This 
female was given 550 IU of PMS intra- 
venously on the 13th day. Seventy-two 
hours later many blut punkte were ob- 
served in each ovary. The left ovary 
(826 mg) anda piece of the uterus were 
removed. Histological examination re- 
vealed 26 corpora lutea in early stages. 
Most had blood in the lumina and ap- 
peared to be freshly ovulated. The uterus 
showed a fairly typical effect of estrogen, 
and no material was present in the uterine 
glands, indicating the near absence of 
progesterone. Eleven days later (26 days 
after the first injection) the female was 
killed and the right ovary was removed 
(2,147 mg). There were 29 early-stage 
corpora lutea, most of them packed with 
luteal cells, but lumina were present in 
2-4 corpora. The cytoplasm and nuclei 
of these luteal cells were more darkly 
stained and the nuclei were smaller than 
usual. The intracellular space exceeded 
the norm. Secretory material was present 
in the uterine glands. 

The third female was about 22 months 
old when caught, but was 53 months of 
age when these experiments were begun. 
Subcutaneous injections of PMS were 
begun 2 months before the breeding 
season, at the rate of 50 IU per day, 
and were continued for a total of 12 
injections. On the 13th day 200 IU were 
injected intravenously. At that time each 
Ovary measured 12 X 6 mm and con- 
tained approximately nine follicles, each 
about 2 mm in diameter. Two days later 
the ovaries and follicles had not changed 
in size, but one follicle was hemorrhagic 
and one had a thin red line across the 
surface at its highest point. Twenty-four 
hours later when the ovaries were exam- 
ined, blood oozed from most or all of 
11 or 12 follicles in each. There were 
tiny holes in the highest points of most, 
and perhaps in all, of them. The ovu- 
lated follicles were partly hollow and 
partly filled with fluid and stringy ma- 
terial. The left ovary, measuring 10 X 6 


ee ae 


MRA RA RN NR FE ERR IN oe 
mm, was removed (242 mg). When ex- 
amined histologically, it was found to 
contain six or more blood-filled, early- 
stage corpora lutea. Among 16 wild rac- 
coons the average number of corpora 
lutea per ovary, determined by histologi- 
cal examination, was 2.1. Thus, in this 
most nearly normal ovulation induced by 
exogenous hormones, the ovaries were 
somewhat less than twice normal weight, 
but the ovulation rate was approximately 
5.7 times normal. 

The female just discussed weighed 
5.35 kg and received a total of 800 IU 
of PMS, a dosage of about 150 IU per 
kg, a rate similar to that used success- 
fully to cause ovulation in ranch mink 
(Hammond 1952:219). 

In a third series of experiments four 
different hormones were used on four 
sexually immature female raccoons from 
2 to 4 months old (the last four animals 
in Table 12) in an attempt to learn how 
immature ovaries respond to hormones 
and to study differential responses to the 
several hormones. The injection of hu- 
man menopausal gonadotropin (HMG- 
J5, largely FSH) subcutaneously twice a 
day for 7 days immediately after 2 days 
of single injections resulted in little stimu- 
lation of either the ovary or the uterus 
in one immature female. In the second 
young raccoon 50 international units 
(IU) of PMS daily for 8 days, followed 
by 100 IU and 250 IU on the 9th and 
10th days, respectively, resulted in a 
slightly more stimulated uterus than did 
the HMG-J5 injected into the animal 
just discussed. In the third animal in 
this age group 10 Armour units of LH in- 
jected subcutaneously daily for 9 days, 
followed by 25 units on the 10th day, 
resulted in larger follicles than did either 
of the two previous treatments. 

In the raccoon that received only LH 
this hormone caused more development of 
the follicles than did either PMS or 
HMG-J5 in the other young females. 
PMS and HMG-J5 contain both FSH 
and LH and might be expected to cause 
greater stimulation than LH alone. The 
ovaries stimulated by HMG-J5 contained 


78 Inurno1is NaTuRAL History SuRVEY BULLETIN 


more interstitial tissue than did the ovar- 
ies of the females that received the other 
hormones, and the ovaries of the raccoon 
that received PMS had less interstitial 
tissue than those of the female that re- 
ceived LH. The uterus of the female 
injected with HMG-J5 was somewhat 
less stimulated (endometrium 650 mi- 
crons) than that (endometrium 820 mi- 
crons) of the female that was given PMS 
although the differences in these uteri 
were slight. The uterus of the female 
that received LH was more stimulated 
(endometrium 1,275 microns) than was 
either of the other two. 

On the basis of the information ob- 
tained from’ our experiments with four 
female raccoons, it appears that 35-50 
IU of PMS given subcutaneously each 
day for 12 days caused the development of 
follicles at any time of year in adult 
females. A dose of 200 IU given on the 
13th day might be expected to cause 
ovulation 48-60 hours later. 


UTERINE MILK 


Uteri, and ovaries containing corpora 
lutea, were sectioned from 18 raccoons 
that had not been treated with hormones. 
In 17 of the 18 secretory material 
(uterine milk) was present in the lumina 
of most, but not all, of the uterine glands 
although it may have been present in all 
18 uteri but overlooked in some of the 
sections. 

Histological sections of ovaries con- 
taining no corpora lutea and the corre- 
sponding uteri were examined from 89 
raccoons collected throughout the year. 
February and March were each repre- 
sented by a single animal, but each other 
month was represented by three or more 
animals. The uterine sections from these 
animals, with two exceptions, contained 
no secretory material in the endometrial 
glands. Small amounts of secretory ma- 
terial were present in the uterine glands 
of one nulliparous adult killed in Septem- 
ber and in another, approximately 7 
months of age, collected in November. 
Secretory material was not abundant in 
either one, but was definitely present. 


Vol. 31, Art. 2 


These data indicate that, in the rac- 
coon, secretory material (presumably 
uterine milk) is present when corpora 
lutea are present. In one female, judged 
to have been only 10 days prepartum, 
secretory material was present. 

Progesterone alone or in combination 
with estrogen was probably responsible 
for the secretion of uterine milk (Table 
13). Progesterone alone was given for 
an insufficient length of time to determine 
whether it alone can cause the uterine 
glands to secrete. Two castrated females 
(No. 1297 and 1786) received a combi- 
nation af progesterone and estrogen for 
several days, and the endometrial glands 
of both contained uterine milk (Table 
13). Any combination of gonadotrophic 
hormones that resulted in the formation 
of corpora lutea caused secretion by the 
uterine glands. Five treatments of 2.5 
mg each of estradiol over periods of 10 
and 20 days, respectively, did not cause 
secretion by the uterine glands in one 
castrated female. One intact female (No. 
2184B) received five daily injections of 
20 units each of FSH, followed on the 
6th day by 50 units of LH. Three days 
later, when she was killed, each ovary 
contained approximately 20-30 follicles 
measuring up to 750 X 1,250 microns, 
but no corpora lutea. The lumina of a 
few endometrial glands contained small 
bits of secretory material. Several hor- 
mones, including various combinations 
of FSH, LH, CGH, PMS, and HMG-J5, 
were given to intact females. Except pos- 
sibly in the female just discussed, none 
of these hormones caused the uterine 
glands to secrete except indirectly by 
causing the formation of corpora lutea. 


Methods. described by Pearse (1960: 
265-271) and by Lillie (1954:274—299) 
were used in an attempt to demonstrate 
the nature of the secretory material. 
In no case did digestion with either pty- 
alin or diastase remove the secretory ma- 
terial from the endometrial glands. This 
finding was taken as evidence that it 
was not glycogen. According to the in- 
formation on the identification of carbo- 
hydrate-containing materials given by 


July, 19/9 SOANDERSON & INALBANDOV. INEPRODUCTIVE UYCLE OF THE KACCOON /Y 


Table 13.—Presence of secretory material in the uterine glands of captive raccoons as related to 


injections of exogenous hormones. 


Estimated 
Raccoon = 
Ri caber Age in Hormone 
Months 
1292 24 PMS* 
1297 14 PMS 
22 Progesterone, 
estrogen 
1298 53 PMS 
1782 14 Estradiol 
Estradiol 
1786 24 Progesterone, 
estradiol 
2184B 38 FSH‘, LH® 
2276 14 FSH, LH, CGH* 
2525 19 FSH, LH, PMS 
2805 3 LH 


Number of 
Days After Corpora Uterine 
First Lutea Milk 

Treatment 
28 ae FP 
49 _ _ 
70 Castrated ae 
15 atpe — 
27 + + 
10 Castrated = 
20 Castrated = 
28 Castrated AF 

8 — T* 

14 = T 
21 + T 
19 — — 
12 — _ 


® Pregnant mare’s serum. 


» The plus symbol indicates the presence of corpora lutea or uterine milk, and the minus symbol indicates 


their absence. 
¢ Early stage. 
4 Follicle-stimulating hormone. 
© Luteinizing hormone. 
tf T=traces. 
® Chorionic gonadotropin hormone. 


Pearse (1960:236—237), it was either a 


mucoprotein or a glycoprotein. 


SUMMARY 


1.—The testes of raccoons in Illinois 
grew at a uniform rate from birth until 
about 10 months of age; at that time the 
average weight of one testis was 5.6 
grams. Most male raccoons reached sex- 
ual maturity as yearlings, but juvenile 
males became sexually potent 3-4 months 
later in the year than did adult males. 
Seasonal variations occurred in testis 
weights; the average weights were min- 
imal in June, July, and August, and were 
highest in December. The average max- 
imum weight of one testis was 2.8 times 
the average minimum. There was a 
positive correlation between testis weight 
and the presence of sperm in the epididy- 
mis, but the weight of the testis did not 
infallibly indicate whether sperm was 
present in the epididymis. In a large 
group of raccoons sperm may be found 
in some animals at any given time, but 
individual males had periods averaging 


3-4 months when they were incapable 
of breeding. 

2.—Ovaries of raccoons showed a near- 
ly steady rate of growth from birth in 
April through the following November. 
The heaviest normal ovaries found were 
in juveniles during November, approx- 
imately 3 months prior to the peak of 
the breeding season. The ovaries of 
juveniles declined in weight from Novem- 
ber through January, and _ perhaps 
through March. Seasonal weights of 
ovaries in parous raccoons followed a 
pattern similar to that found in the 
gonads of adult males. The minimum 
average weight was reached in July, with 
a slow but consistent increase in weight 
occurring from then until November. 
The weights of ovaries of parous raccoons 
declined from November to December 
but increased during January and reach- 
ed their peak average in April, when they 
were slightly heavier than they were in 
November. The average peak weight 
of ovaries of adults in April was slightly 
more than 1.6 times their average weight 
in July. 

3.—The mean birth date for 20 litters 


80 Intinois NaturAL History SurvEY BULLETIN 


conceived in the wild was April 18 
(range, March 9-June 24) and for 11 
litters conceived and born in captivity it 
was April 24 (range, March 16—-June 3). 

4.—The measurement of the largest 
external uterine swelling enabled us to 
estimate birth dates with a maximum 
error of 4 days. 

5.—The sex ratios of young raccoons 
less than 2 months of age and of embryos 
and young at birth were not significantly 
different from 50:50, but there were 
more males among the young less than 
2 months old than among the other 
group, possibly indicating some differen- 
tial mortality of females between birth 
and 2 months of age. 

6.—Yearling females either bred when 
adults bred or did not breed until they 
were almost 2 years of age. If female 
raccoons ovulated but did not become 
pregnant, if they aborted or resorbed 
their young, or if they lost their young 
at or near birth, they sometimes ovulated 
a second time in one season. The inter- 
val between ovulations in five captive 
raccoons held in Urbana, IIl., varied 
approximately from 80 to 140 days. 
Severe weather conditions (extreme cold 
or deep snow) interfered with the normal 
breeding cycle and resulted in an un- 
usually large number of late litters. Fe- 
male raccoons sometimes gave birth to 
two litters in one season, but they did 
not rear more than one litter in one 
season. The vaginal smear was no more 
specific for indicating estrus than was 
gross vulval swelling. 

7.—Contrary to published reports, the 
raccoon is a spontaneous ovulator. 
Ovulation was followed by the formation 
of corpora lutea whether the animal be- 
came pregnant or pseudopregnant. The 
formation of corpora lutea always re- 
sulted in changes in the uteri and nipples. 
The nipples always enlarged; some be- 
came heavily pigmented, some became 
slightly pigmented, and others remained 
unpigmented. Thus, it was possible to 
determine whether a female raccoon had 
ovulated by examining her nipples. 
Corpora lutea in both isolated and non- 
isolated pseudopregnant females formed 


Vol. 31, Art. 2 


from ovulated Graafian follicles and no 
from luteinization of follicles. Corpora 
lutea persisted in pregnant females until 
parturition and apparently disappeared 
14-16 days after parturition. 

8—Raccoons that ovulate become 
either pregnant or pseudopregnant, and 
the corpora lutea persist for about the 
same time in pseudopregnant raccoons as 
they do in those that give birth to young. 
Corpora in females that went at least 
halfway to term persisted about the same 
length of time whether the young wel 
aborted, were resorbed, or were born and 
were removed at birth or nursed until 
weaned. Field evidence indicated that 
in Illinois about 2.5 percent of the adult 
females were pseudopregnant each year. 

9.—Ten of 21 captive yearling females © 
and 9 of 14 wild yearling females were 
sexually immature. ; 

10.—Interstitial tissue occurred in the 
ovaries at some stage of the reproductive ~ 


seldom occurred when corpora lutea 
were present. Ovaries of females less) 


average, more interstitial iene ae h 
relatively and absolutely—than did those 


were not present; from July through ) 
December, ovaries from adults contained | 
more interstitial tissue than did those 
collected earlier in the year. The greatest” 
abundance of interstitial tissue was in 
ovaries taken from juveniles during Oc- - 
tober and November, and the maximum 
ovarian weights recorded during this 
study were those of juvenile females in 
November. 

11—The placenta of Procyon is de ~ 
ciduous, as in the dog, cat, fox, and sea 
and is endotheliochorial. If used wi 
caution, placental scars in raccoons aré 
useful for estimating litter size and ra 
of productivity. The significance 0 
multiple groups of scars is not clear, but 


ae 


si 


July, 1973 Sanperson & NaLBanpov: RepropucTIvE CycLe oF THE Raccoon 81 


it appears that each embryo that reaches 
1 month of age is represented by one 
scar that persists for 10 or more months. 
Scars in wild females with only one 
group of scars probably reflect implanta- 
tion rates for the preceding breeding 
season. Placental scars apparently per- 
sist longer in wild females than in cap- 
tives. 

12._The reproductive system of the 
male raccoon is similar to that of the dog; 
seminal vesicles and Cowper’s glands are 
lacking. 

13.—The uterus of the raccoon is inter- 
mediate between the bicornuate and the 
bipartite uterus. There is a single cervix 
and the horns are distinct, but after they 
join externally to form the single uterine 
body, the uterine lumina remain separate 
to a point near the cervix. The ovoid 
ovary is completely surrounded by the 
bursa ovarii. The sac is intact except 
for a small slit on one side, not large 
enough to permit passage of the ovary, 
as in the mink, dog, and fox. 

14.—The level of sex hormones in the 
male affected the enlargement of the 
preputial orifice and the maturation of 


the penis bone but had little or no effect 
prior to 7 months of age. Castration in 
the male also apparently caused a slight 
delay in the closure of the epiphyseal 
cartilage in the radius and ulna. Re- 
moval of the ovaries, even after raccoons 
had reached sexual maturity but before 
the epiphyses had closed, delayed the 
rate of epiphyseal closure. 

15.—Embryos persisted for about 19 
days after castration in each of two rac- 
coons—to 33 days short of term in one 
female and 7 days short of term in the 
other. 

16.—Limited studies indicated that in- 
jections of androgen did not initiate nor 
prolong spermatogenesis and apparently 
did not influence the size of the testes. 

17.—A dose of 35-50 IU of pregnant 
mare’s serum given subcutaneously each 
day for 12 days caused development of 
Graafian of follicles in adult females at 
any time of the year. A dose of 200 IU 
given on the 13th day caused ovulation 
48-60 hours later; however, in all cases 
of successful ovulation, the ovaries were 
much larger—and the rates of ovulation 
much higher—than normal. 


LITERATURE CITED 


ALLEN, B. M. 1904. The embryonic develop- 
ment of the ovary and testis of the mam- 
mals. American Journal of Anatomy 3(2) : 
89-146 + 7 plates. 

ALTMANN, F. 1927. Untersuchungen tber das 
Ovarium von Talpa europaea mit besonderer 
Beriicksichtigung seiner cyclischen Verander- 
ungen. Zeitschrift fir Anatomie und Ent- 
wicklungsgeschichte 82:482—569. 

AspELL, S. A. 1946. Patterns of mammalian 
reproduction. Comstock Publishing Co., 
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Berarp, E. V. 1952. Evidence of a late birth 
for the raccoon. Journal of Mammalogy 
33 (2) :247-248. 

BraMBELL, F.W.R., and I. H. Mitts. 1948. 
Studies on sterility and prenatal mortality 
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bryos after implantation. Journal of Experi- 
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Burt, W. H. 1960. Bacula of North American 
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of Zoology Miscellaneous Publication 113. 
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Conaway, C. H. 1955. Embryo resorption and 
placental scar formation in the rat. Journal 
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Corner, G. W. 1932. Cytology of the ovum, 
ovary and Fallopian tube. Pages 1567-1607 
in E. V. Cowdry, ed. Special cytology, 2nd 
ed. Vol. 3. Paul B. Hoeber, Inc., New York. 

Davis, D. E., and J. T. Emuen, Jr. 1948. The 
placental scar as a measure of fertility in 
rats. Journal of Wildlife Management 
12(2) : 162-166. 

Deanesty, R. 1935, XI—The reproductive 
processes of certain mammals. Part IX— 
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Deno, R. A. 1937. Uterine macrophages in 
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456 + 8 plates. 

1941. A criterion for distinguishing 
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Dorney, R. S. 1953. Some unusual juvenile 
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34(1) :122-123. 

Exper, W. H. 1952. Failure of placental scars 
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GeorcE, J. L., and M. Stirr. 1951. March 
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igan. Journal of Mammalogy 32(2):218. 

Gotpman, E. A. 1950. Raccoons of North and 
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Interior, Fish and Wildlife Service, North 


82 


American Fauna 60. U.S. Governmer 
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Hammonp, J., Jr. 1952. Gonadotroph 
induced ovulation in mink. Journal of Mam 
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Hansson, A. 1947. The physiology of n 
production in mink (Mustela vison, Schreb) 
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tion. Institute of Animal Breeding, Roy 
Agricultural College of Sweden, Stockhol; 
Acta Zoologica 28. 136 pp. 4 
His, W. 1865. Beobachtungen Uber den Ba 
des Saugethiereierstockes. Archiv flr Mikre 
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Jascer, E. C. 1947. Use of the os phallus ¢ 
the racoon [sic] as ripping tool. Journal | 
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Kincssury, B. F. 1914. The interstitial cel 
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Linu, R. D. 1954. Histopathologic techi 
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ment 17(3) :320-321. 

, and R. K. Enpers. 1954a. Ovulai 
in the raccoon. Journal of Mammalog 
35(3) :440. : 

, and . 1954b. Trans-uterine mi 
gration in the raccoon. Journal of Man 
malogy 35(3) :439. 

MILLARD, C. 1939. Raccoon experiment. Y 
consin Conservation Bulletin 4(3) : 28-29) 

Mompers, H., and C. Conaway. 1956. Th 
distribution “a placental scars of first ami 
second pregnancies in the rat. Journal 1 
Embryology and Experimental Moni 
4(4) :376-384 + 2 plates. 

Montcomery, G. G. 1969. Weaning | 
tive raccoons. Journal of Wildlife wa 
ment 33(1):154-159. 

Mossman, H. W. 1937. Comparative mo 
phogenesis of the fetal membranes and 2 al 
cessory uterine structures. Contributions } 
Embryology 158, Carnegie Institution | 
Washington Publication 479. 129-246 +4 
plates. 

Naxsanpov, A. V. 1958. Reproductive phy! 
iology. W. H. Freeman and Company, ba 
Francisco. 271 pp : 

ParzeLt, V. 1955. "Uber das Ovarium du 
Karnivoren und seine Zwischenzellen. Ze 
schrift fiir Mikroskopisch-Anatomische Fo 
schung 61(3) :309-359. 

Pearse, A. G. E. 1960. Histochemistry: » thi 
oretical and applied, 2nd ed. Little, Brow 
and Company, Boston. 998 pp. 

Porr, C. H. 1944. Attainment of sexual m 


1 


ily, 1973 SanperRson & NALBANDOV: REPRODUCTIVE CycLE OF THE Raccoon’ 83 


turity in raccoons. Journal of Mammalogy 
B5(1'):91. 

sMUSSEN, A. T. 1918. Cyclic changes in 
the interstitial cells of the ovary and testis 
in the woodchuck (Marmota monax). En- 
docinology 2:353-404 + 4 plates. 

NKER, G. C. 1944. Os clitoridis from the 
racoon [sic]. Journal of Mammalogy 25 
(1) :91-92. 

ANDERSON, G. C. 1950. Methods of measur- 

ing productivity in raccoons. Journal of 

Wildlife Management. 14(4) :389-402. 

1961a. Techniques for determining 

age of raccoons. Illinois Natural History 

Survey Biological Notes 45. 16 pp. 

1961b. The lens as an indicator of 

age in the raccoon. American Midland 

Naturalist 65(2) :481—485. 

NypDER, R. L., and J. J. Curistian. 1960. 
Reproductive cycle and litter size of the 
woodchuck. Ecology 41(4) :647-656. 

ooTer, C. A. 1946. Muskrats of Tule Lake 
Refuge, California. Journal of Wildlife 
Management 10(1) :68-70. 

TAFFORD, W. T., and H. W. Mossman. 1945. 
The ovarian interstitial gland tissue and its 
relation to the pregnancy cycle in the guinea 
pig. Anatomical Record 93(1) :97—107. 

brains, H. J. 1956. The raccoon in Kansas: 

| natural history, management, and economic 
| importance. University of Kansas Museum 


of Natural History and State Biological Sur- 
vey of Kansas Miscellaneous Publication 10. 
76 pp. 

Stocxarp, C. R. 1932. Cellular changes in 
the fluid of the mammalian vagina. Pages 
1611-1629 in E. V. Cowdry, ed. Special 
cytology, 2nd ed. Vol. 3. Paul B. Hoeber, 
Inc., New York. 

Stuewer, F. W. 1943a. Raccoons: _ their. 
habits and management in Michigan. Eco- 
logical Monographs 13(2) : 203-257. 

. 19436. Reproduction of raccoons in 
Michigan. ‘Journal of Wildlife Management 
7(1) :60-73. 

U. S. DEPARTMENT OF AGRICULTURE, Bureau 
of Biological Survey. 1936. Raising raccoons. 
Wildlife Research and Management Leaflet 
BS-34. 2 pp. 

U. S. WEATHER Bureau. 1960. Climatological 
data: Illinois. 65(1-3) : 1-43. 

Watson, M. 1881. On the female organs and 
placentation of the racoon (Procyon lotor). 
Royal Society of London Proceedings 
32(213) :272-298 + 4 plates. 

Wuitney, L. F., and A. B. UNDERWoobp. 1952. 
The raccoon. Practical Science Publishing 
Company, Orange, Conn. 177 pp. 

Woop, J. E. 1955. Notes on reproduction and 
rate of increase of raccoons in the Post Oak 
Region of Texas. Journal of Wildlife Man- 
agement 19(3) :409-—410. 


INDEX 


A 


Ampullae, 65 
Androgen (see testosterone) 


Birth date 

estimating, 46-47 

mean, 32, 45-46, 79-80 
Bulbo-urethral glands (see Cowper’s glands) 
Bursa ovarii, 67, 81 


Cc 


Canis lupus (see wolf) 

Castration 
effects on females, 34, 50-51, 68-70, 79, 81 
effects on males, 34, 68, 81 


Cat 
domestic, 57 
Cervix, 66, 81 


CGH (see chorionic gonadotropin) 
Chorionic gonadotropin, 34, 73, 75-76, 78-79 
Corpora albicantia, 55-56 
Corpora lutea, 32-33, 35, 53-61, 71-75, 77-80 
persistence, 53-56, 80 
secretion of progesterone, 57 
Cowper’s glands, 65, 81 


D 
Dog, 65, 67, 80-81 
E 
ECP (see estradiol) 
Embryos 
persistence after castration, 34, 63, 81 
Epiphyses 


closure, 68-69 

effects of castration on closure, 68, 81 
Estradiol, 34-35, 51-53, 72, 79 
Estrous cycle, 32, 48-53 


F 


Felis catus (see cat, domestic) 

Females 
percentage sexually mature as yearlings, 

56-57 

Fimbria, 67 

Follicle-stimulating hormone, 34-35, 52-53, 
71-72, 74-79 

Fox, 65, 67 

FSH (see follicle-stimulating hormone) 


G 


Gonad(s) (see ovary and testis) 
Graafian follicles, 34, 72-76, 80 
effect of pregnant mare’s serum, 81 


84 


H 


Histology, 33 

HMG-J5 (see human menopausal gonadotro- 
pin) 

Hormones, effects of exogenous 
on females, 34-35, 71-78, 81 
on males, 34, 70-71, 81 

Human menopausal gonadotropin, 35, 76-78 


Interstitial tissue, 33, 57-61, 80 


L 


LH (see luteinizing hormone) 
Luteinizing hormone, 34-35, 52-53, 71-72, 
74-79 


M 


Mammae 

pigmentation, 54, 57, 80 
Marmota monax (see woodchuck) 
Mink, 59, 67 
Muskrat, 61 
Mustela erminea (see stoat) 
Mustela vison (see mink) 


N 


Nipples (see mammae) 


fe) 


Ondatra zibethicus (see muskrat) 
Oryctolagus cuniculus (see rabbit, European) 
Os baculum, 66, 68 
Os clitoridis, 67-68 
Os penis (see os baculum) 
Ovary(ies), 67, 81 
seasonal cycle, 30, 43-45, 79 
Ovulation, 32-33, 53-55 
days between, 49 
effect of pregnant mare’s serum, 71-74, 77, 
81 
spontaneous, 54, 80 


Pp 


Placenta, 62, 80 

Placental scars, 33, 61-65, 80-81 

PMS (see pregnant mare’s serum) 

Pregnant mare’s serum, 34-35, 52-53, 60-61, 
71-74, 76-79, 81 

Procyon lotor (see raccoon) 

Progesterone, 34-35, 51-53, 57, 71, 73-74, 
77-79 

Prostate, 65 

Pseudopregnancy, 55-56, 60-61, 80 


ee 


July, 1973 SanpErson & NALBANDov: REPRODUCTIVE CYCLE OF THE Raccoon 


R 


Rabbit, European, 61 
Raccoon(s) 
cage for captive, 29 
captive, 29, 30-31 
Reproductive tract (s) 
female, 34, 66-68, 81 
male, 33, 65-66, 81 


S 


Seminal vesicles, lacking, 65, 81 
Sex ratios 

secondary, 32, 47-48, 80 
Sexual maturity 

females, 55-57, 80 

males, 35-36, 79 
Spermatogenesis 

effect of testosterone, 70, 81 
Stoat, 61 


T 


Testis (es) 
seasonal cycle, 30, 35-43, 79 
Testosterone, 34, 70, 81 


U 
Urethra, 65 
Uterine milk, 35, 78-79 
Uterus, 66-67, 81 


V 


Vaginal biopsies, 52-53 
Vaginal smears, 50-51 
Vasa deferentia, 65 
Vulpes fulva (see fox) 


WwW 


Wolf, 65 
Woodchuck, 57 


85 


YOME FUDINCATIONS OF THO PRLENNAED TNE ENS AR FEE NEN 


BULLETIN 


Volume 30, Article 3.—Migrational Behavior of 
Mallards and Black Ducks as Determined 
from Banding. .By Frank C’ Bellrose and 
Robert D. Crompton. September, 1970. 68 
p., frontis., 25 -fig,,. bibliogr., index. 

Volume 30, Article 4-Fertilization of Estab- 
lished "Trees: = A Report of Field Studies. By 
Dan Neely, E. B. Himelick, and Webster R. 
Crowley,. Jr. September, 1970. 32 p., fron- 
tis:, 8-fig., bibliogr., index. 

Volume-30, Article 5.—A Survey of the Mussels 

*«(Unionacea) of the Illinois River: A Pollut- 
‘ed Stream. By William C. Starrett. February, 
tis., 8 fig., bibliogr., index. 

Volume 30, Article 6—Comparative Uptake 
and Biodegradability of DDT and Methoxy- 
chlor by Aquatic Organisms. By Keturah A. 
Reinbold, Inder P. Kapoor, William F. 
Childers, Willis N. Bruce, and Robert L. 
Metcalf. June, 1971. 12 p., frontis., 5 fig., 
bibliogr., index. 

Volume 30, Article 7—A Comparative Study of 
Two Components of the Poinsettia Root Rot 
Complex. By Robert S. Perry. August, 1971. 
35 p., frontis., 10 fig., bibliogr., index. 

Volume 30, Article 8—Dynamics of Condition 
Parameters and Organ Measurements in 
Pheasants. By William L. Anderson. July, 
1972. 44 p., frontis., 6 fig., bibliogr., index. 

Volume 31, Article 1—The Effects of Supple- 
mental Feeding and Fall Drawdowns on the 
Largemouth Bass and Bluegills at Ridge 
Lake, Illinois. By George W. Bennett, 
H. Wickliffe Adkins, and William F. Chil- 
ders. January, 1973. 
fig., bibilogr., index. 


BIOLOGICAL NOTES 


70.-An Ecological Study of Four Darters of the 
Genus Percina (Percidae) in the Kaskaskia 
River, Illinois. By David L. Thomas. De- 
cember, 1970. 18 p., 11 fig., bibliogr. 

71—A Synopsis of Common and Economic 
Illinois Ants, with Keys to the Genera 
(Hymenoptera, Formicidae). By Herbert 
H. Ross, George L. Rotramel, and Wallace 
E. LaBerge. January, 1971. 22 p., 27 fig., 
bibliogr. 

72.-The Use of Factor Analysis in Modeling 
Natural Communities of Plants and Ani- 
mals. By Robert W. Poole. February, 1971. 
14 p., 14 fig., bibliogr. 

73.-A Distributional Atlas of Upper Mississip- 
pi River Fishes. By Philip W. Smith, Alvin 
C. Lopinot, and William L. Pflieger. May, 
1971. 20 p., 2 fig., 107 maps, bibliogr. 


List of available publications mailed on request 


28 p., frontis., 8 


wyVNTeRe 


74.-The Life History of the Slenderhead Da 
er, Percina phoxocephala, in the Embar 
Rive., Illinois. By Lawrence M. Page a 
Philip W. Smith. July, 1971. 14 p., 10 fi 
bibliogr. 

75.-Illinois Birds: Turdidae. By Richard 
Graber, Jean W. Graber, and Ethelyn 
Kirk. November, 1971. 44 p., 40 fig., bib- 
liogr. 

76.-Illinois Streams: A Classification Based | 
Their Fishes and an Analysis of Factors 
sponsible for Disappearance of Native 
cies. By Philip W. Smith. November, 1 
14 p., 26 fig., bibliogr. 

77-The Literature of Arthropods Associa 
with Soybeans. I. A Bibliography of 
Mexican Bean Beetle, Epilachna vari 
Mulsant (Coleoptera: Coccinellidae) 
M. P. Nichols and M. Kogan. Febr 
1972. 20 p., 1 fig., bibliogr. 

78.-The Literature of Arthropods Assoei 
with Soybeans. II. A Bibliography of 
Southern Green Stink Bug, Nezara vir 
(Linneaus) (Hemiptera: Pentatomic 
By N. B. DeWitt and G. L. Godfrey. 
1972. 23 p., 1 fig., bibliogr. } 

79.-Combined Culture of Channel Catfish 
Golden Shiners in Wading Pools. Bi 
Homer Buck, Richard J. Baur, Charles 
Thoits III, and C. Russell Rose. April, 19; 
12 p., 3 fig., bibliogr. C 

80.-Illinois Birds: Hirundinidae. By Ric 
R. Graber, Jean W. Graber, and Ethelyn 
Kirk. August, 1972. 36 p., 30 fig., bibl 

8i.Annotated Checklist of the Butterflies 
Illinois. By Roderick R. Irwin and John 
Downey. May, 1973. 60 p., 3 fig., $ 
maps, bibliogr. ‘a 

82.Lactate Dehydrogenase Isozymes of Dar- 
ters and the Inclusiveness of the Ger 


Percina. By Lawrence M. Page and Greg- 
ory S. Whitt. May, 1973. 7 p., 5 figs 
bibliogr. a 
CIRCULAR 7 


46.-Illinois Trees: Their Diseases. By J. C 
ric Carter. June, 1964. (Third printi 
with alterations.) 96 p., frontis., 89 fig. : 

49-The Dunesland Heritage of Illinois. By 
Herbert H. Ross (in cooperation with Illin 
Department of Conservation). August, 1963. 
28 p., frontis., 16 fig., bibliogr. a 

51.Illinois Trees: Selection, Planting, and 
Care. By J. Cedric Carter. August, 1966 
123 p., frontis., 108 fig. 4 

52.-Fertilizing and Watering Trees. By D 
Neely and E. B. Himelick. December, 19' 
(Third printing.) 20 p., 9 fig., bibliogr. 

53.—Dutch Elm Disease in Illinois. By J. Ced 
Carter. October, 1967. 19 p., frontis., 17 f 


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Address orders and correspondence to the Chief, 
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Natural Resources Building, Urbana, Illinois 61801 


4 


el 


a 


| ILLINOIS 
@ tural History Survey 
] 7 BULLETIN 


Nutritional Responses 
of Pheasants to Corn, 


with Special Reference 

: to High-Lysine Corn 

* 

ze 

ali NATURAL HISTORY SURVEY 

iam L. Anderson NOV 14 1973 
LIBRARY 


RAL HISTORY SURVEY DIVISION 
‘ NA, ILLINOIS THE LIBRARY, OF THE 


NNV 7. 1079 VOLUME 31, ARTICLE 


| ILLINOIS 
‘atural History Survey 
| BULLETIN 


Nutritional Responses 

of Pheasants to Corn, 
with Special Reference 
to High-Lysine Corn 


| 
/ 
| 
| 
| 


i F. Labisky 
lam L. Anderson 


om ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


AURAL HISTORY SURVEY DIVISION 
‘RANA, ILLINOIS 


VOLUME 31, ARTICLE : 
JULY, 1973 


STATE OF ILLINOIS 


DEPARTMENT OF REGISTRATION AND EDUCATION 


BOARD OF NATURAL RESOURCES AND CONSERVATION 


DEAN BARRINGER, Ph.D., Chairman; THomAS Park, Ph.D., Biology; L. L. Stoss, Ph.D., Geology; (VACANT), 
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Section of Economic Entomology 


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STEVENSON Moore, III, Ph.D., Entomologist, Exten- 
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JAMES E. AppLesy, Ph.D., Associate Entomologist 

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ea 

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cago, Illinois; WILDLIFE RESEARCH, WILLARD D. KLIMSTRA, Ph.D., Professor of Zoology and Director of Co- 
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tistical Design and Analysis, University of Illinois. 


CONTENTS 


2 SEEN LLG nS) crocene: ev AG GO PAE SCs Eten CREE En 87 
VHETRIODS, ado on Sh Soe 5.66 Alo GOCE poe ee Oe TO eee Ae ee 88 
Heedinpeairialslcme vent esHensi na tiyansyoucitt acces teins sidered nae dawiaooek 88 
Heedincaminyalw lay ACUI HELeTES) ree eel aetee ftes cies Said <ts elese vas vege eskaeseeiew s 88 
SoWeciOnMmOo ime Data crn tasers Tee CR EN Hoc Be ceie wre eal sane hee 89 
ANDEINISES, yo nib. 6.5 doc Ace OCS RRREE A cele Re He ic. Ce ae aa 90 

J ESRINES Lo dufecipe edie. ode ee ites EER NOTE Ohta er 90 
Sleaycerarl eet Leni siimge te tortor tey- ner, teres <i> Ae teeta ya caices Re wear SIMA Sida ieses Gi vost Sw gen ahead ere as 90 
pda erehta Ghanges tr. mov ye etree a sekacis ere te Mew siadd asa ae ub oe 90 

[itoreyel: (Crovetsittay ot @) sy Sag Aca An Acie aa retrain ar oP On oe eee ee eet teriesor 91 

Dy ieesti bilttya Coenuclents. rs cer. eurereeee ete suey ato) ete heres aranese tities 20s %, ces era yoretereve le ee 91 
CalsrienWtilization perme mer sets eye esr ye mraite ce eta sy cir ornare ccun ho oh eiewitol te on ee 92 

Israfiein trier 5 SoSqcene Cocoon hacen eae ee Reais eee meter es 92 

HB ysirtemmOMtIMIZAELOMR rors 080-8 on <7 Teeth eer soe cep NOES ae aes soles « elete ohana eo. dias easter eNO 93 

LPRLIGTEAL "GS, TESTS Metaeaet ee ReLO ce. triza ciate Seu cee ene atte eee ieee retaretear 94 
Glandspanca Orcans eyee.t cera sls etek ois sis oe olkaye eigeNe ei seals me-she ahe siakereis eke « 94 
HanpanclahattyaeA Clas gtae ete, crsis theta te care cic ois cto pheteveiaia quate stone suave nishroa andes 94 

PCAN mL eTIS Meme yet fe FS cvenctes into secis xastarcret smite) njsceis eves oucea cave isc ene laieie, ciulelsv aise ye: aieea, Ore 97 
isayeky \WStane (Clete ea epee e es cha Be ch ol ee a ibid MDENee a elor eatRe onic aoe cec 97 
IDiwesinayihiny (CWeiinets Soe oop akon soho ctbmEe ato weehn aterEopeecneterT do 98 

(Callorare: WWiGIrAtn att a5 socio piracnodn OOO CEs BEOUODO SAA aro oR PM oor ar 98 

Pinglietey UWiRITCAIWOla y+ oe SecA cnc omic 30 dq cuter bo Olio nCeena ERB coortceey Nec eee 98 

Ib STG, LOMTKWATI GSE), onan Gnee Genes oon oO EOD an ewes eIppIOSr ce Onno ciimcecia 98 

Gievily aval’ OSA Rae aol coco capo erinoM cmos be Beer apie ac momncbe 99 

[Para cistahlbaning Nets bi ole aprotic 6 he oerionas Ana CRccn DA Ona e mS Ionics aicere 100 

BUTANE SACI toe TP fo) = ass apeRe ames cia- stetoiattee Vou essatiec at Sve. seb oPaayene whee! tice ayes cite emiedess eens 100 
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Nutritional Responses of Pheasants 
to Corn, with Special Reference 


to High-Lysine Corn 


IN LATE 1963, Purdue University sci- 
entists discovered, by amino acid analysis, 
that the endosperm of maize (Zea mays) 
kernels homozygous for the opaque-2 mu- 
tant contained about 70 percent more 
lysine than the endosperms of kernels of 
normal hybrids (Mertz et al. 1964; Mertz 
1966:12). The endosperm of opaque-2 
also contained greater amounts of tryp- 
tophan than that of normal corn (Pickett 
1966:19). 

Lysine and trytophan are among those 
amino acids that are dietary essentials 
for protein synthesis in many animals, 
including man. The proteins in endo- 
sperm of normal corn are of low biologi- 
cal quality. Thus the opaque-2 mutant, 
which alters the amino acid composition 
(particularly that of lysine, tryptophan, 
and leucine) of the maize endosperm, has 
offered the potential of a type of corn 
having exceptional nutritional values. 
The superior nutritional benefits of 
this modified-protein corn (hereinafter 
termed high-lysine corn) for growth have 
already been demonstrated in feeding 
experiments with rats (Mertz et al. 1965; 
Mertz 1966), swine (Pickett 1966; Jensen 
et al. 1967), chicks (Rogler 1966), and 
turkeys (Adams & Rogler 1970). 

The nutritional potential of high-ly- 
sine corn has led to predictions that this 
corn may replace a substantial acreage of 
normal-corn hybrids produced in the 
Corn Belt during the 1970's. The esti- 
mated acreage of high-lysine corn plant- 
ed in the United States in 1972 was 
80,000-100,000 acres (D. E. Alexander, 
University of Illinois, personal communi- 
cation, January 12, 1973). Inasmuch as 
corn is important in the diet of many 
wild animals, the widespread use of high- 


Ronald F. Labisky 
William L. Anderson 


lysine corn offers a potential nutritional 
benefit to wild birds and mammals. 

Corn figures more prominently in the 
diet of midwestern pheasants, particu- 
larly in fall and winter, than it does for 
most wildlife species (Korschgen 1964: 
170, 173). To illustrate, during fall and 
winter, corn constitutes at least 80 percent 
(by weight) of the total food intake by 
pheasants in thriving populations in east- 
central Illinois (Anderson & Stewart 
1969:261; R. F. Labisky, unpublished 
data). Yet despite the importance of 
corn to pheasants, little is known of its 
nutritional attributes for growth, main- 
tenance, or reproduction. Furthermore, 
juvenile hens, in contrast to adult hens, 
suffer a disproportionately high rate of 
nonhunting mortality between fall and 
winter in Illinois (R. F. Labisky, unpub- 
lished data). That the onset of this 
mortality among juvenile hens coincides 
with that time of the year at which waste 
corn from the harvest suddenly becomes 
abundantly available suggests a potential 
causal link between unbalanced nutrition 
and mortality. Hence the objectives of 
this study were to ascertain the physio- 
logical responses of juvenile hen pheas- 
ants in fall, and of adult hen pheasants 
in late winter and early spring, to ex- 
clusive diets of both normal corn and 
high-lysine corn. 


ACKNOWLEDGMENTS 


Acknowledgement is due the following 
members of the Department of Agrono- 
my, University of Illinois. Dr. D. E. 
Alexander supplied the corns, provided 
their lysine and fatty acid profiles, and 
offered advice on various aspects of the 


88 Inuinois Narurat History SuRvEY BULLETIN 


study. Dr. T. R. Peck and G. G. Stone 
offered laboratory facilities for, and ma- 
terially aided in, the analyses of pheasant 
excreta for nitrogen. Dr. I. de la Roche 
analyzed the fat samples for determina- 
tion of fatty acids. Dr. C. M. Wilson 
analyzed the commercial ration for ami- 
no acids. 

Dr. B. G. Harmon, Department of 
Animal Science, University of Illinois, 
supervised the analyses of pheasant ex- 
creta for lysine. 

Dr. G. C. Sanderson, Illinois Natural 
History Survey, offered editorial sugges- 
tions during preparation of the paper, 
and O. F. Glissendorf edited the final 
manuscript. D. R. Vance and J. E. Mc- 
Clendon of the Survey assisted in various 
aspects of the experiment. 

Special thanks are due Drs. Alexander 
and Harmon, and Dr. J. E. Savage, De- 
partment of Poultry Science, University 
of Missouri, for critically reviewing the 
manuscript. 


METHODS 
FEEDING TRIAL I: JUVENILE HENS 


The 21 juvenile hens used in the ex- 
periment were obtained from the Illinois 
State Game Farm, Yorkville, in 1966. 
These hens, which had hatched on June 
20, were transported to Urbana on Sep- 
tember 13. The hens were held in two 
wire-bottomed 3.0 x 3.9 x 1.8-meter 
pens and fed a commercial flight and 
maintenance chow (FMC) until October 
3 when they were individually placed, by 
random assortment, in 70 x 60 x 34-cm 
cages. The cages had thin-walled fiber- 
glass sides, top, bottom, and rear, which 
prevented sight contact between birds. 
The birds were fed a diet of two-thirds 
FMC and one-third normal corn (whole 
kernels) for the period October 3-14 to 
acquaint them with corn, and then an 
exclusive diet of FMC for the period 
October 15-20. 


Inasmuch as 19 of the 21 hens post- 
ed gains in body weight between Octo- 


Vol. 31, Art. 3 


ber 10 and October 20, the feeding trial 
was begun on the latter date. Three 
groups of 7 hens each were randomly 
selected to be fed exclusive, unrestricted 
diets of FMC, normal hybrid corn (Pio- 
neer 3306), or high-lysine corn (Table 
1), and water ad libitum. The FMC was 
pressed into corn-sized pellets for the 
feeding trials (see Frontispiece). ‘The 
experiment was terminated 8 weeks later, 
December 15. One hen from the group 
of hens fed normal corn died from an 
injury during the trial. 


FEEDING TRIAL II: ADULT HENS 


The 12 adult hens, 3 and 4 years old, 
used in the feeding trial were also of 
game-farm origin. These hens had been 
transported to Urbana as juveniles, and 
subsequently maintained in wire-bot- 
tomed outdoor pens, similar to those used 
to house the juveniles. On February 7, 
1967, these hens were individually placed, 
by random assortment, in the same cages 
in which the juveniles of Trial I had 
been held. They had been fed an intro- 


ductory diet of one-half FMC, one- — 


fourth normal corn, and one-fourth high- 
lysine corn for the period February 1-7. 
Because of their quick acceptance of corn, 
they were returned to an exclusive FMC 
diet on February 8. 

All 12 hens posted gains in body weight 
during the interval of February 27—March 
6; therefore, the feeding trial was begun 
on the latter date. Six hens were offered 
a diet of normal corn and six hens a diet 
of high-lysine corn (Table 1); both 
groups had unrestricted access to water. 
The food intake by adult hens was re- 
stricted-to 200 g of corn per bird per 
week. The corn was provided in two 
100-g lots, on the first and fourth days 
of each week. This limited offering of 
corn was judged to be about 60 percent 
of a normal weekly intake, and was in- 


tended to simulate the estimated poten- — 


tial food intake of wild hens subjected to 
the rigors of late winter in Illinois. The 
experiment was terminated after 7 weeks, 
on April 24. 


July, 1973 Lasisky & ANDERSON: 


NutTriTIONAL RESPONSES OF PHEASANTS 


89 


Table 1.—Mean concentrations of calories, crude protein, lysine in protein, and selected min- 
erals in diets of a commercial flight and maintenance chow (FMC), of normal corn, and of high-lysine 


corn that were fed to hen pheasants in 1966 and 1967. 
Diets 
High-Lysine Corn: 
Normal Corn: O paque-2 
FMC Pioneer 3506 Synthetic Ao 
Calories per g* 4,278.9 + 4.4° 4,651.6 + 99.1 4,544.0 + 6.1 
Percentage crude 
protein® 22.8 = 0:03 (0) S55 (Opie ilileyh Se) Co) 
Percentage lysine‘ 
in protein au) 3.24 4.7 
Percentage fiber <12.0 2.0 2.0 
Percentage saturated: 
unsaturated fatty acids’ 0 14:86 19:81 
ppm of major elements 
calcium 14,443 + 1,469 41 sE.3 oy) ass il 
magnesium 1,833 = 24 166 22°48 Cyl Se. aif! 
sodium 2,785 + 49 204 = 9 195 + 4 
potassium 72997 S75159 3,249 = 138 3,924 = 184 
phosphorus ey) 2a) 40) 250 83e == 2900 2,264 + 202 


*Caloric contents of the rations differed 


multiple range test indicated that the caloric 
other. 


> Standard errors. 


significantly (F=11.18, 4; P!< 0.05); application of Duncan’s 
content of both corns differed from FMC but not from each 


° Crude protein compositions of the rations differed significantly (F—989.66,,,; P <0.05); application 
of Duncan’s multiple range test indicated that the caloric content of both corns differed {rom FMC, but not 


from each other. 


4 This particular hybrid contains slightly greater concentrations of protein and of lysine than most normal 


corn hybrids. 
lysine. 
¢ Lysine content is for defatted samples. 


{ Yhe principal fatty acids in the corns are unsaturated: 


The average normal corn hybrid contains about 10.5 percent protein, of which 2.8 percent is 


oleic and linoleic. The normal and high-lysine 


corns contained, respectively, 34 and 22 percent oleic, and 52 and 58 percent linoleic acid. 


9 Data are not available. 


COLLECTION OF DATA 


Body weights of the hens were recorded 
at the onset of the feeding trials and at 
weekly intervals thereafter. Correspond- 
ingly, the amount of food consumed by 
each hen during each week of the feed- 
ing trial was measured to the nearest 
gram. All food consumption was con- 
verted to a dry weight standard. The 
total excreta was collected for each ju- 
venile hen for Weeks 4, 5, 6, 7, and 8 
(final) and for each adult hen for 
Weeks 5 and 7 (final). 

After the final body weight of the hens 
had been recorded, each hen was placed 
in an inverted position and decapitated. 
All birds were then dissected. The fol- 
lowing muscles, fat deposits, organs, and 
glands were excised from both juvenile 
and adult hens, and weighed: muscles of 
right half of the sternum (pectoralis thor- 


acia, ventral head of the supracoracoi- 
deus, and coracobrachialis—nomencla- 
ture as used by Hudson & Lanzillotti 
1964: 13-15); fat strip and visceral fat 
(as described by Breitenbach & Meyer 
1959: 1017); liver; thyroids; parathy- 
roids; and adrenals. The heart, pan- 
creas, gizzard, kidneys, spleen, and thy- 
muses from juvenile hens were also 
excised and weighed, as were the ovary, 
oviduct, and largest ovum from the adult 
hens. The organs and glands, after be- 
ing freed of extraneous material, were 
blotted carefully with paper toweling to 
remove excess blood and moisture prior 
to being weighed. The heart and liver 
were opened and blood clots therein re- 
moved: the gall bladder was excised from 
the liver. The contents, but not the lin- 
ing, were removed from the gizzard be- 
fore the latter was weighed. The weights 


90 


recorded for kidneys and endocrine 
glands are for paired (right and left) 
measurements. 


ANALYSES 


Each sample of food and excreta was 
oven-dried at 60°C for 142 hours, finely 
ground, and then sealed in a sterile plas- 
tic bag for subsequent determination of 
nitrogen (crude protein), lysine, and 
caloric content. 

Nitrogen content of foods and excreta 
was determined by Kjeldahl procedures; 
the crude protein content of each item 
was calculated as nitrogen x 6.25. Crude 
protein determinations were made for five 
samples of each of the three foods, and 
for single samples of dried excreta from 
each juvenile hen for each of the last 
5 weeks of the 8-week experiment and 
for each adult hen for Weeks 5 and 7 
of the 7-week experiment. 

Lysine in the pheasant excreta was 
measured, following acid hydrolysis under 
vacuum for 16 hours, by chromatographic 
analysis (Beckman Amino Acid Analyzer, 
Model 120). Lysine determinations for 
excreta were made from composites of 
the weekly samples for Weeks 4—8 of the 
8-week experiment for each juvenile hen, 
and for Weeks 5 and 7 of the 7-week 
experiment for each adult hen. The 
amount of lysine in the foods (defatted) 
was also measured by chromatographic 
analysis; approximately 80 percent of the 
nitrogen in the foods was recovered as 
amino acids. 


Caloric content of foods and excreta 
was measured by standard caloric-bomb 
techniques. Calories were measured 
from three samples of each of the three 
foods, and from a composite of the five 
and two weekly collections of excreta 
from each juvenile and each adult hen 
pheasant, respectively. 

The mineral content of the foods was 
derived by atomic absorption spectropho- 
tometry (for Ca and Mg), flame spectro- 
photometry (for Na and K), and color- 
imetry (for P). 

The null hypothesis, in all tests for de- 
termination of statistical differences, was 
accepted or rejected at the 0.05 level of 
probability. 


FINDINGS 


JUVENILE HENS 
Body Weight Changes 


The juvenile hens that were fed ex- 
clusive diets of FMC or high-lysine corn 
posted gains in body weight that aver- 
aged 98.4 and 23.4 g, respectively, during 
the 8-week feeding trial; those fed normal 
corn suffered losses that averaged 8.7 g 
(Table 2). Both groups of hens to 
which corn was fed exhibited marked 
declines in body weight during the first 
week of the feeding trial (Fig. 1). In 
the final analysis, all of the seven hens 
fed FMC, five of the seven hens fed 
high-lysine corn, and three of the six hens 
fed normal corn gained weight during the 


Table 2.—Body weight statistics for juvenile hen pheasants fed exclusive diets of flight and 
maintenance chow (FMC), of normal corn, or of high-lysine corn for an 8-week period, October 20— 


December 15, 1966. 


Mean Body Weight (g) or Weight Change 
for Specified Diet 


FMC Normal Corn High-Lysine Corn F Valuesary 
(n=7 Hens) (n=6 Hens) (n=7 Hens) 
Initial weight (Oc- 
tober 20) MAAS =) 2OE2e 742.2 = 36.9 152 = 324: 0.452,37 
Final weight (De- 
cember 15) 809.7 + 29.6 733.5 = 35.8 776.1 += 39.6 1.132,17 
Weight change +98.4 = 7.6 —8.7 + 14.7 Sree! Se D7 fa 4.702,17* 


* Denotes statistical significance, P <0.05. All combinations of paired means differed significantly. 


* Standard errors. 


Inuinois NatrurAt History Survey BULLETIN Vol. 31, Art. 3 


| 


July, 1973 Lasisky & ANDERSON: NurritionaL RESPONSES OF PHEASANTS 91 


+100 


+80 


+60 


GRAMS GAIN OR LOSS IN BODY WEIGHT 


eee 


Sie 


<7 FMC (n=7) 


+20 
HIGH-LYSINE 
CORN 
(n=7) 
oO 
pee gr 7349) 
Bea —_ (7349 
NORMAL CORN 
(n=6) 
-40 
(0) | a 3 4 5 6 Tf 8 


(OCTOBER 20) (DECEMBERIS) 


WEEKS OF STUDY 
Fig. 1.—Mean change in body weight (g) by weekly periods among juvenile hen pheasants fed 
exclusive diets of FMC, of normal corn, or of lysine corn; the vertical lines transecting the means are 


standard errors. 


right of the graph, respectively. Statistically significant 
occurred among diets for three of the eight successive weekly periods: 


3 (F= 3.93); and 6 to 7 (F=3.59). 


8 weeks. The extremes in weight change 
among hens on each of the diets were: 
FMC, +128 and +59 g; high-lysine 
corn, +177 and —88 g; and normal 
corn, +44 and -71 g. 


Food Consumption 


The three diets, fed ad libitum, were 
consumed by the juvenile hens in signifi- 
cantly different amounts; the greatest in- 
take was of FMC and the lowest was of 
normal corn (Table 3). Inasmuch as 
the caloric, crude protein, and lysine con- 
tents of the high-lysine corn were either 
similar to or greater than those of normal 
corn (Table 1), thereby discounting com- 
pensatory nutritional needs, the greater 
rate of consumption of high-lysine corn 


The mean initial and final body weights for each group are given at the left and 


(P<0.05; 2 and 19 df) 
initial to 1 


weight changes 
(F = 3.73); 2 to 


per hen suggested that it may have been 
more palatable’ to pheasants than nor- 
mal corn. Changes in body weight per 
100 g¢ of food consumed averaged +-4.3, 
+1.3, and -0.5 g on FMCG, high-lysine 


corn, and normal corn, respectively. 


Digestibility Coefficients 


Significant differences in the digesti- 
bility coefficients? were exhibited by hens 


1 Food intake is often depressed if the animal’s 
diet is deficient in either protein or an indispensable 
amino acid (see review by Harper, 1967). Therefore, 
the greater consumption of high-lysine corn over nor- 
mal corn by the hens may have reflected its higher 
lysine content rather than any superiority in palat- 
ability. 

2 Digestibility coefficient = 


[ = dry weight of excreta )x 100 | 
Total dry weight of food consumed 


92 Ituois Naturat History Survey BuLLETIN 


on the different diets (Table 3). Hens 
fed FMC digested substantially less (59.1 
percent) of their ration than did those 
hens fed either normal corn (82.5 per- 
cent) or high-lysine corn (81.2 percent). 
In contrast to the corns, the compara- 
tively low rate of digestibility of FMC, 
in part, reflected its higher fiber content. 
Despite the similarity in the mean digesti- 
bility coefficients of the two corns, they 
were significantly different because of the 
extremely narrow range of variation in 
the digestibility of each of the corns by 
the individual hens. 


Calorie Utilization 


Although the total intake of calories 
by the juvenile hens was significantly 
greater for those fed FMC than for those 
fed either normal corn or high-lysine 
corn, the utilization of the calories by the 
hens receiving the FMC was markedly 
less than for those receiving either of 
the corns; the metabolizability coeffi- 


Vol. 31, Art. 3 


cients* were 66.3, 84.8, and 83.3 for 
FMCG, normal corn, and high-lysine corn, 
respectively (Table 4). Because of this 
different proportionate utilization of cal- 
ories, there was no significant difference 
in the total number of calories utilized 
per hen for birds on the three diets during 
the 8-week trial. Juvenile hens obtained 
2,837, 3,945, and 3,808 kcal of metab- 
olizable energy per kg of FMC, normal 
corn, and high-lysine corn consumed, re- 
spectively. 


Protein Utilization 


The intake of crude protein by the 
juvenile hens was significantly different 
among the birds fed the three diets, being 
more than twice as great for FMC as 
for either normal or high-lysine corn (Ta- 
ble 3). The high intake of crude protein 
by the hens fed FMC reflected not only 


8% Metabolizability coefficient = 
be 


Total calories in excreta 
Total calories in food consumed 


)x 100] 


Table 3.—Comparative consumption and utilization of three foods—flight and maintenance 
chow (FMC), normal corn, and high-lysine corn—fed as exclusive diets to different groups of juvenile 


hen pheasants for an 8-week period, October 20—December 15, 1966. 


dry weight. 


All values are expressed as 


Mean Value per Hen per Week 
for Specified Diet 


FMC Normal Corn High-Lysine Corn F Valuescs) 
(n=7 Hens) (n=6 Hens) (n=7 Hens) 

Food consumed (g) 282.4 = 12.9° 205.6 = 10.8 234.6 += 17.0 65.1 12,130* 
Crude protein 

consumed (g)? 64.4 = 1.0 DAN ae aU{ay<2= (0).7/ 888.5 1>,136* 
Lysine consumed (¢) ° 3.2 = 0.06 0.8 + 0.01 Lesa O10 Ss 2,305.5 12, 136* 
Excreta (g)" UPoroy ese 36.0 + 0.6 44.2 + 1.0 716.8 12,53 * 
Digestibility 

coefficient 

(percent) * 5911 S=) 0:3 8220 == 02) Siz 2e= se OS 2,266.672,s5 * 
Crude protein in 

excreta (percent) “ SOSh ee 5 47.9 + 0.9 47:2 + 0.7 20.920,s3 * 
Crude protein 

utilized (g)* 16.2 + 1.8 5 + 0.4 5.7 + 0.4 24.240 53 * 

(percent) * 26:0° == 2:5 D2 ENG 21.0 + 1.4 3.1 12,53 

Change in body 

weight (g) +12.3 —1.1 sess) 


* Denotes statistical significance, P < 0.05, 


different. 
* Standard errors. 


>See Table 1 for protein content of foods. 
© Product of crude protein consumed and amount of lysine in protein. 
4 Based on data for Weeks 4 through 8 only. 


L z Those means underscored by the same line are not significantly 
Interactions among weeks yielded no significant F values in any category. 


July, 1973 Lasisky & ANDERSON: NutriTionaL RESPONSES OF PHEASANTS 93 


Table 4.—Comparative consumption and utilization of calories by juvenile hen pheasants fed 
exclusive diets of a flight and maintenance chow (FMC), of normal corn, or of high-lysine corn for 


an 8-week period, October 20-December 15, 1966. 


Mean Value per Hen per 8-Week Period 
for Specified Diet 


FMC Normal Corn High-Lysine Corn F Valuescar) 
(n=7 Hens) (n=6 Hens) (n=7 hens) 

Calories consumed 

(kcal) 9,688 + 244% 7,501 + 179 8,357 + 542 6.762,17* 
Calories per g 

excreta? 31359) EP 19 3,892 + 23 3,888 + 30 156.992..7* 
Calories utilized 

(kcal)? 6,409 + 147 6,486 + 176 6,959 + 477 0.91217 

(percent) ° 66.3 + 0.4 84.8 + 0.3 83.3 + 0.2 1,005.632,17* 


* Denotes statistical significance, P<0.05. Those means underscored by the same line are not significantly 


different. 
* Standard errors. 


> Product of the 8-week consumtpion of calories and the 5-week (Weeks 4-8) percentage utilization of calories 


for each hen. 
© Calorig values for Weeks 4-8. 


their high rate of consumption of the ra- 
tion but also the ration’s high protein 
content (22.8 percent). Whereas the pro- 
portionate utilization of the crude protein 
consumed by the hens did not differ sig- 
nificantly among diets (Table 3), those 
hens fed FMC utilized nearly 214 times 
more crude protein than did hens fed 
either of the corns. Changes in body 
weight of the juvenile hens were related 
directly (r = 0.56, 18 df; P < 0.05) to 


the amount of crude protein utilized. 


Lysine Utilization 
The intake of lysine by the juvenile 
hens also differed significantly among the 


three diets; the hens fed FMC consumed 
about 24 and 4 times as much as those 
hens fed lysine corn and normal corn, 
respectively (Table 5). The propor- 
tionate utilization of lysine, however, was 
greatest for hens fed high-lysine corn 
(99.2 percent), and differed significantly 
from that for hens fed either normal corn 
(88.4 percent) or FMC (85.6 percent). 
The total amount of lysine utilized per 
hen during the 8-week feeding trial dif- 
fered significantly among the diets, ‘aver- 
aging 21.6 g on FMC, 10.2 g on high- 
lysine corn, and 5.6 g’on normal corn. 
The response in body weight of the ju- 
venile hens was strongly dependent (7 = 


Table 5.—Comparative consumption and utilization of lysine by juvenile hen pheasants fed 
exclusive diets of a flight and maintenance chow (FMC), of normal corn, or of high-lysine corn for 


an 8-week period, October 20—December 15, 1966. 


Mean Value per Hen per 8-Week Period 
for Specified Diet 


FMC Normal Corn High-Lysine Corn F Valuescat) 
(n=7 Hens) (n=6 Hens) (n=7 Hens) 
Lysine consumed (g) 25e2) == 0162 Gisae=n Or? 10i3 == (0:7 314.2417 
Lysine utilized 
(g)” 21.6 + 0.4 Gi Se (Ohl iMOeY ==) (0.7) 311.672,17* 
(percent) ° So bes 15 88.4 1.0 99.2 + 0.1 51.240.17* 


* Denotes statistical significance, P<0.05. Those means underscored by the same line are not significantly 


different. 
® Standard errors. 


> Product of the 8-week consumption of lysine and the 5-week (Weeks 4-8) percentage utilization of lysine 


for each hen. 
© Utilization values for Weeks 4-8. 


94 


0.73, 18 df; P < 0.05) on the amount of 
lysine utilized. 


Protein vs. Lysine 


Significant differences existed in the 
quantitative utilization of both crude pro- 
tein and lysine by the juvenile hens on 
the three diets, which warranted a more 
definitive examination of the contribution 
of these two variables to the growth and 
maturation processes of pheasants. Hence, 
the influences of the quantitative utiliza- 
tion of crude protein and of lysine, ir- 
respective of diets, on the corresponding 
gain or loss in body weight of the juvenile 
hens were measured by multiple regres- 
sion analysis (Table 6). This analysis 
revealed, as previously demonstrated, that 
both the amount of crude protein utilized 
and the amount of lysine utilized, when 
considered separately, significantly influ- 
enced the body weights of juvenile hens 
in autumn. After accounting for crude 
protein, the amount of lysine utilized 
made a significant contribution to re- 
gression; however, the interjection of 
crude protein after accounting for lysine 
did not reveal a significant contribution 
to regression. Thus although the body 
weight of juvenile hen pheasants was 
significantly dependent on the utilized 
amounts of both crude protein and lysine 
when the two variables were considered 
singly, it was significantly dependent only 
on the utilized amount of lysine when 


Inuinors NaturaL History SurvEY BULLETIN 


Vol. 31, Art. 3 


the two variables were considered to- 
gether (Table 6). 


Glands and Organs 


The mean weights of gizzards, para- 
thyroid glands, adrenal glands, and kid- 
neys differed significantly among the hens 
fed diets of FMC, normal corn, or high- 
lysine corn (Table 7). These differences, 
except in the case of kidneys, also were 
evident when the weights of the glands 
or organs were expressed as percentages 
of body weight. 

The size of the adrenal glands, when 
expressed as an index percentage of body 
weight, was inversely correlated with the 
gain (or loss) in body weight of the juve- 
nile hens (Fig. 2), and thus adrenal size 
was generally greatest for hens fed nor- 
mal corn, intermediate for those fed high- 
lysine corn, and least for those fed FMC. 


Fat and Fatty Acids 

Although the deposits of fat, whether 
strip or visceral, did not differ statistical- 
ly among the juvenile hens fed the three 
diets, they were greatest for juvenile hens 
fed high-lysine corn, intermediate for 
those fed FMC, and least for those fed 
normal corn (Table 7). The accumula- 
tions of fat by hens fed high-lysine corn 
averaged nearly three times greater than 
accumulations of fat by hens fed normal 
corn. 

The distribution of the fatty acids con- 


Table 6.—Analysis of variance, as derived from multiple linear regression analysis, of the ef- 
fects of the utilized amounts of crude protein (g) and lysine (g) on the gains or losses in body 


weight (g) of juvenile hen pheasants in autumn. 


The null hypothesis is that the contribution to re- 


gression from X; is zero, where i=1 (crude protein), 2 (lysine). 

Source DF SS MS F 
Xi regression 1 27,974 27,974 10.73* 
X: regression|X: 1 16,515 16,515 6.33* 
Residual 17 44,318 2,607 

Total 19 88,807 

X. regression 1 44,392 44,392 17.03* 
X. regression|X.2 1 97 97 0.04 
Residual 17 44,318 2,607 

Total 19 88,807 


* Denotes statistical significance, P <:0.05. 


NutritionaAL RESPONSES OF PHEASANTS 


Lapisky & ANDERSON: 


July, 1973 


6971 =Ae ‘99 CI =As “898=—de “106=—Ap “SEOL=Ao “97 TE=—Aq “89° bE=—do 
9 = FOO! Oe Lo == sO Ge) eo “OL = OLL 96 + co oo = 06S sjeuarpy 
MOLE oat ee Lal Slee a “SO 58. a1 = 6G oe Ty sprosdyyereg 
61T + TIS LOR a Be Vol + L+9 c8 + G8E Sr + 66 “lL = €0S sprordyy, 
Ill = 09F 8S = $9E 88 = Ih £8 += 666 c& = 666 $8 += 096 sasnurAy ], 
(,01 X) suecreg sureISTIT 
F00';0 += £0°0 600'0 += 40°0 8000 + +00 0.0 + 960 100 + L¢°0 800 + T€0 uaalds 
c00 = 0GS°0 400 = 9F0 c0'0 = #90 lO + BE 060+ FE FO + EF sXoupry 
 cOMuaeaen a) FLO = 20e "E00 soe Tt Sal Obes 1a 0 + OFT SsGiUmssemc Ol pavzzip 
100 + <cl0 100 + &10 100 + &10 S00 + 60 400 + S60 G00 = 90'T SvaTOURd, 
90'0 = £9'T 800 = IGT 90'0 = 6FT 80 + 9cl OT + ctl L0 + Val JOAVT 
thO + 8c TT0 + 660 660 += GSL0 Sb + 601 OT + TE 6T + «a9 | EAP ELAN 
c0'0 += LT0 600 + £00 €0°0 + cI0 6G 0 += SHT clo + £90 8o0 + S0T drys eq 
90 + 96 OT + 946 90 = 692 08 = T06! 891 = F68l - 64 + OGIC SSTOSDMSEUeLS 
WysIaM Apog [vury Jo JUsIeg SUIvIL) 
XO) ite} 9) OWA (4=u) (g=u) (£=u) [eAS5} 9) 2X2) 
aursh']-y sty] JeUMON ul0yn ulor OWA ‘ursIOQ, 
autsd']-Ystyy [euLIONy ‘ansst J, 


“(GP ZL PUD Z #500 >d) 
juasayip AjjuD>2yIUBIs 91D aul] Jo puly auWOs ayy Aq PasOrsiapUN sUDaW “996| ‘G| Jaquiareq—OZ 192qG0j20 ‘poliad yaaM-g UD JO} UJOD auUIsAj-YyBIYy jo 40 ‘UO jOWJOU 
$0 ‘(DW4) Moy> ar2uDUajUIDW puUD j4YBIY D jo sjaIp aAIsnj>xa pa} syuDsDaYyd Usy ajiuaAn! Woy spuDjB puD ‘suDBio ‘sanssi, pajrajes yo syyBlaMm UDaW—'/ ajqd] 


Inuunors NATURAL 


roa) 


150 e 


+ =FMC (n=7) 
* = NORMAL CORN(n=6) 
© = HIGH LYSINE CORN(n=7) 


ADRENALS AS PERCENTAGE (X 107) OF BODY WEIGHT 


-50 {0} +50 


tained in the visceral fat of juveniles was 
almost the same for birds on diets of 
normal corn and high-lysine corn; the 
ratio of saturated to unsaturated fatty 
acids in the visceral fat was about 27:73 
for both corn diets (Table 8). The vis- 


History Survey BULLETIN 


+100 


Vol. 31, Art. 3 


Y=105.51-0.25X 
r=—0.49(P<0.05) 


— Relation- 
ship, as indicated by 
linear correlation, be- 
tween gain or loss in 
body weight (g) and 
the corresponding size 
of adrenal glands (ex- 
pressed as 10° percent- 
age of final body 
weight) for juvenile hen 
pheasants fed exclusive 
diets of either FMC, nor- 
mal corn, or high-lysine 
corn for the total 8- 
week feeding trial, Oc- 
tober 20-December 15, 
1966. 


Fig. 2s 


+150 
GRAMS GAIN OR LOSS IN BODY WEIGHT 


ceral fat of juvenile hens fed FMC con- 
tained a higher proportion of saturated 
fatty acids than did fat from hens fed 
either of the two corns, the principal dif- 
ferences being in the distribution of pal- 
mitic, stearic, and linoleic acids. 


Table 8.—Percentage distribution of fatty acids contained in the visceral fat of juvenile and adult 
hens fed exclusive diets of flight and maintenance chow (FMC), of normal corn, or of high-lysine corn. 


Values represent the mean 


of two replicated composite samples of visceral fat from hens in each age- 


diet group. 
Fatty Acids Juvenile Hens Adult Hens 
am Normal High-Lysine Normal  High-Lysine 
Visceral Fat FMC Corn Corn Corn Corn 
Saturated acids 
Lauric (12:0)" 1.0 0.9 0.6 0.8 0.5 
Myristic (14:0) 0.4 0.3 0.2 0.3 0.2 
Palmitic (16:0) 24.2 21.0 21.4 20.5 16.6 
Stearic (18:0) 8.2 4.8 5.2 6.2 5.8 
Subtotal 33.8 27.0 27.4 27.8 23.1 
Unsaturated acids 
Palmitoleic (16:1) 7.4 7.4 6.6 7.7 6.0 
Oleic (18:1) 37.8 37.6 39.6 39.1 41.8 
Linoleic (18:2) 17.8 27.9 25.8 22.4 28.3 
Linolenic (18:3) 1.0 > U) 3.0 U 
Arachidonic (20:4) 2.2 b 0.7 0.1 0.5 
Subtotal 66.2 72.9 WORT. 72.3 76.6 
Total (percentage) 100.0 99.9 100.1 100.1 99.7 


® The numbers preceding and following the colons represent the number of carbon atoms and the number of 


bonds, respectively. 
> Not isolated in analysis. 


July, 1973 Lasisky & ANDERSON: 


ADULT HENS 
Body Weight Changes 


Adult hen pheasants that were fed 
either normal corn or high-lysine corn at 
the restricted rate of 200 g each per week 
for a 7-week period in late winter and 
early spring suffered losses in body weight 
that averaged 5.8 g for those on normal 


NuTRITIONAL RESPONSES OF PHEASANTS 97 


corn and 65.5 g for those on high-lysine 
corn (Table 9). Both groups of hens 
suffered rather drastic losses in weight 
during the first week of the feeding trial. 
However, the hens fed normal corn es- 
sentially recovered their first-week loss in 
weight during the subsequent 6 weeks 
whereas those fed high-lysine corn con- 
tinued to lose weight throughout the 
remainder of the feeding trial (Fig. 3). 


Table 9.—Body weight statistics for adult hen pheasants fed exclusive diets of 200 g of normal 
corn or of 200 g of high-lysine corn per hen per week for a 7-week period, March 6~April 24, 1967. 


Mean Body Weight (g) or 
Weight Change for 
Specified Diet 


Normal Corn High-Lysine Corn F Values«ar) 
(n=6 Hens) (n=6 Hens) 
Initial weight (March 6) 907.3 = 40.6° 900:3pe) 2221 0.02: ,10 
Final weight (April 24) 901.5 + 38.6 834.8 + 26.3 2.031,10 
Weight change —5.8 = 25.9 —65.5 + 26.4 2.593, 10 


@ Standard errors. 


+20 


-60 


GRAMS GAIN OR LOSS IN BODY WEIGHT 
I 
3 


-80 


ee 4A ( { (9029) 
NORMAL CORN 


(n=6) 


HIGH-LYSINE 
CORN 
(n=6) 


( 8359) 


0 \ 2 
(MARCH 6 ) 


(APRIL 24) 


WEEKS OF STUDY 
Fig. 3—Mean change in body weight (g) by weekly periods among adult hen pheasants fed exclu- 


sive diets of normal corn or of lysine corn; the vertical lines transecting the means are standard errors. 
The mean initial and final body weights for each group are given at the left and right of the graph, 
respectively. Statistica.ly significant (P<0.05; 1 and 10 df) weight changes occurred between diets 
for only one of the seven successive weekly periods: 2 to 3 (F—= 6.14). 


98 


The changes in body weight recorded for 
the 7-week period for the six hens in 
each group were +73, +40, +40, —54, 
—60, and —68 ¢ for normal corn, and 
+16, —19, —39, —72, —126, and —153 g 


for high-lysine corn. 


Digestibility Coefficients 


With both corn diets restricted to 200 
g per hen per week, each hen consumed 
the total offering, and hence an identical 
amount of corn. The digestion of the 
two corns by the adult hens, however, 
differed significantly, averaging 81.3 per- 
cent for normal corn and 76.9 percent 
for high-lysine corn (Table 10). 


Calorie Utilization 


The number of calories consumed by 
the adult hens was similar for the diets 
of normal corn and high-lysine corn 
(Table 11). Yet, both the proportionate 
and absolute utilization of calories dif- 
fered significantly between the two groups 


Inurnois Natura History Survey BULLETIN 


Vol. 31, Art. 3 


of hens, being greater for those fed nor- 
mal corn than for those fed high-lysine 
corn. The adult hens metabolized 3,921 
(84.3 percent) and 3,617 (79.6 percent) 
kcal per kg of normal corn and high-ly- 
sine corn, respectively. 


Protein Utilization 


The intake of crude protein was simi- 
lar for the adult hens on each of the 
two corns because the corns were fed at 
identical rates and had similar protein 
contents (Tables 1 and 10). However, 


both the proportionate and absolute utili- 


zation of crude protein by the adult hens 
differed significantly between the two 
diets, with the efficiency of protein utili- 
zation being greater for hens fed normal 
corn than for those fed high-lysine corn 
(Table 10). 


Lysine Utilization 


The intake of lysine by hens fed high- 
lysine corn was about 42 percent greater 


Table 10.—Comparative utilization of exclusive diets of normal corn and of high-lysine corn, 
fed at a restricted rate of 200 g per hen per week, by adult hen pheasants for a 7-week period, 
March 6~April 24, 1967. All values expressed as dry weight. 


Mean Value per Hen per Week 
for Specified Diet 


Normal Corn 


High-Lysine Corn F Valuescaz) 


(n=6 Hens) 

Food consumed (g)” 200 
Crude protein? 

consumed (g) 24.0 
Lysine consumed (g)° 0.8 
Excreta (g)* 37.1 + 0.6° 
Digestibility 

coefficient (percent)? 81.3 + 0.2 
Crude protein in 

excreta (percent) 47.3 + 21 
Crude protein 

utilized (g)? 6.3 + 0.7 

(percent) 26.3 + 2.8 
Change in body 

weight (¢g) — 0.8 


(n=6 Hens) 

200 

23.4 
1.1 

45.0 = 1.6 18.85: ,10* 
76.9 + 0.5 54.07:,10* 
44.0 + 2.7 4.241 10 
3.6 + 0.7 8.641,10* 
15a = 19) 6.23110 

—9.4 


* Denotes statistical significance, P< 0.05. Interactions between weeks yielded no significant F values in any 


category. 


2 Each hen consumed the 200 g of food presented to it each week. 


> See Table 1 for protein content of food. 


© Product of crude protein consumed and amount of lysine in protein. 


4 Based on data from Weeks 5 and 7 only. 
¢ Standard errors. 


bho 


ys 


July, 1973 Lasisky & ANDERSON: 


NutriTIONAL RESPONSES OF PHEASANTS 99 


Table 11.—Comparative utilization of calories by adult hen pheasants fed a restricted diet of 200 
g of normal corn or 200 g of high-lysine corn per hen per week for a 7-week period, March 6— 


April 24, 1967. 


Mean Value per Hen per 7-Week Period 
for Specified Diet 


Normal Corn 


(n=6 Hens) 
Calories consumed (kcal) 6,512° 
Calories per g excreta” S905 = 262 
Calories utilized 
(kcal)? 5,492 + 16 
(percent)? 84.3 + 0.2 


* Denotes statistical significance, P< 0.05. 


High-Lysine Corn F Valuescar) 
(n=6 Hens) 
6,362 
4,121 + 28 3,897.40: ,10* 
5,066 + 55 17,095.46:,10* 
79.6 + 0.9 27.48:1,10 * 


¢ Each hen consumed the 200 g of food presented to it each week; hence, all hens on ecah of the two diets 
consumed the same number of kcal. See Table 1 for caloric content of corns. 


> Caloric values for Weeks 5 and 7. 
© Standard errors. 


7 Product of the 7-week consumption of calories and the 2-week (Weeks 5 and 7) percentage utilization of 


calories for each hen. 


than among those hens fed normal corn 
(Table 12), the difference being attribu- 
table to the different proportions of ly- 
sine contained in the two corns (Table 1). 
The difference in proportionate utiliza- 
tion of the lysine between the two corns, 
although not statistically significant, was 
slightly greater among hens fed the high- 
lysine corn. In the final analysis, the 
adult hens fed high-lysine corn utilized 
about 33 percent more lysine during the 
feeding trial than did those hens fed an 
equivalent amount of normal corn. The 
response in body weight of the adult hens 
was not related to the amount of lysine 
utilized during the 7-week trial (r = 0.56, 
10 df; P > 0.05). 


Glands and Organs 


Although the adult hens fed normal 
corn and those fed high-lysine corn ex- 
hibited striking differences in the sizes 
of their organs, glands, and tissue masses 
(Table 13), none of the differences were 
statistically significant; there was, how- 
ever, a pronounced pattern in the differ- 
ences. Either the mean weight or the 
percentage of body weight (or both) of 
sternal muscles, fat strip, visceral fat, 
liver, ovary (and ovum), oviduct, and 
thyroids was larger for hens fed normal 
corn than for those fed high-lysine corn. 
The parathyroids and adrenals, however, 
were larger among hens on the high-ly- 


Table 12.—Comparative utilization of lysine by adult hen pheasants fed a restricted diet of 200 
g of normal corn or 200 g of high-lysine corn per hen per week for a 7-week period, March 6— 


April 24, 1967. 


Mean Value per Hen per 7-Week Period 
for Specified Diet 


Normal Corn 


High-Lysine Corn F Valuescar) 


(n=6 Hens) (n=6 Hens) 
Lysine consumed (g) 5.4° Ted 
Lysine utilized 
(g)? 4.6 + 0.1 6.9 + 0.2 192.88, 10* 
(percent) ° 85.4 + 0.9% 89.0 + 2.0 2.761,10* 


* Denotes statistical significance, P < 0.05. 
2 All hens consumed the 200 fi 
group consumed the same amount of lysine. 


g of food presented to them each week; hence, each hen within each dietary 


> Product of the 7-week consumption of lysine and the 2-week (Weeks 5 and 7) percentage utilization of 


lysine for each hen. 
© Utilization values for Weeks 5 and 7. 
4 Standard errors. 


100 


Inurnors NAaTrurAL History Survey BULLETIN 


Vol. 31, Art. 3 — 


Table 13.—Mean weights of selected tissues, organs, and glands from two groups of adult hen — 
pheasants that were fed, respectively, normal corn or high-lysine corn for a 7-week period, March 6— — 
April 24, 1967. There were no significant differences between paired means (P>0.05; 1 and 10 df), 


Tissue Normal High-Lysine 
Organ, Corn Corn Normal High-Lysine 
or Gland (n=6) (n=6) Corn Corn 
Grams Percent of Final Body Weight 
Sternal muscles 20225) == 18 205.2 + 4.7 23.5 + 0.6 24.7 + 1.0 
Fat strip 2.99) Se) O57 1.46 + 0.45 0.32 + 0.06 0.17 + 0.05 
Visceral fat 95.3 ch 5. 134 +1416 2.79 + 0.63 1.53 + 051 
Liver 150: Silo ee 206 1.65 + 0.07 1.60 + 0.08 
Ovary 7.6.) 2:8 3.1 + 0.3 0.83 + 0.30 0.37 + 0.03 
Largest ovum 2.65 = 1.13 0.70 + 0.21 0.29 = 0.12 0.08 = 0.03 
Oviduct 116 + 1.4 Ih eal 1.27 = 0.13 0.93 + 0.16 
Milligrams Percent (X 10*) 
Thyroids 64.9 = 11.9 Oo: ==) 754 SEB 66.5 + 7.9 
Parathyroids 9:95 == 208 phe ak 9-8) ily Wi eo mee) 14:3 
Adrenals 99.0 = 45) | 055 2249 103.7, = 5.6 Gai 


sine diet than among those on the normal 
corn diet. This pattern of difference in 
weights of tissues, organs, and glands be- 
tween the normal corn diet and the high- 
lysine corn diet for adult hens was nearly 
the exact opposite of that observed among 
the juvenile hen pheasants. 


Fat and Fatty Acids 


The visceral fat from adult hens fed 
normal corn showed slightly greater 
amounts of saturated fatty acids and 
correspondingly lesser amounts of unsat- 
urated fatty acids than did that from hens 
fed high-lysine corn (Table 8). The ra- 
tios of saturated to unsaturated fatty acids 
in the visceral fat were about 28:72 for 
the adult hens fed normal corn and 


23:77 for those fed high-lysine corn. 


DISCUSSION 


The 8-week feeding trial with juvenile 
hen pheasants, which was conducted be- 
tween mid-October and mid-December, 
coincided with that period of the year 
during which wild hen pheasants of com- 
parable ages are gaining in body weight. 
To illustrate, the body weights of wild 
juvenile hens captured by nightlighting 
in east-central Illinois during the 6 years 
1956-1961 averaged 833 g (n=447) in 


October and 943 g (n=92) in December 
(R. F. Labisky, unpublished data), 
which represented gains of 110 g, or 13.2 
percent per hen during the 2 months. 

The juvenile hens in the present study 
posted 2-month changes in mean body 
weight of +13.8, +3.1, and —1.2 per- 
cent on exclusive, ad libitum diets of 
FMC, high-lysine corn, and normal corn, 
respectively (Table 2). Wild juvenile 
hens must supplement their corn-domi- 
nated autumn diets with food items more 
nutritious than corn to post the weight 
gains that they normally exhibit in au- 
tumn. However, as a staple food, high- 
lysine corn appears to be more favorable 
than normal corn in supplying the de- 
mands for growth by young pheasants, 
at least when both corns are available 
in an unrestricted supply. 

If not restricted in their food intake, 
yearling and adult hens usually gain 
weight throughout late winter and early 
spring, reaching their maximum weight 


in April just prior to the onset of egg-— 


laying (Kirkpatrick 1944:178; Breiten- 
bach et al. 


The hens in this study, when fed restrict- 
ed diets of corn during a 7-week span in 
March and April, averaged weight losses 
of 0.6.and 7.3 percent on diets of normal 


1963 : 25-26; Gates & Woeh- 
ler 1968: 235-238; Anderson 1972:467). 


July, 1973 


corn and high-lysine corn, respectively; 
their final body weights averaged 200- 
300 ¢ below normal weights reported for 
adult hens at the initiation of egg-laying 
by Breitenbach (1963:26), Labisky & 
Jackson (1969:720), and Anderson 
1972459). 

Any meaningful discussion of the 
physiological responses of pheasants to 
diets of normal corn and high-lysine corn 
must be prefaced by some knowledge of 
the nutritional requirements of the spe- 
‘cies. Unfortunately, data regarding nu- 
tritional requirements of pheasants are 
scarce, particularly for subadult birds, 
and must be extrapolated from informa- 
tion available for poultry. The National 
Research Council (1971:15-16) lists the 
minimum metabolizable energy (ME) 
requirements for domestic chickens (Gal- 
lus gallus) and turkeys (Meleagris gallo- 
pavo) as 2,900-3,095 kcal per kg of food 
for 14- to 20-week-old birds, and as 2,850 
kcal for mature breeders. Thus, the ME 
obtained by pheasants, both juveniles 
and adults, from the normal corn and 
high-lysine corn fed in this study well 
exceeded the energy levels required by 


Lapisky & ANDERSON: NUTRITIONAL RESPONSES OF PHEASANTS 


101 


poultry (Table 14). The FMC, however, 
only provided growing pheasants 2,837 
kcal of ME per kg of ration, slightly less 
than the minimum ME required by grow- 
ing poultry. However, Barrett & Bailey 
(1972:14, 16-17) recently reported that 
breeding pheasants can perform satis- 
factorily on diets containing about 2,500 
kcal of ME per kg of ration if the protein 
level is above 13 percent. 

The rates of metabolizability of both 
normal corn and high-lysine corn by ju- 
venile and adult hen pheasants in this 
study paralleled closely the general 80 
percent metabolizability rate of normal 
corn by chickens (Ewing 1963:83). How- 
ever, among pheasants, the metaboliza- 
bility of normal corn was slightly greater 
than for the high-lysine corn regardless of 
whether the corns were fed ad libitum to 
juveniles or in restricted quantities to 
adults (Tables 3 and 10). Also, ju- 
veniles metabolized proportionately more 
of each of the two corns than did adults, 
a difference that may have reflected the 
relative demands of growth. 

Our study showed that although the 
juvenile hen pheasants utilized a similar 


Table 14.—Levels of protein, metabolizable energy, and selected amino acids recommended for 
poultry feeds in relation to quantities supplied in diets fed to pheasants during this study. All levels, 
except those for metabolizable energy, are expressed as percentages of total diet. 


Dietary Requirements* 


Foods Supplied 14- to 22-Week-Old 
Pheasants in This Study 


Chickens: Turkeys: Chickens: Normal High-Lysine 
14-20 Weeks 14-20 Weeks Breeders FMC Corn Corn 
Crude protein 12.0 16.5 15.0 22.8 12.0 OS7, 
Metabolizable energy : 
(kcal/kg) 2,900 3,095 2,850 2,837° 3,945° (3,921) ° 3,808"(3,617)° 
Selected amino acids 
Arginine 0.72 1.0 0.8 1.42 0.46,0.457 0.79° 
Lysine 0.66 0.9 0.5 eal 0.38,0.18" 0.55 
Methionine 0.24 0.31 0.28 0.32 0.14,0.097 0.16° 
Cystine 0.21 0.21 0.25 0.23 0.14,0.097 0.20° 
Tryptophan 0.12 0.15 0.11 u 0.09,0.09% 0.15° 


“From National Research Council (1971:15—-16); these levels are recommended for achieving satisfactory 


dietary responses by poultry. 
° ME tor juvenile hen pheasants. 
© ME for adult hen pheasants. 


@ Amino acids not measured in this particular normal corn (Pioneer 3306). First value, except for lysine, 


from Cromwell et al. (1967:/0b) for normal corn containing 9.10 percent protein, and 


second value from Na- 


tional Kesearch Council (1971:28, 40) for normal corn (No. 2 yellow dent) containing 8.90 percent protein; 


values are probably conservative as regards Pioneer 3306. 


. £ Amino acids not measured in high-lysine corn. 
taining 11.60 percent protein. 
' Data not available. 


Value from Cromwell (1967:706) for opaque-2 corn con- 


102 


number of calories on all three diets— 
FMCG, normal corn, and high-lysine corn 
—the proportionate utilization of calor- 
ies was inversely associated with caloric 
consumption (Tables 4 and 11). Barrett 
& Bailey (1972:20), however, reported 
that breeder pheasants compensated for 
foods with low ME levels by increasing 
food consumption, and consequently 
maintained reasonably similar levels of 
caloric intake on diets containing from 
2,100 to 3,400 kcal of ME per kg. This 
type of compensatory action did not oc- 
cur among juvenile hen pheasants fed 
different diets in our study. The diets of 
these birds, however, were more variable 
in ME, protein content, and amino acid 
patterns than the rations fed by Barrett 
and Bailey, and hence are not totally 
comparable. 

In our study, the similarity in caloric 
utilization by juvenile pheasants among 
the diets characterized by different levels 
of ME was achieved not by compensatory 
caloric intake but by compensatory me- 
tabolizability. Furthermore, adult hens 
did not show a greater proportionate uti- 
lization of calories from the corn than 
did juveniles, even though they were fed 
a restricted ration and consumed fewer 
calories than they would have consumed 
on an ad libitum diet of corn. 

The National Research Council (1971: 
19) listed the dietary protein require- 
ments for starting and growing pheasants 
at 30 percent. Dale & DeWitt (1958: 
292) reported that the growth rates of 
young pheasants, to 10 weeks of age, 
were less on diets containing 15, 18, and 
22 percent protein than on diets contain- 
ing 28 percent protein. The Council 
(1971:15-16) also listed the protein re- 
quirements of the chicken as 20-23 per- 
cent for chicks, 12-16 percent for growing 
chickens, and 15 percent for laying 
(breeding) chickens; comparable levels 
for domestic turkeys were 28, 14-20, and 
14 percent, respectively. The reported 
protein content in the diets of wild hen 
pheasants in the Midwest ranged season- 
ally from a minimum of about 12 percent 


Ituinors NaturAL History SurvEY BULLETIN 


1 


Vol. 31, Art. 3— 


to a maximum of about 19 percent 
(Korschgen 1964: 169, 174). Collectively, — 
these findings suggest that the protein — 
needs of pheasants are probably satisfac- 
torily met at levels of 16-20 percent for 
juveniles older than 14 weeks and 15 © 
percent for adult breeders. Therefore, in ~ 
this study, the dietary protein levels for 
pheasants were sufficient in the FMC ~ 
(22.9 percent), but insufficient in both 
normal corn (12.0 percent) and high- 
lysine corn (11.7 percent). | 

Whereas consumption of crude protein — ; 
by the juvenile hen pheasants differed — 
among birds fed FMC, normal corn, and 
high-lysine corn (Table 3), the propor- f 
tionate utilization of the protein con- 
sumed, irrespective of amount, was simi-— 
lar on all diets. Thus, among juvenile 
pheasants the total amount of protein 
utilized was related directly to the amount ~ 
consumed—a situation opposite that for 
caloric utilization. 

Although adult hens consumed similar 
amounts of protein from the two corns, 
which were fed at a restricted rate, they 
utilized 41 percent less of the protein from 
high-lysine corn than from normal — 
corn (Table 10). Thus, although both 
corns yielded dietary protein levels that 
were unsatisfactory to juveniles and 
adults, the pheasants still failed to utilize 
about three-fourths of all the protein they 
consumed in corn. 

Eleven of the 23 verified amino acids 
in proteins are essential to birds; that is, 
they cannot be sufficiently synthesized by 
the bird and must be taken in via the diet. 
Ewing (1963:201, 203) points out that 
arginine, lysine, methionine, cystine, and 
tryptophan are particularly important 
to birds because they are essential amino 
acids that are in critical demand during 
avian growth and development; the other 
amino acids are either synthesized by the 
bird or are present in ample quantities 
in most foods. j 

Important to the understanding of the 
amino acid complex is the fact that a 
deficiency of any essential amino acid will 


not only reduce the utilization of other 


July, 1973 


amino acids, but will also reduce the utili- 
zation of the entire diet. Thus, although 
this paper is concerned principally with 
the growth-associated amino acid, lysine, 
other amino acids that are potentially 
important to pheasants cannot be ignored. 
The FMC ration provided to pheasants 
in this study offered adequate quantities 
of protein and the essential amino acids 
for both subadult and adult birds (Table 
14). The normal corn and high-lysine 
corn diets, while providing minimal 
amounts of protein, did not supply ade- 
quate quantities of amino acids. The 
amino acids most lacking in the corns 
were lysine and methionine. High-lysine 
corn, however, offered an amino acid 
profile superior to that of normal corn, 
and the profiles of both corns were more 
aligned with the requirements of adult 
birds than of growing pheasants. 

The importance of lysine to growth 
was well illustrated in a study by Baldini 
et al. (1953:946-948). They demon- 
strated that young bobwhites (Colinus 
virginianus), which reportedly required 
diets with 28 percent protein, could sur- 
vive and grow well on diets containing 
as little as 20 percent protein as long 
as the diets contained adequate amounts 
of lysine. In their experiments, the addi: 
tion of 0.3 percent lysine to a base diet 
of 20 percent protein and 1.0 percent 
lysine produced a ration with growth and 
survival qualities for bobwhites that were 
equal to those provided by a diet contain- 
ing 28 percent protein and 1.0 percent 
lysine; thus, 0.3 percent lysine essentially 
replaced 8.0 percent crude protein. Such 
findings offer support for our conclusion 
that the amount of lysine utilized by ju- 
venile pheasants contributed more signif- 
icantly to their growth than. did the 
amount of crude protein utilized (Table 
6). 

As with crude protein, pheasants ex- 
hibited no compensatory utilization of 
lysine; the utilization of lysine was related 
directly to its intake for both juvenile and 
adult birds (Tables 5 and 12). Perhaps 


the most interesting observation was the 


Laspisky & ANDERSON: NutTRITIONAL RESPONSES OF PHEASANTS 103 


extremely high utilization (99.2 percent) 
of lysine from high-lysine corn by juvenile 
pheasants. This rate of utilization was 
not maintained by adult hens. 

The role of inadequate nutrition— 
quantitative or qualitative—as regards the 
physiology of stress in vertebrates is poor- 
ly understood. Among mammals, the 
term “stress” has become almost synony- 
mous with increased adrenocortical ac- 
tivity (see review by Christian 1963). Pre- 
sumably, some adverse stimulus triggers, 
via the hypothalamus, an increased re- 
lease of adrenocorticotropic hormone 
(ACTH), which in turn results in the 
increased production and secretion of 
corticosteroids from the adrenal cortex 
that are necessary for maintaining physi- 
ological homeostasis under the given 
stress. (Prolonged exposure by the ani- 
mal to an adverse stimulus may result 
in exhaustion of the adrenal cortex, the 
subsequent failure of corticosteriod pro- 
duction, and finally death.) To produce 
the additional corticosteriods, the adrenal 
cortex undergoes hyperplasia and hyper- 
trophy—hence, enlargement of the gland. 
Thus, enlarged adrenals have become 
generally recognized as clinical evidence 
of acute or chronic distress in mammals. 

Whether enlarged adrenals are a mea- 
sure of stress in birds is not clear. Like 
Christian & Davis (1966:11-13), who 
found a direct relationship between 
adrenal size (of mature females) and 
population density for vole ( Microtus 
pennsylvanicus) (Neave & Wright 1968: 
634) reported a positive correlation be- 
tween adrenal-weight indices and popu- 
lation density for ruffed grouse (Bonasa 
umbellus). Breitenbach et al. (1963:34) 
reported that the adrenals of adult hen 
pheasants that were restricted in their 
food intake (45 g per day) did not hy- 
pertrophy; however, the adrenals of in- 
dividual hens, in noticeably poor condi- 
tion, exhibited a marked increase in size. 
Also, Newlon et al. (1964:538-539) ob- 
served that the adrenal weights of bob- 
whites were greatest for birds fed those 
foods which yielded the poorest perform- 


104 Iruinois Natura History Survey BULLETIN 


ance in maintaining body weight. These 
observations, coupled with our findings 
that the adrenal weights of hen pheasants 
were inversely related to changes in body 
weight (Table 7 and 13; Fig. 1-3), sug- 
gest to us that enlarged adrenals offer 
diagnostic symptoms of the stresses of 
inadequate nutrition in pheasants, and 
possibly other birds. 

The deposition and mobilization of 
depot fats are dynamic processes—even in 
an animal in reasonably stable energy 
balance (White et al. 1968:500). We 
found that depot fat, both strip and vis- 
ceral, was greatest for juvenile hen pheas- 
ants fed high-lysine corn, intermediate 
for those fed FMC, and least for those 
fed normal corn (Table 7). In contrast, 
adult hens, fed restricted but equal 
amounts of the two corns, accumulated 
greater fat deposits on a diet of normal 
corn than on a diet of high-lysine corn 
(Table 13). The fat deposits from these 
adult hens, irrespective of the type of 
corn diet, were many times smaller than 
those reported in spring for confined 
hens fed a high-protein ration ad libitum 
(Breitenbach 1963:32) or for wild hens 
(Anderson 1972:461). Breitenbach et 
al. (1963:34) presented evidence that 
the storage of fat may be stimulated by 
increased amounts of adrenocorticoster- 
oids. Hence, if the production of corti- 
costeroids paralleled increased adrenal 
size, as would be expected, fat deposits 
should have been related directly to 
adrenal size. We did not observe this 
relationship among the hens in this study. 
The birds’ depot fats, which represent 
their largest reservoir of energy, were re- 
lated to body weight, and therefore in- 
versely reflected the hens’ day-to-day 
energy demands. 

Depot fat consists chiefly of triglycer- 
ides; fatty acids, both saturated and un- 
saturated, are hydrolized from triglycer- 
ides via the action of the lipases. We 
found that the ratios of saturated to un- 
saturated fatty acids in the visceral fat 
from juvenile and adult hen pheasants 
fed normal corn and from juvenile hens 


were decreasing in body weight, and 


Vol. 31, Art. 3 — 
fed high-lysine corn were about 27:73 
(Table 8). Correspondingly, normal — 
corn and high-lysine corn contained sat- — 
urated to unsaturated fatty acid ratios of — 
14:86 and 19:81, respectively (Table 1).— 
Hence, there was some disparity in the — 
distribution of saturated and unsaturated — 
fatty acids between the depot fat of | 
pheasants and their corn diets. Although — 
the distribution of fatty acids in the depot — 
fat of herbivorous galliform birds gener- ' 
ally reflects the composition of the diet — 
(Moss & Lough 1968:559; West & Meng — 
1968b:438), West & Meng (1968a:539) — 
have also provided evidence that, at least — 
for the redpoll (Acanthis flammea), en-— 
vironmental conditions and the physio- 
logical state of the bird also influence the 
fatty acid composition of the visceral fat. 
Interestingly, the ratio of saturated to’ 
unsaturated fatty acids in the visceral fat — 
of the adult hens fed the restricted intake 
of high-lysine corn was 23:77, which 
represented an increase in unsaturated 
fatty acids over the 27:73 ratio recorded © 
for juvenile hens fed either high-lysine — 
corn or normal corn ad libitum and for — 
adult hens fed the restricted diet of nor-— 
mal corn. The shift by adult hens fed — 
high-lysine corn to a fatty acid composi- 
tion of depot fat that more closely re- 
flected that of their high-lysine corn diet ~ 
is not surprising because White et al. 
(1968:499) reported that fatty acid pro-~ 
files of depot fat resemble the dietary 
profiles more closely when the depot fat — 
is being depleted. The adult hens fed 
high-lysine corn at the restricted rate 


ra 


therefore probably drawing on the stores 
of saturated fatty acids for reserve energy. 
Under these conditions the replacement 
fatty acids reflected the high proportion 
of unsaturates in the high-lysine corn diet.” 
Plant seeds abound in unsaturated fat-— 
ty acids. The principal unsaturated fat- 
ty acids in the corns were oleic and lino-— 
leic. Correspondingly, the principal fatty” 
acids in the visceral fat of pheasants were 
oleic and linoleic (Table 8); however, 
the samples of visceral fat contained pro- 


July, 1973 Lasrsky & ANDERSON: Nurtritionat RESPONSES OF PHEASANTS 105 


portionally more oleic acid and consider- 
ably less linoleic acid than either of the 
corns. Linoleic was the principal fatty 
acid in the depot fats of the heather-eat- 
ing red grouse (Lagopus lagopus scoticus) 
(Moss & Lough 1968:560-561) and of 
the willow-eating willow ptarmigan (La- 
gopus lagopus alascensis) (West & Meng 
1968b:438). However, as we found for 
the pheasant, Walker (1964:63-64) re- 
ported the predominant fatty acid in the 
depot fat of the seed-eating bobolink 
(Dolichonyx oryzivorus) to be oleic acid. 
Oleic acid and linoleic acid seem to be 
the principal unsaturated fatty acids that 
characterize the depot fats of granivorous 
and herbivorous birds, respectively. 
The parathyroid glands secrete a hor- 
mone that functions importantly as a 
regulator of calcium, and probably phos- 
phorus, metabolism; high and low levels 
of circulating calcium act on the glands 
to inhibit or stimulate, respectively, se- 
cretion of the parathyroid hormone 
(Geschwind 1961:434-436). Hypertro- 
phied parathyroid glands and reduced 
levels of blood calcium are characteristic 
responses of laying chickens to low-calci- 
um diets (Bloom et al. 1960:207). In 
our study, we fed juvenile hen pheasants 
diets that ranged from about 14,000 ppm 
calcium for FMC to about 40 ppm calci- 
um for the corns (Table 1); corns in 
general are notoriously low in calcium 
content. The parathyroids from these 
juvenile hens fed normal corn and high- 
lysine corn weighed at least twice as 
much as the glands from those fed FMC 
|(Table 7), and nearly twice as much as 
\the glands from their wild counterparts 
| (Anderson 1972:485). Furthermore, the 
jadult hens fed exclusive diets of the corns 
\during late winter and early spring (Ta- 
|ble 13) had parathyroids substantially 
jlarger than those reported by Anderson 
|(1972:485) for wild adult hens at a 
\comparable time of the year. The hyper- 
\trophied parathyroid glands from hen 
\pheasants fed exclusive diets of calcium- 
‘deficient corns constituted strong clinical 
jevidence that the birds were suffering 
from a negative calcium balance. 


The gonadal recrudescence among 
pheasants in spring is a response, medi- 
ated via the hypothalamo-hypophyseal 
axis, to increasing photoperiod (Bisson- 
nette & Csech 1936:106; Hiatt & Fisher 
1947:538, 543; Greeley & Meyer 1953: 
353-354). In Illinois, complete gameto- 
genesis among hens, as evidenced by egg- 
laying, is attained between late March 
and mid-April (Labisky & Jackson 1966: 
382; Labisky 1968:69; Labisky & Jack- 
son 1969:719). In our study, none of 
the adult hens fed corn diets restricted 
to an intake of 200 g per week had initi- 
ated egg-laying at the conclusion of the 
experiment on April 24. Furthermore, 
the reproductive tracts of these hens, 
when compared to hens on unrestricted 
diets or in the wild (Breitenbach et al. 
1963:29; Anderson 1972:484) were se- 
verely underdeveloped physiologically for 
late April. The lag in ovarian and ovi- 
ducal development was more pronounced 
for hens fed high-lysine corn than for 
those fed normal corn (Table 13). 

The findings from this and previous 
investigations of confined pheasants (Ger- 
stell 1942:68; Kozlik 1949:62; Breiten- 
bach et al. 1963:27; Gates & Woehler 
1968: 240) have demonstrated that delays 
in egg-laying are related to poor physical 
condition in spring; Edwards et al. 
(1964:278) hypothesized a similar situa- 
tion for wild pheasants. Also, poor physi- 
cal condition, usually the result of mal- 
nutrition, signifies reduced reserves of 
energy. Fisher (1967:121) cited evidence 
to show that domestic hens would cease 
egg production as soon as their protein 
reserves were exhausted after being 
placed on a protein- or amino acid-defi- 
cient diet. Corns do not abound in pro- 
tein, and are deficient in one or more of 
the essential amino acids. 

The inhibitory effects of inadequate 
nutrition on reproduction in galliform 
birds, however, seem to be mediated 
through the hypothalamo-hypophyseal 
axis and not directly by protein or 
amino acid imbalances. Morris & Nal- 
bandoy (1961:687) demonstrated that 
undernourished or starved domestic pul- 


106 


lets failed to produce sufficient gonado- 
tropins to promote functional ovarian and 
oviducal development. Gates & Woehler 
(1968:243) sum it up aptly: “It would 
appear that restoration of body condition 
[in pheasants] . . . is a requisite for re- 
crudescence, normal rates of egg laying 
being delayed until energy is available 
for seeded and the secretory integ- 
rity of the pituitary is restored.” 


The nutritional responses of pheasants 
to normal corn and high-lysine corn are 
not consistent for birds of different ages. 
The weight gains and physiological par- 
ameters for juvenile hen pheasants fed 
high-lysine corn in autumn were distinct- 
ly more favorable than for those corres- 
pondingly fed normal corn. Yet adult 
hens fared far better on normal corn 
as an emergency food in late winter than 
they did on high-lysine corn. Thus, al- 
though potentially of benefit to growing 
pheasants, high-lysine corn may well be 
a detriment to mature pheasants. These 
conflicting results are at least partially 
explainable. The greater nutritional val- 
ue to young pheasants of high-lysine corn, 
when contrasted with normal corn, un- 
doubtedly lies in its higher content of 
the growth-oriented amino acid, lysine. 
Our findings indicated strongly that the 
weight gains of juvenile pheasants were 
dependent directly on the amount of ly- 
sine utilized. Evidence in support of 
these findings is provided by the research 
of Cromwell et al. (1967:711), who 
studied the nutritional responses of do- 
mestic chicks to the two corns: “. 
the beneficial effects of opaque-2 corn 
over normal corn appeared to be medi- 
ated solely through its higher lysine con- 
tent.” These workers, however, fed the 
corns as a part of nutritionally balanced 
basal diets, and not as exclusive food 
items. They also pointed out (p. 712) 
that the beneficial responses exhibited by 
young animals to opaque-2 corn would 
probably be much less for mature animals 
that have a lower protein requirement. 
Furthermore, the probability of selecting 


Inuinots Natura History SurvEY BULLETIN 


genetically for a pheasant that responds to — 


higher than normal dietary levels of ly- 
sine by exhibiting an improved rate of 
growth seems remote, as Godfrey (1968: 
1565) found that the heritability for ly- 
sine utilization among Japanese quail 
(Coturnix coturnix japonica) was very 
low. Thus opaque-2 corn apparently is 
not a panacea for assuring adequate nu- 
trition in pheasants. Another modified- 
protein corn called floury-2, which con- 


Vol. 31, Art. 3 
; 
j 


tains higher concentrations of both lysine ~ 


and methionine than those in most other 
corns (Nelson et al. 1965:1470; Crom- 
well et al. 1968:846), may offer nutrition- 
al benefits for birds that are potentially 
normal corn or opaque-2 corn. 4 

Corn, an important food source to — 
many species of wildlife, is a nutritive sta- _ 
ple for pheasants. To illustrate, Newlon — 
et al. (1964:536-537), in evaluating — 
foods for sustaining bobwhites, reported — 
that the mean survival duration for ju- — 
venile and adult birds fed an exclusive, 
ad libitum diet of normal corn in No-— 
vember was 22.6 days; no bobwhite sur-~ 
vived the 38-day feeding trial. As ob- : 
served in this study, however, exclusive - 
ad libitum diets of normal corn sustained © 
juvenile pheasants, with only minor 
weight losses, for 8 weeks in autumn with-_ 
out any mortality. Also, adult hens fed — 
a restricted intake of normal corn (200 
g per week) in late winter and early 
spring maintained their body weights for 
a 7-week feeding trial. If, however, 
pheasants are to parallel the annual cycle 
of body weights that normally character-— 
ize thriving populations in the wild they 
must. supplement their corn-dominated 
diets with food items more nutritiously 
balanced than high-lysine or normal corn. 
A plausible hypothesis, therefore, is that 
the high rates of nonhunting mortality 
observed among wild juvenile pheasants — 
in Illinois during autumn (R. F. Labisky, _ 
unpublished data) may be directly or in- 
directly related to nutritive imbalances ~ 
resulting from the surging availability of 
waste corn in the birds’ diet. 


ee eye 


Be 


superior to those provided by either — 


} 


July, 1973 


SUMMARY 


Corn is an important food for many 
wild animals and is especially prominent 
in the diet of wild pheasants in the United 
States. Yet despite its importance as a 
staple food of pheasants, knowledge of 
its nutritional value to the species is still 
quite limited. In 1963, scientists added 
another dimension to corn nutrition by 
discovering a modified-protein corn, 
opaque-2 corn (herein called high-lysine 
corn), which has substantially greater 
amounts of lysine in its endosperm than 
does normal corn. Lysine is one of the 
essential growth-promoting amino acids. 


The objectives of this investigation 
were to ascertain the physiological re- 
sponses of juvenile hen pheasants in au- 
tumn, and of adult hens in late winter 
and early spring to exclusive diets of 
normal corn and high-lysine corn. In 
the 8-week feeding trial for juveniles, 
October 20—December 15, 1966, 21 hens 
in three groups of 7 each were fed ex- 
clusive, ad libitum, diets of a balanced 
ration (FMC), normal corn, or high- 
lysine corn. Analyses of the three foods 
yielded the following: FMC—4.28 kcal 
per g, 22.8 percent protein, and 4.9 per- 
cent lysine in protein; normal corn—4.65 
kcal per g, 12.0 percent protein, and 3.2 
percent lysine in protein; and high-lysine 
corn—4,54 kcal per g, 11.7 percent pro- 
tein, and 4.7 percent lysine in protein. 
The feeding trial for juveniles coincided 
with the season in which juvenile hens, 
in contrast to adult hens, suffer dispro- 
portionately high rates of nonhunting 
mortality in Illinois, and also simultane- 
ously with the time that waste corn from 
the harvest suddenly becomes an abun- 
dant food source. Hence this phase of 
the study was designed partially to de- 
termine if juvenile mortality among wild 
pheasants was related to unbalanced nu- 
trition. In the 7-week feeding trial for 
adult hens, March 6—April 24, 1967, 12 
hens in two groups of 6 each were fed 
exclusive diets of normal corn or high- 
lysine corn at the restricted rate of 200 


Lasisky & ANDERSON: NutriTIONAL RESPONSES OF PHEASANTS 


107 


g per bird per week. This restricted in- 
take of food was intended to simulate 
the conditions that wild hens in the 
Midwest often confront in late winter 
and early spring. 

The juvenile hen pheasants fed ex- 
clusive diets of FMC and _ high-lysine 
corn for the 8-week period in autumn 
posted gains in body weight that aver- 
aged 98.4 g (13.8 percent) and 23.4 g 
(3.1 percent), respectively; correspond- 
ingly those juvenile hens fed normal corn 
suffered average losses of 8.7 g (1.2 per- 
cent). Wild juvenile hens averaged 
gains of 110 g (13.2 percent) for the 
comparable autumn period. 

Adult hens, each fed a restricted in- 
take of 200 g of corn per week for the 
7-week period in late winter and early 
spring, averaged losses in body weight of 
5.8 g (0.6 percent) for normal corn and 
65.5 g (7.3 percent) for high-lysine corn. 
Whereas hen pheasants usually exhibit 
gains in body weight in late winter, 
reaching their maximum weight just 
prior to the onset of egg-laying (usually 
in April), the adult hens on the restrict- 
ed intake of corn averaged 200-300 g 
below the normal body weights of wild 
hens in April. 

The kcal of energy metabolized per kg 
of food consumed by juvenile hens was 
2,837 (66.3 percent efficiency) for FMC, 
3,945 (84.8 percent) for normal corn, 
and 3,808 (83.3 percent) for high-lysine 
corn; adult hens metabolized 3,921 kcal 
per kg (84.3 percent) of normal corn 
and 3,617 kcal per kg (79.6 percent) of 
high-lysine corn. Juvenile hens, despite 
the differences in the yield of metaboli- 
zable energy among the foods, utilized 
the same number of calories on all three 
diets; the similarity in caloric utilization 
was achieved by compensatory metaboliz- 
ability and not by compensatory calor- 
ic intake. 

Unlike the situation for calories, hen 
pheasants exhibited no compensatory 
utilization of either crude protein or ly- 
sine. The total amounts of both protein 
and lysine utilized by hen pheasants 


108 


were related directly to dietary intake. 
Even though the dietary protein levels 
of both corns were unsatisfactory, the 
birds utilized only about one-fourth of 
the protein they consumed. The propor- 
tionate utilization of lysine by either 
adult or juvenile hens exceeded 85 per- 
cent for all diets. Interestingly, juvenile 
hens utilized 99 percent of the lysine 
consumed from high-lysine corn—a rate 
not achieved by adult hens. Most im- 
portant was the finding that the body 
weights of juvenile pheasants in autumn 
were more dependent on the amount of 
lysine utilized than on the amount of 
crude protein utilized. 

The adrenal weights of hen pheasants 
were inversely related to changes in body 
weight, which in turn was a reflection 
of the consumption of diets of different 
nutritional offerings. These findings in- 
dicated that enlarged adrenal glands in 
pheasants, and possibly other avian 
species, may offer diagnostic evidence for 
detecting the physiological stresses of un- 
balanced or inadequate nutrition. 

Depot fats of hen pheasants were di- 
rectly related to changing body weight, 
a relationship that reflected changing 
demands for energy for growth or main- 
tenance. The ratio of saturated: unsatur- 
ated fatty acids in the visceral fat of 
hens fed corn was about 1:3, which was 
greater than-that found in the corns. 
Oleic and linoleic were the principal 
fatty acids in corn, and, correspondingly, 
the most prominent in the depot fats of 
pheasants. 

Hypertrophied parathyroid glands 
from hen pheasants fed exclusive diets 
of calcium-deficient corns offered strong 
clinical evidence that wild hen pheasants 


Inurnois NATuRAL History Survey BULLETIN 


Vol. 31, Art. 3 


= 


on corn-dominated diets would suffer 
from a negative calcium balance. | 


The reproductive tracts of adult hen 
pheasants fed restricted diets of normal 
corn or high-lysine corn, unlike those of 
wild hens or hens fed unrestricted diets, 
were severely underdeveloped in late 
April. The lag in ovarian and oviducal 
development was substantially greater 
for hens fed high-lysine corn than for 
those hens fed normal corn. The effects — 
of inadequate nutrition are apparently 
mediated through the Bem e | 
pophyseal-gonadal axis. 


The nutritional responses of young 
and adult hen pheasants to normal corn ~ 
and high-lysine corn were not similar, — 
The physiological profiles of juvenile 
hens fed high-lysine corn in autumn were 
distinctly more favorable than of those 
fed normal corn. The greater nutritional 
value to young pheasants of high-lysine — 
corn, in contrast to normal corn, very 
likely is associated with its higher content 
of lysine, an essential growth-promoting 
amino acid. However, as an emergency 
food for adult hens in late winter or early 
spring, normal corn proved superior to- 
high-lysine corn. 


To attain the physiological plateaus 
that normally characterize self-maintain- — 
ing populations in the wild, pheasants 
must supplement their corn-dominated 
diets with foods more nutritiously bal-— 
anced than corn—high-lysine corn not” 
excepted. Dietary imbalances, resulting; 
from the surging availability of waste» 
grain from the corn harvest, may be» 
associated with the high rates of non-— 
hunting mortality among juvenile hen) 
pheasants in Illinois during autumn. 


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Gescuwinp, Irvine I. 1961. Hormonal con- 
trol of calcium, phosphorus, iodine, iron, 
sulfur, and magnesium metabolism, p. 387- 
472. In C. L. Comar and Felix Bronner 
(Editors), Mineral metabolism, Volume 1, 
Part B. Academic Press, New York and 
London. P. 387-879. 


Goprrey, Epwarp F. 1968. Ten generations 
of selection for lysine“ utilization in Japan- 
ese quail. Poultry Science 47(5):1559- 
1566. 

GREELEY, FREDERICK, and RoLtanp K. MEYER. 
1953. Seasonal variation in testis-stimulat- 
ing activity of male pheasant pituitary 
glands. Auk 70(3) :350-358. 

Harper, ALFRED E. 1967. Effects of dietary 
protein content and amino acid pattern on 
food intake and preference, p. 399-410. In 
Handbook of physiology, Section 6 (Charles 
F. Code, Editor): Alimentary canal, Vol- 
ume 1. Control of food and water intake. 
American Physiological Society, Washing- 
ton, D.C. 459 p. 

Hiatt, Ropert W., and Harvey I. FIsHer. 
1947. The reproductive cycle of ring-necked 
pheasants in Montana. Auk 64(4) :528- 
548. 

Hupson, Georce E., and Patricia J. LAN- 
ZILLOTTI. 1964. Muscles of the pectoral 
limb in galliform birds. American Midland 
Naturalist 71(1):1-113. 


109 


110 


Jensen, A. H., D. E. Becker, and B. G. 
Harmon. 1967. Opaque-2 corn, milo and 
wheat in diets for finishing swine. Journal 
of Animal Science 26(6):1473 (Abstract). 

Kirkpatrick, C. M. 1944. Body weights 
and organ measurements in relation to age 
and season in ring-necked pheasants. Ana- 
tomical Record 89(2):175-194. 

Korscucen, Leroy J. 1964. Foods and nu- 
trition of Missouri and midwestern pheas- 
ants. North American Wildlife and Natural 
Resources Conference Transactions 29:159- 
180. 

Kozurx, Frank M. 1949. Pheasant-quail 
management research: pheasant section. 
Wisconsin Wildlife Research 8(1) :51-64. 

Lapisky, RoNALD F. 1968. Ecology of pheas- 
ant populations in Illinois. Ph.D. Thesis. 
University of Wisconsin, Madison. 511 p. 

—, and Gary L. Jackson. 1966. Char- 

acteristics of egg-laying and eggs of yearling 

pheasants. Wilson Bulletin 78(4) :379-399. 

, and 1969. Production and 
weights of eggs laid by yearling, 2-, and 3- 
year-old pheasants. Journal of Wildlife 
Management 33(3) :718-721. 

Mertz, Epwin T. 1966. Growth of rats on 
opaque-2 maize, p. 12-18. In Edwin T. 
Mertz and Oliver E. Nelson (Editors), 
Proceedings of the high lysine corn confer- 
ence. Corn Industries Research Founda- 
tion, Corn Refiners Association, Inc., Wash- 
ington, D.C. 186 p. 

, Lynn S. Bates, and Ouiver E. 

Netson. 1964. Mutant gene that changes 

protein composition and increases lysine 

content of maize endosperm. Science 145 

(3629) :279-280. 

, Orrvia A. VERNON, Lynn S. BATEs, 
and Otiver E. Netson. 1965. Growth of 
rats fed on opaque-2 maize. Science 148 
(3678) : 1741-1742. 

Morris, T. R., and A. V. NaLBanpov. 1961. 
The induction of ovulation in starving pul- 
lets using mammalian and avian gonadotro- 
pins. Endocrinology 68(4) :687-697. 

Moss, R., and A. K. Loucu. 1968. Fatty 
acid composition of depot fats in some game 


birds (Tetraonidae). Comparative Bio- 
chemistry and Physiology 25(2) :559-562. 
NaTionAL ReseARCH Councit. 1971. Nu- 


trient requirements of domestic animals. 


Inurnors NaturaL History Survey BuLLETIN 


Vol. 31, Art. 3 


Number 1, Nutrient requirements of poul- 


try. Sixth edition (revised). National | 
Academy of Sciences, Washington, D.C. . 
54 p. i 


Neave, Davi J., and Bruce S. WRIGHT. | 
1968. Ruffed grouse adrenal weights re- 
lated to population density. Journal of 
Wildlife Management 32(3) :633-635. 

NEtson, Otiver E., Epwin T. Mertz, and 
Lywn S. Bates. 1965. Second mutant gene + 
affecting the amino acid pattern of maize » 
endosperm proteins. Science 150(3702): 
1469-1470. a 

NEWwLOon, CuHartes F., THomas S. BASKETT, 
Rospert P. BrEITENBACH, and Jack A. 
Stanrorp. 1964. Sustaining values of 
emergency foods for bobwhites. Journal of 
Wildlife Management 28(3) :532-542. a 

Pickett, Ricwarp A. 1966. Opaque-2 corn } 
in swine nutrition, p. 19-22. In Edwin T. ° 
Mertz and Oliver E. Nelson (Editors), Pro- - 
ceedings of the high lysine corn conference. — 
Corn Industries Research Foundation, Corn 
Refiners Association, Inc., Washington, D.C 
186 p. 

Rocier, Joun C. 1966. A comparison of | 
opaque-2 and normal corn for the chick, 
p. 23-25. In Edwin T. Mertz and Oliver 
E. Nelson (Editors), Proceedings of the 
high lysine corn conference. Corn Indus- 
tries Research Foundation, Corn Refiners 
Association, Inc., Washington, D.C. 186 p. 

WALKER, ALMA TOERS. ; 
acids in migratory bird fat. ; 
Zoology 37(1) :57-64. i 

West, Georce C., and MartHa S. MENG. 
1968a. The effect of diet and captivity on 1 
the fatty acid composition of redpoll (Acan- - 
this flammea) depot fats. Comparative © 
Biochemistry and Physiology 25(2) :535— 
540. 


ne 


, and . 1968b. Seasonal changes 
in body weight and fat and the relation of 
fatty acid composition to diet in the willow ¥ 
ptarmigan. Wilson Bulletin 80(4) :426-— 
441. 

WuitTe, ABRAHAM, PHILIP HANDLER, and | 
Emit L. Smitu. 1968. Principles of bio-~ 
chemistry. Fourth edition. The Blakiston * 
Division, McGraw-Hill Book Company, 
Inc., New York. 1187 p. 


INDEX 


A 


Acanthis flammea (see redpoll) 
Adrenal glands 

as indicator of stress, 103, 108 

mean weights of, 95, 100 

weight correlated with body weight, 94, 

96 

Adrenocortical activity, 103 
Adrenocorticosteroids, 104 
Adrenocorticotropic hormone (ACTH), 103 
Amino acids 

analyzer, 90 

requirements for, 101 

essential to birds, 102 
Arachidonic acid, 96 
Arginine, 101, 102 
Atomic absorption spectrophotometry, 90 


Bobolink, 105 

Bobwhite, 103, 106 

Body weight 
changes in, 90-91, 97, 107 
correlated with adrenal weight, 94, 96 
correlated with lysine utilization, 93-94 
correlated with protein utilization, 93, 94 
of wild pheasants, 100 


procedures for recording, 89 
Bonasa umbellus (see ruffed grouse) 


Cc 


Calcium, 89, 90, 105, 108 
Calories 
consumption of, 93, 98, 99 
determination of, 90 
in experimental diets, 89, 107 
utilization of, 92, 93, 98, 99 
Care and housing of pheasants, 88 
Chicken, 87, 101, 102, 105, 106 
Colinus virginianus (see bobwhite) 
Colorimetry, 90 
Coracobrachialis, 89 
Corn 
as food for wild animals, 87, 107 
chemical composition of, 89, 101 
metabolizable energy (ME) of, 101 
Pioneer 3306, 88, 89 
Corticosteroids, 103, i104 
Coturnix coturnix japonica (see Japanese 
quail) 
Cystine, 101, 102 


D 

Diets 

chemical composition of, 89, 101 
Digestibility coefficients 

defined, 91 

for experimental diets, 91-92, 98 
Dissecting procedures, 89 
Dolichonyx oryzivorus (see bobolink) 


Egg-laying 
timing of, 105 

Excreta 
mean weights of, 92, 98 
procedures for collecting, 89 


F 


Fat deposits 
deposition and mobilization, 104 
mean weights of, 95, 100 
Fat strip (see fat deposits) 
Fatty acids 
in experimental diets, 89 
in pheasants, 94, 96, 100 
saturated: unsaturated ratios, 89, 96, 100, 
104, 108 
Feeding trials 
procedures for, 88 
Flame spectrophotometry, 90 
Flight and maintenance chow (FMC) 
chemical composition of, 89, 101 
Floury-2, 106 
Food consumption 
by experimental hens, 91, 92, 98 
procedures for measuring, 89 


G 


Gallus gallus (see chicken) 

Gizzard, 95 

Glands, 89-90, 94, 95, 96, 99, 100, 103-104, 
105, 108 

Gonadal recrudescence, 105, 106 


H 


High-lysine corn 

defined, 87 
Hypothalamo-hypophyseal axis, 105, 108 
Hypothalamus, 103 


Japanese quail, 106 


K 


Kidneys, 95 
Kjeldahl procedures, 90 


L 


Lagopus lagopus alascensis (see willow 
ptarmigan ) 

Lagopus lagopus scoticus (see red grouse) 

Lauric acid, 96 

Leucine, 87 

Linoleic acid, 96, 104-105, 108 


111 


112 Inurnors Narurat History Survey BULLETIN 


L (cont.) 


Linolenic acid, 96 

Liver, 95, 100 

Lysine 
consumption of, 92, 93, 99 
correlated with body weight, 93-94 
determination of, 90 
importance of in nutrition, 87, 103, 108 
in experimental diets, 89, 101, 107 
requirements for, 101, 102 
utilization of, 93, 99 


M 


Magnesium, 89, 90 
Meleagris gallopavo (see turkey) 
Metabolizability coefficient 
defined, 92 
for experimental diets, 92, 93, 98, 99, 107 
Metabolizable energy (ME) 
requirements for, 101 
supplied by experimental diets, 101 
Methionine, 101, 102, 103 
Microtus pennsylvanicus (see vole) 
Minerals, 89, 90, 105, 108 
Mortality 
related to diet, 106, 108 
Muscles (see sternal muscles) 
Myristic acid, 96 


N 


Nitrogen 
determination of, 90 
Null hypothesis 
level accepted or rejected, 90 


oO 


Oleic acid, 96, 104-105, 108 
Opaque-2, 87, 106, 107 
Organs, 89-90, 94, 95, 99, 100 
Ovary, 100 

Oviduct, 100 

Ovum, 100 


Palmitic acid, 96 
Palmitoleic acid, 96 
Pancreas, 95 


Vol. 31, Art. 3 


Parathyroid glands 


as indicator of negative calcium balance, 
105, 108 
functions of, 105 
mean weights of, 95, 100 
Parathyroid hormone, 105 
Pectoralts thoracia, 89 
Phosphorus, 89, 90 
Potassium, 89, 90 
Protein 
consumption of, 92, 98 
correlated with body weight, 93, 94 
determination of, 90 
in diets of wild pheasants, 102 
in excreta, 92, 98 
in experimental diets, 89, 107 
requirements for, 101, 102 j 
utilization, 92-93, 98, 102 


R 
Rats, 87 
Red grouse, 105 
Redpoll, 104 
Ruffed grouse, 103 

s) 


Sodium, 89, 90 

Spleen, 95 

Stearic acid, 96 

Sternal muscles, 95, 100 

Stress, 103 

Supracoracoideus 
ventral head of, 89 

Swine, 87 


Thymus glands, 85 
Thyroid glands, 95, 100 
Triglycerides, 104 
Tryptophan, 87, 101, 102 
Turkey, 87, 101, 102 


V 
Visceral fat (see fat deposits) 
Vole, 103 

WwW 


Willow ptarmigan, 105 


Some Publications of the ILLINOIS NATURAL HISTORY SURVEY 3 


BULLETIN 


Volume 30, Article 3—Migrational Behavior of 
Mallards and Black Ducks as Determined 
from Banding. By Frank C. Bellrose and 
Robert D. Crompton. September, 1970. 68 
p., frontis., 25 fig., bibliogr., index. 

Volume 30, Article 4.-Fertilization of Estab- 
lished Trees: A Report of Field Studies. By 
Dan‘Neely, E. B. Himelick, and Webster R. 
Crowley, Jr. September, 1970. 32 p., fron- 
tis., 8 fig., bibliogr., index. 

Volume 30, Article 5A Survey of the Mussels 
(Unionacea) of the Illinois River: A Pollut- 
ed Stream. By William C. Starrett. February, 
tis., 8 fig., bibliogr., index.’ 3 

Volume 30, Article 6—Comparative Uptake 
and Biodegradability of DDT and Methoxy- 
chlor by Aquatic Organisms. By Keturah A. 
Reinbold, Inder P. Kapoor, William F. 
Childers, Willis N. Bruce, and Robert L. 
Metcalf. June, 1971. 12 p., frontis., 5 fig., 
bibliogr., index. 

Volume 30, Article 7—A Comparative Study of 
Two Components of the Poinsettia Root Rot 
Complex. By Robert S. Perry. August, 1971. 
35 p., frontis., 10 fig., bibliogr., index. 

Volume 30, Article 8—Dynamics of Condition 
Parameters and Organ Measurements in 
Pheasants. By William L. Anderson. July, 
1972. 44 p., frontis., 6 fig., bibliogr., index. 

Volume 31, Article 1—The Effects of Supple- 
mental Feeding and Fall Drawdowns on the 
Largemouth Bass and Bluegills at Ridge 
Lake, Illinois. By George W. Bennett, 
H. Wickliffe Adkins, and William F. Chil- 
ders. January, 1973. 28 p., frontis., 8 
fig., bibilogr., index. 


BIOLOGICAL NOTES 


71—A Synopsis of Common and Economic 
Illinois Ants, with Keys to the Genera 
(Hymenoptera, Formicidae). By Herbert 
H. Ross, George L. Rotramel, and Wallace 
E. LaBerge. January, 1971. 22 p., 27 fig., 
bibliogr. 

72.-The Use of Factor Analysis in Modeling 
Natural Communities of Plants and Ani- 
mals. By Robert W. Poole. February, 1971. 
14 p., 14 fig., bibliogr. 

73.-A Distributional Atlas of Upper Mississip- 
pi River Fishes. By Philip W. Smith, Alvin 
C. Lopinot, and William L. Pflieger. May, 
1971. 20 p., 2 fig., 107 maps, bibliogr. 

74.-The Life History of the Slenderhead Dart- 
er, Percina phoxocephala, in the Embarras 


List of available publications mailed on request 


No charge is made for publications of the ILt1nors NATURAL History Survey. A sing! 
of most publications will be sent free to anyone requesting it until the supply becomes low. 
publications, more than one copy of a publication, and publications in short supply are sub 
for special correspondence. Such correspondence should identify the writer and explain 
to be made of the publication or publications. 


Address orders and correspondence to the Chief, 


River, Illinois. By Lawrence M. 
Philip W. Smith. July, 1971. 14 p., 
bibliogr. ‘ 
75—Illinois Birds: Turdidae. By 
Graber, Jean W. Graber, and Ei 
Kirk. November, 1971. 44 p., 40. 
liogr. 4 
76.-Illinois Streams: A Classification 1 
Their Fishes and an Analysis of F; sto 
sponsible for Disappearance of N: 
cies. By Philip W. Smith. Novemb 
14 p., 26 fig., bibliogr. 
77-The Literature of Arthropods — 
with Soybeans. I. A Bibliograp! 
Mexican Bean Beetle, Epilachna 
Mulsant (Coleoptera: Coccinel 
M. P. Nichols and M. Kogan. 
1972. 20 p., 1 fig., bibliogr. 
78—The Literature of Arthropods / 
with Soybeans. II. A Bibliography o 
Southern Green Stink Bug, Nezara u 
(Linneaus) (Hemiptera: Penta’ 
By N. B. DeWitt and G. L. Godfrey. 
1972. 23 p., 1 fig., bibliogr. 
79.-Combined Culture of Channel Cai 
Golden Shiners in Wading Po 
Homer Buck, Richard J. Baur, C 
Thoits III, and C. Russell Rose. ; 
12 p., 3 fig., bibliogr. g 
80.-Illinois Birds: Hirundinidae. By R 
R. Graber, Jean W. Graber, and E 
Kirk. August, 1972. 36 p., 30 fig., bi 
81.—Annotated Checklist of the Butt 
Illinois. By Roderick R. Irwin and Jc 
Downey. May, 1973. 60 p., 3 fi 
maps, bibliogr. 
82.—Lactate Dehydrogenase Isozymes of Da 
ters and the Inclusiveness of the G 
Percina. By Lawrence M. Page and 
ory S. Whitt. May, 1973. 7 p., 
bibliogr. 


CIRCULAR 


46—Illinois Trees: Their Diseases. By 
ric Carter. June, 1964. (Third p 
with alterations.) 96 p., frontis., 89 fig 
49-The Dunesland Heritage of Illinois. 
Herbert H. Ross (in cooperation with III 
Department of Conservation). August, 
28 p., frontis., 16 fig., bibliogr. 
51-Illinois Trees: Selection, Planting 
Care. By J. Cedric Carter. August. 
123 p., frontis., 108 fig. ag 
52.-Fertilizing and Watering Trees. By D 
Neely and E. B. Himelick. December, 193 
(Third printing.) 20 p., 9 fig., bibli 
53.—Dutch Elm Disease in Illinois. By J. C 
Carter. October, 1967. 19 p., frontis., 1 


ai | ILLINOIS 
atural History Survey 
| BULLETIN 


An Urban Epiphytotic 
Of Phloem Necrosis and 
Dutch Elm Disease, 1944-1972 


| 
NATURAL HISTORY SURVEY 
AUG 13 1974 
LIBRARY 


THE LIBRARY OF TRE 


AUG 9 1974 


ssi TY OF ILLINOIS 
AT NW RBANA-CHANPAIGN 


E OF ILLINOIS 
ARTMENT OF REGISTRATION AND EDUCATION 


JRAL HISTORY SURVEY DIVISION 


BANA, ILLINOIS 
VOLUME 31, ARTICLE 
MAY, 1974 ~— 


a — 
" 
Be. | 


| 
| 


ILLINOIS 


Tatural History Survey 
| BULLETIN 


An Urban Epiphytotic 
Of Phloem Necrosis and 
Dutch Elm Disease, 1944-1972 


dric Carter 
ile Rogers Carter 


} OF ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


URAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 


VOLUME 31, ARTICLE 4 
MAY, 1974 


DEPARTMENT OF REGISTRATION AND EDUCATION 
BOARD OF NATURAL RESOURCES AND CONSERVATION 


DEAN BARRINGER, Ph.D., Chairman; THomMas Park, Ph.D., Biology; L, L. Sioss, Ph.D., Geology; Herpert §. 
GuTowsky, Ph.D., Chemistry ; Ropert H. ANDERSON, B.S.C.E., Engineering ; Cuartes E, Oumstep, Ph.D,, Forestry ; 
W. L. Everirt, E.E., Ph.D., Representing the President of the University of Illinois; E.perr H, Hapuey, Ph.D, 
Representing the President of Southern Illinois University. 


NATURAL HISTORY SURVEY DIVISION, Urbana, IlIlinois 
SCIFNTIFIC AND TECHNICAL STAFF 


GEORGE SPRUGEL, JR., Ph.D., Chief 
AuicE K, Apams, Secretary to the Chief 


STATE OF ILLINOIS 


Section of Economic Entomology 


WitiiamM H. Luckmann, Ph.D., Entomologist and Head 
Wits N. Bruce, Ph.D., Entomologist 
Wayne L, Howe, Ph.D., Entomologist 
STEVENSON Moorg, III, Ph.D., Entomologist, Extension 
Howarp B. Perry, Ph.D., Entomologist, Extension 
James E. APPLEBY, Ph.D., Associate Entomologist 
Epwarp J. ARMBRUST, Ph.D., Associate Entomologist 
Marcos Kocan, Ph.D., Associate Entomologist 
JosePpH VY. Mappox, Ph.D., Associate Entomologist 
Ronatp H. Meyer, Ph.D., Associate Entomologist 
Ropert D. PauscH, Ph.D., Associate Entomologist 
Raupu E. SecwrRiest, Ph.D., Associate Entomologist 
Joun K. BousemMan, M.S., Assistant Entomologist 
GeorGE L, Goprrey, Ph.D., Assistant Entomologist 
WixuiaM G. Ruesink, Ph.D., Assistant Entomologist 
JAMES R. Sanborn, Ph.D., Assistant Entomologist 
DovuGuas K. Seu, Ph.D., Assistant Entomologist 
CuaRENCcE E. Wuits, B.S., Assistant Entomologist 
Keun S. Park, M.S., Assistant Chemist 
Sue E. Warkins, Supervisory Assistant 
DonaLtp E. KuHLMAN, Ph.D., Assistant Professor, Exten- 
sion 
Rosco—E RANDELL, Ph.D., Assistant Professor, Extension 
Tim Coouey, M.A., Assistant Specialist, Extension 
Joun F. Watt, M.S., Assistant Specialist, Extension 
JEAN G. Wiuson, B.A., Supervisory Assistant 
DaniEt P. BarTELL, Ph.D., Research Associate 
MartTHa P. Nicuous, M.S., Research Associate 
Susan Bonn, B.S., Research Assistant 
STEPHEN D. Cowan, B.S., Research Assistant 
STEPHEN K. Evrarp, B.S., Research Assistant 
Barbara J. Forp, M.A., Research Assistant 
Raymonp A. Korex, M.Mus., Research Assistant 
RosE ANN Meccoul, B.S., Research Assistant 
BarRBARA E. PETERSON, B.S., Research Assistant 
ANNEMARIE REpgoRG, B.S., Research Assistant 
KETURAH REINBOLD, M.S., Research Assistant 
STEPHEN Roserts, B.S., Junior Professional Scientist 
JOHN T. SHaw, B.S., Junior Professional Scientist 
Lowe. Davis, Technical Assistant 
CHARLES G, HELM, M.S., Technical Assistant 
Linpa IsENHOWER, Technical Assistant 
Lu-pine LEE, M.S., Technical Assistant 


Section of Botany and Plant Pathology 

J. Cepric Carter, Ph.D., Plant Pathologist and Head 
Rosert A. Evers, Ph.D., Botanisi 

Junius L. Forsperc, Ph.D., Plant Pathologist 
EvuGeNeE B. HIMEvick, Ph.D., Plant Pathologist 

R. Dan NEELY, Ph.D., Plant Pathologist 

D. F. ScHOENEWEISS, Ph.D., Plant Pathologist 

J. LELAND CRANE, Ph.D., Associate Mycologist 
Water Hartstirn, Ph.D., Assistant Plant Pathologist 
Berry S. Neuson, Junior Professional Scientist 

Gene E. Rew, Technical Assistant 


Section of Aquatic Biology 

D. Homer Buck, Ph.D., Aquatic Biologist 

R. Wexpon Larimore, Ph.D., Aquatic Biologist 

Rosert C. HILTIBRAN, Ph.D., Biochemist 

Witiiam F, Cuinpers, Ph.D., Associate Aquatic Biolo- 
gist 

ALLISON BricHaM, Ph.D., Assistant Aquatic Biologist 

WarREN U. BriGHaM, Ph.D., Assistant Aquatic Biologist 

RicuarpD E. Sparks, Ph.D., Assistant Aquatic Biologist 

JoHN TRANQUILLI, M.S., Assistant Aquatic Biologist 

DonaLp W. Durrorp, M.S., Junior Professional Scientist 

Mary Frances Martin, Junior Professional Scientist 

Joun M. McNurney, M.S., Junior Professional Scientist 


CONSULTANTS AND RESEARCH AFFILIATES: 


life Research, Southern Illinois University ; 


of Illinois. 


Systematic Entomouocy, Ropertck R. Irwin, Chicago, Illi 
nois ; WILDLIFE RESEARCH, WILLARD D. Kuimstra, Ph.D., Professor of Zoology and Director of Cooperative Wi 
ParasitoLoGy, NormMAN D. Levine, Ph.D., Professor of Veterina 
Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, University 
Illinois ; ENTomMoLoGy, Ropert L. Metcaur, Ph.D., Professor of Zoology and of Entomology and Head of the De- 
partment of Zoology, University of Illinois; and GitBERT P. WAtDBAUER, Ph.D., Professor of Entomology, Un 
versity of Illinois; Statistics, Horace W. Norton, Ph.D., Professor of Statistical Design and Analysis, Universi 


Tep W. Storck, B.S., Junior Professional Scientist 
RicHaRp J. Baur, M.S., Research Assistant 

Tom Hit, M.S., Research Assistant 

RicwarD Kocuer, B.S., Research Assistant 

Rospert Moran, M.S., Research Assistant 

Linpa Kurppert, B.S., Technical Assistant 

C. Russet Rose, Field Assistant 


Section of Faunistic Surveys and 
Insect Identification 


PuHiuip W. SmitH, Ph.D., Taxonomist and Head 
Watuace E, LaBerGE, Ph.D., Taxonomist 

Mitton W. Sanperson, Ph.D., Taxonomist 
Lewis J. STANNARD, JR., Ph.D., Taxonomist 
Larry M. PaGe, Ph.D., Assistant Taxonomist 

Joun D. Unzicker, Ph.D., Assistant Taxonomist 
DonaLp W. WEBB, M.S., Assistant Taxonomist 
BERNICE P. SWEENEY, Junior Professional Scientist 


Section of Wildlife Research 


GuEN C. SanpEeRSON, Ph.D., Wildlife Specialist and Head 
Frank C, Beuurose, B.S., Wildlife Specialist q 
JEAN W. GraBeER, Ph.D., Wildlife Specialist 
RicHarp R. GRaBeR, Ph.D., Wildlife Specialist 
Haroutp C, Hanson, Ph.D., Wildlife Specialist 
Ronaup F. Lapisky, Ph.D., Wildlife Specialist Z 
WivuiaM L, AnprersoN, M.A., Associate Wildlife Special- 
ist 
W. W. Cocuran, IJR., B.S., Associate Wildlife Specialisi 
Wixuiam R. Epwarps, M.S., Associate Wildlife Special- 
ist 
Jack A. Exuis, M.S., Associate Wildlife Specialist 
G. Buarr JosELYN, M.S., Associate Wildlife Specialist — 
CuHarLes M. Nixon, M.S., Associate Wildlife Specialist 
KENNETH E. SmitH, Ph.D., Associate Chemist 
Ronatp L. WESTEMEIER, M.S., Associate Wildlife Spe- 
cialist 
STEPHEN P. Havera, M.S., Assistant Wildlife Specialist 
Davin R. Vance, M.S., Assistant Wildlife Specialist 
Ronaup E. Duzan, Junior Professional Scientist 
HeLen C. Scuuutz, M.A., Junior Professional Scientist 
ELEANORE WILSON, Junior Professional Scientist 
SHARON FRADENBURGH, B.A., Laboratory Technician 
Rogert D. Crompton, Field Assistant 
James W. SEEtS, Laboratory Assistant 


Section of Administrative Services 
Rosert O. Watson, B.S., Administrator and Head 


Supporting Services 
Vernon F, Bittman, Maintenance Supervisor q 
Witma G. Dituman, Property Control and Trust Ac- 
counts 

Ropert O. Euuis, Assistant for Operations 
Luoyp E, HurFMAN, Stockroom Manager R 
J. Witt1am Lusk, Mailing and Distribution Services 
MELVIN E. ScHwartz, Financial Records 
James E. SERGENT, Greenhouse Superintendent 


‘Publications and Public Relations 
Owen F. GuissenporF, M.S., Technical Editor % 
Ropert M. ZEWADSKI, M.S., Associate Technical Editor 
SHIRLEY MCCLELLAN, Assistant Technical Editor 
Luoyp LeMeErRE, Technical Illustrator 
LawrENCE S. Fartow, Technical Photographer 


Technical Library 
Doris F. Dopps, M.S.L.S., Technical Librarian J 
Doris L. SuBLETTE, M.S.L.S., Assistant Technical Li- 
brarian 


| CONTENTS 
| 
ee gered ge. Se OS eee eer ee ee ane ae 114 
MEERUT TRIMER VTLWi Stet eacioes earthen ol sore clays cnoed artys ava ate eas ahe sab) atte che Sayetecs 115 
—«. THETRTUNTES ZN) IMIS OS, Gon ogog5 sou sceasucooen ods HUpSSerocHoaasaBaed 116 
| 
RESULTS NN Sr ree Puen thee UNG ihts Bie la cet cnansla els lorciale ol eho tevaten's) aimtah sc are% 122 
ease sarcadior Phlocny NECIOSISi a)! 5.j2s'0. oe vids yee = ale eae a 122 
| Appearance and Spread of Phloem Necrosis in the 700 Block of South 
iMmynnestreety Champaign |O4OSN Gay eee cysts leieeia)- settle ee eres: 123 
Spread of Phloem Necrosis in 1951 and 1952 and Influence of Phloem 
| Necrosis on the Early Spread of Dutch Elm Disease .............. 123 
| Appearance of Dutch Elm Disease and Locations of Elms Having 
Dntehelrmns Disease. Oa 19D Bel ice vaciecele ose ek ieiels ee « Sleqee ole sis wie 124 
| Spread of Dutch Elm Disease in the 700 Block of West Michigan 
| Avenue nbana Ono 959 wicca cies. cae sects as vas sce Hoylake 127 
Spread of Phloem Necrosis and Dutch Elm Disease on Springfield 
AVeniiem @hamipargnr Goa Q09 en. cari.) aeeetsiel sale sie wleresatels «ells a cie-\= 128 
Phloem Necrosis-Affected Elms That Later Showed Symptoms of 
Witchy blmM Seas: eis see, sce yetes ose ie.crs ys sh2y Ren Serine p ee one 130 
Incidence of Phloem Necrosis and Dutch Elm Disease in Champaign- 
. MOlaatyerracr eel GAA Gi aera ete epee sya clea syne MMe (os ow ieee ise chine 0 ai8,sisceons 130 
| Palen INGORCSE feec pookanca oases ceca nen Tee ereaen cual cic 132 
iDinieln, Tila DIES: 6m Secor ere OIG Oe eee eS econ 133 
Effect of Each Disease on the Residual Population of Elms ....... 134 
Accumulated Percentages of Elms Killed by Each Disease, 
) IN aL-1072 Lath. 2 eee ica ea ce eaten Serr amien tee, 134 
| Time of Year in Which Elms Died from Phloem Necrosis and Dutch 
ilimy IDFRGRGS" 5 2.5 suo8 Aoens cle Gece Oe ae ae ene nor ice 135 
} Period of Time in Which Elms Died Following the Appearance of 
Foliage Wilt of Phloem Necrosis or Dutch Elm Disease .......... 136 
BDISCUSSION -- «1122... eet eet 137 
_ SEMAUTN? Gyo Bigs 26 tints na eg trae ert ero et are areca 139 
REPU EAS IED IA a yi eRe ea eae ete eo cnas' tele Slavs auscnie fo ave eschews. erp = 141 
SESE o o'n'o.0 1d OBER Ete O.GLOr0 0.6. GSS Uae nek UES one ae ea 142 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. 
| It is a contribution from the Section of Botany and Plant Pathology of the Illinois Natural 
| History Survey. 

: 

J. Cedric Carter is Plant Pathologist and Head, Section of Botany and Plant Pathol- 
| ogy, Illinois Natural History Survey. Lucile Rogers Carter, Catalog Department, Univer- 
| sity Library, University of Illinois, assisted in various aspects of the study reported here. 


(31556—3M—5-74) 


Frontispiece.—These before and after pictures along the Broadwalk on the University of Illi 
campus illustrate vividly how a cathedral-like archway of stately American elms can be destroyed quit 
and completely by phloem necrosis and Dutch elm disease. The upper photograph was taken in 5 


tember 1954. The lower photograph was taken in September 1966, several years after the univ: 
sity pad begun replacing the elms with other species. (Upper photo by W. E. Clark;- lower photo) 
. D. Zehr) 


An Urban Epiphytotic of Phloem Necrosis 


and Dutch Elm Disease, 


THE AMERICAN ELM was used 
more extensively than any other species 
as a shade tree in Illinois prior to 1940. 
It was especially valued for its rapid 
growth, majestic size, vase shape, and 
extensive shade. It was used for streets, 
boulevards, and park drives, as well as 
in lawns. In many cities throughout 
the state it represented half or more 
of the shade trees lining the streets. 
In some cities it was estimated that 
three-fourths of the shade trees were 
American elms. Because of the loss 
to diseases of millions of elms, espe- 
cially American elms, since 1940, most 
cities and homeowners now plant other 
trees. The elm problem has made 
many people aware of the importance 
of using diversified plantings to avoid 
such a catastrophe in the future. 


Extensive dying of elms in central 
Illinois, especially in Bloomington, Nor- 
mal, and Champaign, occurred as early 
as 1883 (Forbes 1885:112; 1912:3). 
This dying of elms subsided within 
a few years. Although the cause of the 
elm deaths was not determined, the 
symptoms reported are not typical of 
any current vascular wilt disease. 

In 1907 elms were reported dying 
in southern Illinois in and around 
Fairfield in Wayne County. By 1912 
numerous elms were dying in 14 towns 
(Cairo, Carbondale, Centralia, Clayton, 
Du Quoin, Edwardsville, Fairfield, Gala- 
tia, McLeansboro, Mt. Vernon, Quincy, 
Robinson, Sumner, and Vandalia) in 
13 counties in southern and western 
Illinois (Forbes 1912:5). Careful ex- 
amination of affected trees revealed 
that although the small fibrous roots 
were dead, some of the main roots 
were alive, as were some of the leaves. 
This early dying of elms was generally 


1944-1972 


J. Cedric Carter 
Lucile Rogers Carter 


referred to as “elm blight” and at- 
tributed to various conditions and agen- 
cies, including drought, exhaustion of 
soil nutrients, insect attack (borers and 
bark beetles), and diseases of unknown 
causes (Forbes 1912:7-10). 


Following 1912 no reports of ex- 
tensive dying of elms appeared until 
1930, when many elms were killed in 
and near several large and small cities, 
notably Hillsboro in south-central Illi- 
nois, Danville in east-central Illinois, 
Peoria in north-central Ilinois, Quincy 
in west-central Illinois, and Cairo 
at the southern tip of the state 
(Carter 1945:23). By 1940 (Carter 
1954) elms were dying throughout 
the southern half of the state and as 
far north as Danville on the east and 
Quincy on the west. Other towns on 
the northern border of this area in- 
cluded Charleston, Shelbyville, Taylor- 
ville, and Pittsfield. The greatest num- 
ber of affected trees were in a broad 
belt extending diagonally southwest- 
ward from Danville and Paris on the 
east to Alton and Belleville on the west. 
This area is north of the southern part 
of the state where heavy losses of elms 
occurred earlier. This dying of elms 
continued to spread northward to Ur- 
bana in 1944, to Mattoon, Springfield, 
and Lincoln in 1945, and to Decatur, 
Bloomington, and the area around Bur- 
lington, Iowa, in 1948. General and 
widespread dying of elms became 
stabilized by 1948 with the northern 
boundary of the affected area repre- 
sented by a line extending from Dan- 
ville on the east through Champaign- 
Urbana, Bloomington, and Peoria to 
the area around Burlington, Iowa, on 
the west. North of this area dying elms 
were found in only six cities — Melvin 


118 


114 


and Dwight in 1945, Rockford in 1946, 
Chebanse in 1950, and Chenoa and 
Onarga in 1953. 


Following the 1938 report that a dis- 
ease of a virus nature was killing elms 
in Ohio (Swingle 1938), it was soon 
determined that the widespread dying 
of elms in the Ohio Valley region of 
the Midwest was caused by the same 
disease. It was determined to be a 
virus disease called phloem necrosis 
(Swingle 1940 and 1942). However, 
recent work by Wilson et al. (1972) 
indicates that elm phloem necrosis may 
be caused by a mycoplasmalike or- 
ganism (MLO). 


The symptoms exhibited by dying 
elms in Illinois as early as 1912 were typ- 
ical of those described later for phloem 
necrosis. It appears that this disease 
was the major cause of elms dying in 
southern Illinois from 1907 to 1950. 


Dutch elm disease, caused by Cera- 
tocystis ulmi (Buism.) C. Moreau, was 
discovered in one American elm in 
Coles County (east-central Illinois) in 
1950, and the disease spread rapidly, 
especially in the area where elm 
phloem necrosis was killing thousands 
of elms annually. By 1969 it had spread 
into all 102 counties of the state. At 
present each disease kills many elms 
annually, phloem necrosis in the south- 
ern two-thirds of the state and Dutch 
elm disease throughout the state. 


The first symptom of phloem necrosis 
is the dying of fibrous roots. This 
symptom is followed by foliage symp- 
toms, which appear as drooping leaf 
blades and upward curling leaf mar- 
gins. Next the leaves turn yellow, 
brown, or both, and drop from the 
tree. These symptoms may occur Over 
one or more growing seasons. On some 
trees the foliage wilts rapidly within a 
few weeks and turns brown, but many 
leaves remain attached to the branches. 
Occasionally a tree may have one or a 
few large branches dying simultane- 
ously over a period of 1 or more years. 


The infection of individual branches 
by the Dutch elm disease fungus re- 


ILLINOIS NATURAL History Survey BuLiLetin 


Vol. 31, Art. 4 


sults in wilting and dying foliage on 
the affected branches. Frequently, the 
early symptom of Dutch elm disease 
is wilting foliage on one or a few 
branches. Wilt of the entire tree oc- 
curs after the fungus becomes systemic 
in the tree. The yellowing of leaves 
over the entire crown is uncommon 
in Dutch elm disease but common in 
phloem necrosis. 


The present study was initiated, fol- 
lowing the appearance of phloem 


necrosis in Urbana in 1944, to obtain’ 


data on the number of elms affected 
annually by the disease and to study 
the pattern of spread in a municipal 
area where no city-wide control pro- 
gram was practiced. With the appear- 
ance of Dutch elm disease in Urbana 
in 1951, the study was expanded to 
include both diseases and the relation- 
ship of the two diseases in a municipal 
area. 


ACKNOWLEDGMENTS 


The authors wish to express their 
sincere appreciation to all those who 
have assisted or advised in any way 
during the course of this investigation. 
Special recognition is given to. Drs. 
Dan Neely, J. L. Forsberg, and J. W. 
Gerdemann for reviewing and making 
numerous helpful suggestions in the 
preparation of the manuscript. 


The photograph for Fig. 9 was taken 
by William Clark, formerly Natural 
History Survey Photographer. All other 
illustrations, except as noted, were 
photographed by Wilmer Zehr, Survey 
Photographer. The illustrations in Fig. 
5, 7, 8, 10, 11, and 12 were prepared 
by Lloyd LeMere, Survey Illustrator. 


We are grateful to Robert M. Zewad- 
ski, Survey Associate Editor, for his 


painstaking efforts in editing the manu-— 


script and to Mrs. Betty Ann Nelson 
for typing the manuscript. 


The research policies of the Natural 
History Survey made it possible for the 
authors to carry on this study over a 
period of 29 years. 


May, 1974 CARTER & CARTER: 


LITERATURE REVIEW 


Field-grown elms do not show symp- 
toms of phloem necrosis for 1 or more 
years following the introduction of the 
MLO (Baker 1949:729, 730; Campana 
1958). Elms affected by this disease 
usually die during the growing season 
in which symptoms first appear. How- 
ever, some elms may show symptoms 
for two or sometimes three growing 
seasons before dying (Campana 1958; 
Swingle 1938:757). Phloem necrosis 
affects only the American elm (Ulmus 
americana L.) and its cultivars and 
the winged elm (U. alata Michx.). The 
phloem necrosis MLO is carried by the 
elm leafhopper (Scaphoideus luteolus 
Van D.). The MLO is introduced into 
the leaves of healthy elms by the feed- 
ing activity of elm leafhoppers that 
have previously fed on leaves of phloem 
necrosis-affected elms. 


Elms affected by Dutch elm disease 
may die during the growing season 
when symptoms first appear, or they 
may die during the following growing 
season. Occasionally an affected elm 
may not die until the third growing 
season (Banfield, Rex, & May 1947). 
Dutch elm disease affects all elm spe- 
cies, but the Asiatic species are rela- 
tively resistant (Bretz, Swingle, & 
Parker 1945). The American elm ap- 
pears to be more susceptible than the 
other native and European species 
(Neely & Carter 1965). 


Published data on the loss of elms 
in municipal areas where both diseases 
occur are limited. However, data are 
available on losses caused by each 
disease in areas where only one was 
present (Campana & Carter 1955 and 
1957; Carter 1954, 1955, and 1961; 
Neely, Carter, & Campana 1960; Neely 
1967). 


In municipal plantings of elms, losses 
resulting from phloem necrosis may 
increase from a few scattered trees 
when the disease is first discovered to 
several hundred trees annually within 
3 years. In three municipalities in 
central Illinois the number of phloem 


PHLOEM NECROSIS AND DuTcH ELM DISEASE 115 


necrosis-affected elms increased from 9 
in 1945 to 370 in 1948 in Lincoln, 
from 5 in 1945 to 417 in 1948 in 
Mattoon (Carter 1950:50), and from 6 
in 1948 to 154 in 1951 in Bloomington.' 
In Mt. Pulaski, which had approxi- 
mately 600 elms in 1942 when phloem 
necrosis was first found there, all but 
19 elms were killed by the disease by 
1948 (J. C. Carter, unpublished data). 


Phloem necrosis also kills elms in 
rural areas. In a survey made in south- 
ern Illinois in 1945, the numbers of 
elms recorded as dying from phloem 
necrosis were 1,644 in rural areas and 
1,655 in municipal areas (Carter 1954). 


In municipal plantings of elms, Dutch 
elm disease losses may increase from 
a few scattered trees when the disease 
is first discovered to hundreds of trees 
each year in a few years. In five munici- 
palities in northern Illinois where Dutch 
elm disease was discovered in 1955 and 
where phloem necrosis was not present, 
the total numbers of diseased elms 
from 1955 through 1961 were over 
4400 in Aurora, 1,600 in Elgin, 2,500 
in Joliet, 1,400 in Waukegan, and 1,000 
in Zion. The percentages of the original 
elm populations that became diseased 
in these five cities during this period 
were 48, 23, 33, 11, and 27, respectively 
(Neely 1967: 513). In Bloomington 
in north-central Ilinois, Dutch elm dis- 
ease was first found affecting 10 elms 
in 1954 following a 6-year period in 
which 932 elms were affected by 
phloem necrosis. The number of elms 
affected annually by Dutch elm dis- 
ease increased to 242 in 1955 and to 
507 in 1956 (Campana & Carter 1957: 
636). 


In areas where both diseases are 
present, the incidence of phloem ne- 
crosis results in an increase in the oc- 
currence of Dutch elm disease (Cam- 
pana & Carter 1955). The native elm 
bark beetle (Hylurgopinus rufipes 
Eichh.) and the smaller European elm 


1 The data on the loss of elms in Bloomington 
were obtained by annual surveys made by the 
senior author. 


116 


bark beetle (Scolytus multistriatus 
Marsh.), vectors of the Dutch elm 
disease fungus, colonize and overwinter 
in the bark of weakened, dying, and 
recently killed elms, including those 
affected by phloem necrosis. 


The incidence of Dutch elm disease 
does not increase the occurrence of 
phloem necrosis (Campana 1958 ). Elms 
killed by either disease and left stand- 
ing are not colonized by the elm leaf- 
hopper (S. luteolus Van D.), vector of 
the phloem necrosis MLO. This insect 
overwinters in the egg stage. The 
eggs are embedded in the soft cork 
parenchyma of elm bark (Baker 1949: 
731). 


Dutch elm disease may not only ob- 
scure the presence of phloem necrosis, 
but it may kill phloem necrosis-af- 
fected trees before external symptoms 
of phloem necrosis become apparent 
(Campana & Carter 1957; Campana 
1958). The rate of increase of Dutch 
elm disease exceeds that of phloem 
necrosis in areas where Dutch elm 
disease appears after phloem necrosis 
has been present for several years 
(Campana & Carter 1957:639; Carter 
1955:36-37; Neely, Carter, & Campana 
1960:167, 169). 


Phloem necrosis-affected elms colo- 
nized by smaller European elm bark 
beetles infested with the Dutch elm 
disease fungus will harbor that fungus. 
Of 40 such trees examined in Urbana 
in 1952, 8 contained the Dutch elm 
disease fungus (Campana 1954:358).? 
Populations of the smaller European 
elm bark beetle increase rapidly in 
areas where dying and recently killed 
elms are present. This situation was 
common in the 1950’s in the southern 
two-thirds of Illinois where thousands 
of elms killed annually by phloem ne- 
crosis were not removed immediately 
upon discovery of the disease (Cam- 
pana 1954), 


2 Data originally gathered by E. B. Himelick, 
Illinois Natural History Survey, and published 
by Campana, 


Intinois NaturaAL History Survey BULLETIN 


Vol. 31, Art. 4 


When symptoms of both diseases 
occur in one elm, infection by the 
phloem necrosis MLO most likely oc- 
curs first, since the MLO is in the 
tree 1 year or more before symptoms 
of phloem necrosis become apparent 
(Baker 1948 and 1949:729, 730; Cam- 
pana 1958; Swingle 1938). Therefore, 
elms showing early symptoms of phloem 
necrosis may show symptoms of Dutch 
elm disease later in the same year or 
in the following year before they die. 
Elms infected by the phloem necrosis 
MLO and the Dutch elm disease fungus 
in the same year may show symptoms 
of Dutch elm disease in that year, but 
not symptoms of phloem necrosis. 


Some of the data presented in this 
bulletin on the loss of elms from phloem 
necrosis and Dutch elm disease in 
Champaign and Urbana have been re- 
ported previously (Campana & Carter 
1955 and 1957; Carter 1955; Neely, 
Carter, & Campana 1960). 


MATERIALS AND METHODS 


The twin cities of Champaign and 
Urbana represent a contiguous munici- 
pal area with a common boundary. This 
area is approximately 5 miles [8 km] 
east and west and 2.5 miles [4 km] 
north and south. It is traversed by 
approximately 200 miles [320 km] of 
streets, mostly lined with trees on each 
side and with additional trees in lawns. 


In 1950 all elms and all other species 
of trees were counted in 75 city blocks 
to obtain an estimate of the entire tree 
population as well as the elm popula- 
tion. To obtain the estimate, the entire 
tree population was divided into 25 
areas. The areas were selected so that 
each area included elms of a given 
size and age. The sizes of trees counted 
varied from approximately 4 to 20 


inches [100-500 mm] in trunk diameter 


and 20-75 feet [6-23 m] in height. 
Most of the larger elms were present 
in the older residential areas. 

All trees were counted in each of 
three blocks selected at random in each 


May, 1974 


of the 25 areas. From the data ob- 
tained (Table 1) it was estimated that 
there were 12,195 elms in the munici- 
pal study area. The data also showed 
that the tree population was composed 
of 51 percent elms and 49 percent 
other species. An estimated 2,000 elms 
in Crystal Lake Park in the north part 
of Urbana were later included in the 
total elm population, bringing to 14,195 
the estimated number of elms in the 


Table 1.— Estimated tree population in 
Champaign-Urbana, Illinois, in 1950. 


Elms Other Species 

City Num- Per- Num- Per- 

ber cent ber cent 

Urbana 4,235 51.8 3,946 48.2 

Champaign 7,960 50.6 7,775 49.4 
Total or 

percent 12,195 51.0 11,721 49.0 


Fig. 1.—Phloem necrosis symptoms. 
phloem necrosis in contrast to the healthy tree on the right. Phloem necrosis is indicated by the 


cupping or rolling and yellowing of foliage over the entire tree crown, 


CARTER & CARTER: PHLOEM NECROSIS AND DutcH ELM DISEASE a lp E7¢ 


twin-city area. The actual number of 
elms recorded during this 29-year study 
was 14,103. Of these elms, 66 percent 
were on public property and 34 per- 
cent on private property. The distance 
between elms along many streets was 
30 feet [9 m] or less. This close spacing 
was common in blocks and along streets 
planted almost exclusively to elms. 
Some lots with frontages of about 50 
feet [15 m] in residential areas had 
two elms on the parkway and one or 
more elms on the front lawn. 


The elms under observation included 
all American and European species of 
elms that could be observed from the 
street on public and private property 
within the confines of Champaign and 
Urbana. In addition to the American 
elm and its cultivar, the Moline elm, 
there were 25 slippery elms and 9 


The tree on the left shows typical symptoms of 


(Photo by J. C. Carter) 


118 In,inois Naturat History Survey BULLETIN Vol. 31, Art, 4 


English elms. Asiatic species were not years to control phloem necrosis, and 
included, as they are immune to phloem _ therefore they were not included in this : 
necrosis and resistant to Dutch elm _ study. ¥ 

a 


disease. The elms on the University Elms removed from the study area 
of Illinois campus were sprayed for 5 but not affected by phloem necrosis or 


Hi 

Fig. 2.—Dutch elm disease symptoms. Dutch elm disease frequently appears as wilt- 
ing and browning of foliage on one or a few branches. This tree has foliage wilting on 
the left major branch. (Photo by J. C. Carter) 


119 


“stounj|| ‘euequp-uBbledweu> ul gp6l YSnoiy} pp6] wor sisoidau waojyd Aq payayye swja 9| 4O suo1ye907—'¢e “Biy 


CARTER & CARTER: PHLOEM NECROSIS AND DuTcH ELM DISEASE 


te s| eee, See | L 
| = a Rae i] a 
=| Sel 
Se el i ea if 
ta Saal. | ae (ral 
SUT nite oo : = bei a 
COPS eo) CLLeiemeceaen ie Coc 
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VNVEYN-NOIVIWNVHO | 


120 


Dutch elm disease, except seven elms 
removed in 1967 because of ice damage, 
were not included in the total number 
of trees recorded in this study. How- 
ever, elms that became large enough 
to be seen from the street during the 
29-year period were included. 

The appearance and spread of each 
disease were recorded, starting with the 
first two trees attacked by phloem 
necrosis in 1944. Dutch elm disease 
did not appear until 1951. 


ILLINoIs NATURAL History Survey BULLETIN 


Vol. 31, Art. 4 


Observations were made at irregular 
intervals during the growing season 
from 1944 through 1950 to determine 
the incidence and pattern of spread 


of phloem necrosis. Diseased trees were — 
located by frequent scoutings. As many 
as 8-14 scoutings were made in a grow- © 


ing season. 

After 1950, the authors made two 
surveys annually through 1972, one in 
June and one in September. However, 
in some years the first survey was not 


af 


“|= 


(sie 


besa) 


Le] | 
| | 


Phloem necrosis 
1944-1948 = Dated on map 
| 1949-1950= @ 


—susa. 


as! 


| = 


Fig. 4.—Distribution of 16 Champaign-Urbana, Illinois, elms affected by phloem necrosis 
from 1944 through 1948 (dated according to year when symptoms appeared) and of 412 


May, 1974 


completed until early July and the sec- 
ond survey was not completed until 
early October. Each survey was made 
by observing all elms visible from the 
street while traveling by automobile. 
This type of survey necessitated driving 
200 miles [320 km] of streets, which 
required a maximum driving time of 
24 hours. We used our personal auto- 
mobile, and the surveying was done 
in the evenings and on Saturdays. 


The presence of phloem necrosis dur- 


AND VICINITY 


CARTER & CARTER: PHLOEM NECROSIS AND DuTcH ELM DISEASE 121 


ing the years of low initial incidence, 
1944-1948, was determined by examin- 
ing the inner phloem for the butter- 
scotch color and wintergreen odor char- 
acteristic of the disease. 

Dutch elm disease during the years 
of low initial incidence, 1951-1952, was 
identified by a laboratory culture test 
for the fungus. 

Following the years of low initial 
incidence, each disease was recognized 
by foliage symptoms. Phloem necrosis- 


CHAMPAIGN-URBANA ; 


Je | / 


al E fa Ee a | 
pi = = 2/ 
| tee gtctoe Wek 
=e 72 / 
aha | jad = J} 
= alm } a) =a a 
TRS SSS. : J he Ee 
fests laa = 
ISSSSsser4— : 
SSS Ssaess _~= = 
SSeS2s44o See 
SS SS eS Sella 
ISSSS| sSss me a 
= ed ] has if = =j— 
Sees Ssss coral i. 
iF tala lal te 3 Ti =] ae 1h 
=) ea 
=i bs 


[a] 

Co 
= lefed 
Fal 
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‘cao 
— all 
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—[ 


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pee 
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elms affected by phloem necrosis in 1949 and 1950 (black dots). Some dots represent more 
than one tree where the elms were less than 30 feet [9 m] apart. 


122 


affected trees commonly show yellow- 
ing and wilting of leaves over the en- 
tire crown (Fig. 1), while Dutch elm 
disease-affected trees frequently have 
wilted foliage only on one or a few 
branches (Fig. 2). However, some 
Dutch elm disease-affected trees show 
general wilting on the entire crown. 
Leaves on such trees usually display 
less yellowing but more rapid browning 
and wilting than leaves show on phloem 
necrosis-affected trees. When there was 
doubt as to which disease was involved, 
a diagnosis was usually made by exam- 
ining wood and bark samples. Also, the 
accuracy of diagnosis by observation 
from an automobile was tested by ex- 
amining an occasional elm for wood and 
bark symptoms. 


Data were recorded as to which dis- 
ease was involved and whether a tree 
was wilting or dead. The stage or 
amount of wilt evident at the time 
of each survey was indicated by listing 
each tree showing the early, intermedi- 
ate, or late stage of wilt. All dead 
elms were listed as dead, and all trees 
were listed by street address and 
whether on public or private property. 


RESULTS 


INITIAL SPREAD OF 
PHLOEM NECROSIS 


Following the discovery in Urbana 
of phloem necrosis in two elms in 1944, 
one elm was affected by this disease 
in 1945, two in 1946, three in 1947, 
and eight in 1948 (Fig. 3). The tree 
affected in 1945 was about 300 feet 
[90 m] west of the two elms affected 
in 1944. Of the two elms affected in 
1946, one was about 400 feet [120 m] 
farther west and one was about 1,900 
feet [580 m] southeast of the elms 
affected in 1944 and 1945. Of the three 
elms affected in 1947, one was about 
150 feet [45 m] northeast of a tree 
affected in 1946. The other two were 
on a line between trees affected in 
1944 and 1946 and over 600 feet [180 m] 
from the nearest previously diseased 
elm. One of the eight elms affected 


ILLInoIs NATURAL History Survey BULLETIN 


Vol. 31, Art. 4 


in 1948 was about 150 feet [45 m] 
north of an elm affected in 1947. The 
remaining seven elms were scattered 
west of previously affected trees. They 
ranged from about 700 to 8,000 feet 
[210-2,440 m] from the nearest previ- 
ously diseased elms. Although the di- 
rection of the spread of phloem necrosis 
was generally west and south, each dis- 
eased tree was surrounded by numerous 
healthy elms. 


The two elms that showed phloem 


7 
‘ 


necrosis symptoms in 1944 were in the — 


200 block of West Main Street in 
Urbana. They were among the oldest 


elms in the city. They were surrounded ~ 


by numerous elms except for the three- 
square-block area of the business dis- 
trict of Urbana, starting in the 100 
block of West Main Street. East of 
the business district, elms were abun- 
dant to the east city limit. Each of 
the 14 elms that became diseased from 
1945 through 1948 represented a sep- 
arate infection center, as none of them 
was within root-grafting distance of 
other affected trees (Himelick & Neely 
1962; Verrall & Graham 1935). Each 
diseased tree was surrounded by 
healthy elms. 


In 1949 and 1950 the number of 
diseased trees increased rapidly in 
Urbana. The disease was confined al- 
most entirely to an area in south-central 
Urbana approximately 12 blocks square 
and extending south from the 200 block 
of West Main Street, the location of 
the two elms that first contracted 
phloem necrosis. By 1951, when Dutch 
elm disease was first found in Urbana, 
phloem necrosis was concentrated in 
this 12-block area and scattered at 
random in all directions around 7 of 
the 12 previously diseased Urbana elms, 
as shown in Fig. 4. The disease had 
not invaded an area six blocks wide 
along the west boundary of Urbana. 
In Champaign the disease appeared in 
the 700 block of South Lynn Street 
in 1949. By 1951 it had spread to 
many surrounding elms, as described 
subsequently. A few scattered elms 
north of this area and two elms within 


May, 1974 CARTER & CARTER: 
two blocks of the west boundary of 
Champaign were also diseased. No 
elms were affected within the immedi- 
ate areas surrounding the four Cham- 
paign elms that became diseased be- 
fore 1949. 


APPEARANCE AND SPREAD OF 
PHLOEM NECROSIS IN THE 700 
BLOCK OF SOUTH LYNN STREET, 
CHAMPAIGN, 1949-1951 


Only four elms showed symptoms of 
phloem necrosis in the entire city of 
Champaign in 1948, the year before 
the disease appeared in the 700 block 
of South Lynn Street. The nearest of 
these four elms was over 1,200 feet 
[370 m] northeast of the South Lynn 
Street 700 block (Fig. 4). Of the 36 
elms in that block, 23 were on the 
west side of the street and 13 were on 
the east side. On the west side 21 
were parkway trees and 2 were on 
private property, while 8 were park- 
way trees and 5 were on private prop- 
erty on the east side of the street. Each 
of the 21 parkway elms and 2 private- 
property elms on the west side of the 
street were within root-grafting dis- 
tance of one or more elms (Fig. 5). 
The number of elms having phloem 
necrosis each year and the month in 
which symptoms were first observed are 
shown in Fig. 5. 


On South Lynn Street, 8 elms wilted 
in 1949, 21 in 1950, and 7 in 1951. Of 
the eight elms that wilted in 1949, 
seven were on the west side of the 
street, and each was separated from 
the others by at least one elm that did 
not show wilt symptoms in 1949. Of 
11 closely planted parkway elms on 
the west side of the street, every other 
one (5 trees) wilted in 1949, and the 
remaining 6 trees wilted in 1950. 


The wilting of eight elms in 1949 
suggests that the elm leafhopper, vector 
of the phloem necrosis MLO, fed ex- 
tensively on the trees in this limited 
area before 1949. Since all 36 elms 
wilted within 3 years, all transmission 
of the MLO may have been by the 
elm leafhopper. However, most of the 


PHLOEM NECROSIS AND DuTcH ELM DISEASE 


123 


JOHN ST. 


703 | 
9-49 ——_. 
6-50__705 |* 


ee 
ogee 


6-50 _ 707 |, 
9 scree 
6-50-09 |. 
9-49 


e-50e 


e \ 
7I6 6-50 


Reo 


718 


720 


722 6-50 


724 


726 


WILLIAM ST. 


5.—The spread of phloem necrosis, 


Fig. 
1949-1951, in the 700 block of South Lynn 
Street, Champaign, Illinois. 


elms that wilted in 1950 and 1951 were 
adjacent to elms that had wilted during 
the previous year, and they could have 
become infected through root-graft 
transmission of the MLO. Only three 
of the 36 elms were beyond root- 
grafting distance of other elms. 


SPREAD OF PHLOEM NECROSIS IN 
1951 AND 1952 AND INFLUENCE 
OF PHLOEM NECROSIS ON THE 
EARLY SPREAD OF 

DUTCH ELM DISEASE 


The spread of phloem necrosis in 
Champaign-Urbana in 1951 and 1952 
occurred among elms both within and 


124 


Intrnois NaturAL History SuRvEY BULLETIN 


Vol. 31, Art. 4 


Table 2.—Spread of phloem necrosis in Champaign-Urbana, Illinois, in 1951 and 1952 
in relation to the location of elms affected by phloem necrosis in the previous year. 


Elms Within Root-Grafting 
Distance of Previously 
Diseased Trees 


Venn Diseased 
Elms 
Number 
1951 359 116 
1952 555 186 
Total or percent 914 302 


beyond root-grafting distance of previ- 
ously affected trees. Of the 914 elms 
affected by phleom necrosis in 1951 
and 1952, 33 percent were within and 
67 percent were beyond root-grafting 
distance of previously affected trees 
(Table 2). The affected elms that were 
beyond root-grafting distance of previ- 
ously affected elms were widely and 
randomly scattered at distances of 50 
feet [15 m] to more than 1,000 feet 
[300 m] from previously diseased trees. 


With the appearance of Dutch elm 
disease in a single elm in the 800 block 
of West Pennsylvania Avenue in south- 
west Urbana in 1951, our data on the 
incidence of phloem necrosis were ex- 
amined to determine the influence of 
phloem necrosis on the appearance and 
early spread of Dutch elm disease. The 
relationship of phloem necrosis to the 
appearance and early spread of Dutch 
elm disease is indicated by the fact that 
phloem necrosis-affected elms can har- 
bor both the smaller European elm 
bark beetle and the Dutch elm disease 
fungus. Data originally gathered in 
Urbana by E. B. Himelick in 1952 
showed that 8 of 40 elms dying from 
phloem necrosis were infested with the 
bark beetles and the fungus (Campana 
1954:358). Apparently the Dutch elm 
disease fungus had been introduced 
into the phloem necrosis-affected elms 
by the bark beetles. 


The population of the smaller Euro- 
pean elm bark beetle is largely deter- 
mined by the amount of elm material 
available for colonization. Since there 
was no community-wide program for 
the control of either disease in Cham- 
paign-Urbana, most of the diseased 


Elms Beyond Root-Grafting 
Distance of Previously 
Diseased Trees 


Percent Number Percent 
32.3 243 67.7 
33.5 369 66.5 
33.0 612 67.0 


trees were left standing for several 
weeks, months, or a year or more after 
they had died. These trees served as 
abundant colonizing locations for the 
bark beetles. The number of standing 
dead elms in June and September in 
1951, 1952, and 1953 (Table 3) was 
greater than the number of elms wilting 
from phloem necrosis in these same 
months except in June 1951. Under 
these conditions millions of the smaller 
European elm bark beetles were pres- 
ent in the area and were potential car- 
riers of the Dutch elm disease fungus. 


Table 3.—Number of phloem _necrosis- 
affected elms, wilting and dead, standing in 
Champaign-Urbana, Illinois, in June and Sep- 
tember 1951, 1952, and 1953. 


Elms Dead 
from Phloem 
Necrosis 


Elms Wilting 
from Phloem 
Necrosis 


Year 


June September June September 


1951 213 187 218 141 
1952 324 558 239 316 
1953 589 441 193 195 
Total 1,126 1,186 650 652 


APPEARANCE OF DUTCH ELM 
DISEASE AND LOCATIONS OF ELMS 
HAVING. DUTCH ELM DISEASE, 
1951-1953 


Dutch elm disease first appeared in 
the Champaign-Urbana area in 1951 
when a single affected elm was found 
in the 800 block of West Pennsylvania 
Avenue in southwest Urbana. We veri- 
fied the disease by culturing the fungus 
from samples of the brown, discolored 
sapwood of wilted branches. Phloem 
necrosis also spread to this immediate 
area in 1951 when two elms showed 
symptoms of that disease. 


ee 


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et 


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125 


= 


aspasip wia yong = % 


area in 1952. Of these 11 elms, 3 
were adjacent to elms that were wilting 


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CARTER & CARTER: PHLOEM NECROSIS AND DurcH ELM DISEASE 


Se POND 


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126 


from or had been killed by phloem 
necrosis. Of these three, one was ad- 
jacent to a phloem necrosis-affected elm 
that wilted in 1950 and 1951 and was 
dead by 1952. One was adjacent to an 
elm that wilted in 1950, and one was 
adjacent to two elms. One of these two 
elms wilted in 1950, and the other 
wilted in 1951. The remaining eight 
elms affected by Dutch elm disease in 
1952 were isolated trees that were not 
adjacent to phloem necrosis-affected 
elms or to the one elm affected by 
Dutch elm disease in 1951. 


ILt1noIs NATURAL History Survey BULLETIN 


me nl 


Vol. 31, Art. 4 


The locations of 164 elms affected by 


Dutch elm disease in 1953 and their 
relation to phloem _necrosis-affected 
elms are shown in Fig. 6 (some loca- 
tions represent more than one tree). 
Twenty-seven locations of Dutch elm 
disease-affected elms were in areas 
where phloem necrosis was abundant, 
as indicated by the three areas enclosed 
by heavy lines in Fig. 6. Ten other 
locations were immediately adjacent to 
one or more phloem necrosis-affected 
elms. The locations of the remaining 
41 Dutch elm disease-affected trees 


No cross street between the 600 and 700 block 


MICHIGAN AVENUE 


° Teri) 
LO 7-155 
ears eee 


Onl toe 


This tree had | 
phloem necrosis 


~<z— \@ —— BUSEY AVENUE 


Fig. 
Michigan Avenue, Urbana, IIlinois. 


ee 
7.—The spread of Dutch elm disease, 1955-1959, in the 700 block of West 


May, 1974 


were not adjacent to phloem necrosis- 
affected elms but were widely scattered 
throughout much of the Champaign- 
Urbana area. 

Only one of the 176 elms affected 
by Dutch elm disease during this 3-year 
period, 1951-1953, was within root- 
grafting distance of an elm previously 
affected by Dutch elm disease. This 
one elm wilted in 1953 and was ad- 
jacent to an elm that wilted in 1952. 
Therefore, the Dutch elm disease fun- 
gus was carried to the remaining 175 
elms by elm bark beetles. 


SPREAD OF DUTCH ELM DISEASE 
IN THE 700 BLOCK OF 
WEST MICHIGAN AVENUE, 
URBANA, 1955-1959 

Elms began dying in the 700 block 
of West Michigan Avenue, Urbana, in 
1955, when four elms wilted, three from 
Dutch elm disease and one from phloem 
necrosis. Elms affected by Dutch elm 
disease before 1955 within one block 
of this area were: one in 1951, two 
in 1953, and three in 1954. Phloem 
necrosis did not occur within one block 
of this area before 1955. 


There were 29 American elms (23 
in the parkway and 6 on private prop- 
erty) in this area in June 1955 when 
the first 3 showed wilt symptoms of 
Dutch elm disease. The location of 
each tree is indicated in Fig. 7. The 
date is shown when wilt symptoms 
were first observed on each tree. The 
annual loss of elms is given in Table 4. 


Of the 28 elms killed in this block 
by Dutch elm disease in 5 years, more 
(10 trees) were killed in the third year 
than in any other year of the study. 
No elm within root-grafting distance of 
previously diseased trees showed wilt 
symptoms in the second year. Thirteen 
(46 percent) of the trees were within 
root-grafting distance of previously dis- 
eased trees. Therefore, they may have 
become infected through roots grafted 
to those of previously diseased elms. 
The remaining 15 elms (54 percent), 
not within root-grafting distance of pre- 
viously diseased trees, became infected 


Table 4.—Elms killed annually by Dutch elm disease in the 700 block of West Michigan Avenue, Urbana, IHlinois, 1955-1959. 


Elms Beyond Root-Grafting 


Elms Within Root-Grafting 


Distance of Previously 


Distance of Previously 


Elms Killed 
by Dutch Elm Disease 


Number 


Diseased Trees 


Diseased Trees 


Residual Elm 


Percent 


Number 


Percent 


Number 


Percent 


Population 


Year 


Carter & CARTER: PHLOEM NEcROSIS AND DuTcH ELM 


Residual 


Population Population 


Original 


Residual 


Population Population 


Original 


Residual 
Population 


Original 
Population 


DISEASE 


OMIM NA 


Total or 


a4 


46 


13 


100 


percent 


127 


8 One elm died from phloem necrosis in 1955 and is not included in this table. 


128 IntInoIs NATURAL History SurRvEY BULLETIN Vol. 31, Art. 4 


through insect transmission of the fun- = 38 
= as ARS 
SPREAD OF PHLOEM NECROSIS = Par aS 
AND DUTCH ELM DISEASE ON Se leat evens 
SPRINGFIELD AVENUE, E |= |e a2 
Ells 2 
CHAMPAIGN, 1955-1959 5 ||5 = BSIN“Aes § 
n pest 
Elms in a six-block section of East os ||\8s| 6s rt 
Springfield Avenue, Champaign, be- 2 allie fl . 
tween First and Wright Streets, began Z mars 
dying in 1955 when one elm wilted from ae) Fe aint Fe 
Dutch elm disease. Phloem necrosis did Ps iS 
not appear in this area until 1957, when = 
one elm contracted that disease. Sev- a 
eral elms within three blocks of Spring- % 
field Avenue were attacked by each of uw a8 
these diseases before 1955. Two elms x By l|rwaevs 
died from phloem necrosis in 1953, one 2 & |e a3 = 
located two blocks northwest of the = @ | | o 
Om |e 
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we ae 2 25 
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wn 
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d ST. = 
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DED Esi4-srt E a3 
9-6-58—* in a3 S933 7 
aS: [ aS 
: in. 
Sa] Ist ST. eo om =} a 
Fig. 8.—The spread of phloem necrosis ao 5 § 
a ae Dutch elm disease (DED) on East eT as i 2 
pringfield Avenue, Ch i inoi inp Or 0D Sd 
TOURS BSG are eae mee o é Bae Ss ~ 
; = al aan a ds 


ee 


May, 1974 


100 block and one located two and one- 
half blocks south of the 600 block of 
East Springfield Avenue. Twelve elms 
died from Dutch elm disease, two in 
1953 and ten in 1954. One of the two 
elms that died in 1953 was two blocks 
west and one was one block north of 
the 100 block of East Springfield Ave- 
nue. Of the 10 elms that died in 1954, 
7 were one block distant from the 100 
and 200 blocks, and 3 were two blocks 
from the 400 block of East Springfield 
Avenue. 

The parkway along this six-block 
stretch was lined with 40 Moline elms 
in 1950. Each tree was approximately 
1 foot [0.3 m] in trunk diameter. Seven 
larger American elms along the same 
section were located on private prop- 
erty. The locations of these 47 elms 
are indicated in Fig. 8. The 25 elms 
that wilted in 1955 on streets that cross 
East Springfield Avenue were beyond 
root-grafting distance of the elms along 
Springfield Avenue. All 47 elms on 


Pi.” 


PEE Epa i i We is 2 
t X \s i 
P y ¢) * : * 
Oh 0/4 0/ e 
iy ea, y . 
\ WZ BN 
\ Nt ip; Vi © 


Fig. 9.—Moline elms on the south side of the 100 block of East Springfield Avenue, 


CARTER & CARTER: PHLOEM NECROSIS AND DUTCH ELM DISEASE 129 


East Springfield Avenue became _af- 
fected by phloem necrosis or Dutch 
elm disease and died within 5 years, 
1955-1959. 

Of the 47 elms killed in 5 years along 
East Springfield Avenue, Champaign, 
17 (36 percent) were killed in 2 years 
by phloem necrosis, while 30 (64 per- 
cent) were killed in 5 years by Dutch 
elm disease. The disease involved and 
the year that each tree became diseased 
are shown in Table 5. 

A rapid increase in the number of 
trees affected by each disease occurred 
in the year following the appearance 
of phloem necrosis and in the third 
year following the appearance of Dutch 
elm disease. Two of the trees killed 
by phloem necrosis on the south side 
of the 100 block of East Springfield 
Avenue in 1958 were within root-graft- 
ing distance of a tree killed by phloem 
necrosis in 1957 (Fig. 9). Six of the 
trees on East Springfield Avenue killed 
by Dutch elm disease between 1955 and 


RRC 
XS i 28, ‘ 
he 
NMA 

Q 
VAL. ‘ 


Pia 0 — 


Champaign, Illinois. The first four elms on the right were killed by phloem necrosis. The 
fifth elm and one elm east of this fifth tree, removed before the picture was taken, were 
killed by Dutch elm disease. The second elm from the right died in 1957, and the others 
died in 1958. (Photo by W. E. Clark) 


130 


1959 were within root-grafting distance 
of trees killed by Dutch elm disease 
in the previous 1 or 2 years. Three of 
these six elms were on the north side 
of the 200 block, one on the north side 
of the 100 block, one on the south side 
of the 300 block, and one on the south 
side of the 500 block. 

Since 39 of the 47 elms in this area 
were not within root-grafting distance 
of previously diseased trees, the spread 
of each disease occurred mainly by 
insect transmission of each causal agent. 


PHLOEM NECROSIS-AFFECTED 
ELMS THAT LATER SHOWED 
SYMPTOMS OF DUTCH ELM 
DISEASE 

Some elms affected by phloem ne- 
crosis subsequently became affected by 
Dutch elm disease before dying. The 
number of elms that showed phloem 
necrosis symptoms followed by Dutch 
elm disease symptoms from 1954 to 
1960 is listed in Table 6. During this 
7-year period 1,234 trees were affected 
by phloem necrosis and 10,714 trees 
were affected by Dutch elm disease. 
The data are arranged according to the 
time between the first appearance of 
phloem necrosis symptoms and the 
subsequent appearance of Dutch elm 
disease symptoms. 

Of the 153 elms affected by both 
diseases, 28 (18.3 percent) were af- 
fected by Dutch elm disease in Sep- 


Table 6.—Phloem necrosis-affected elms in Champaign-Urbana, 
960. 


quently contracted Dutch elm disease, 1954-19) 


ILtInoIs NATuRAL History Survey BULLETIN 


tember following the appearance of 


Vol. 31, Art.4 


La 


< 


phloem necrosis symptoms in the pre- — 


vious June, 123 (80.4 percent) 

the year following the appearance of 
phloem necrosis symptoms, and only 
two (1.3 percent) in the second year 
following the appearance of phloem 
necrosis symptoms. In general, the 
percentage of phloem necrosis-affected 


elms that subsequently showed Dutch ~ 


elm disease increased as the number of 
elms affected by Dutch elm disease 
increased. Many elms that become 


- 


infected with the Dutch elm disease 


fungus in June wilt during the same 
growing season. Therefore, it is not 
surprising that 28 elms that showed 


phloem necrosis symptoms in June © 


showed Dutch elm disease symptoms 
by the following September. Elms 


which had been wilting from phloem ~ 


necrosis in June and were wilting from 


Dutch elm disease in September most — 


likely became infected by the Dutch 
elm disease fungus in June, when 


phloem necrosis symptoms were already — 


evident. 


INCIDENCE OF PHLOEM NECROSIS © 


AND DUTCH ELM DISEASE IN 
CHAMPAIGN-URBANA, 
1944-1972 


To determine the incidence of phloem 
necrosis and Dutch elm disease in 
Champaign-Urbana, the number of 
elms affected annually by each disease 


Illinois, that subse- 


Elms Showing 


Phloem Necrosis-Affected 
Elms That Later Showed 


Year Elms Affected by Dutch Elm Disease Symptoms Dutch Elm Disease 
Phloem Necrosis Symptoms 
Same Year® 1 Year Later 2 Years Later Number Percent 
1954 179 13 13 1.3 
1955 123 ae oe steele 
1956 60 1 5 6 10.0 
1957 368 1 42 2 45 12.2 
1958 344 26 45 70 20.3 
1959 148 #5 17 17 11.5 
1960 12 1 1 2 16.7 
Total or 
percent 1,234 28 123 2 158 12.4 


* Trees listed in this column showed phloem necrosis symptoms in the June survey and Dutch 
elm disease symptoms in the following September survey. 


131 


PHLOEM NECROSIS AND DuTcH ELM DISEASE 


CarTER & CARTER: 


May, 1974 


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132 Intinois NaturAL History SurvEY BULLETIN 


was recorded from 1944, when phloem 
necrosis first appeared, through 1972. 
The number of elms affected annually 
and the percentages of the original 
and residual elm populations lost each 
year are recorded in Table 7. The 
percentages of the original elm popula- 
tion lost annually to each disease are 
illustrated in Fig. 10. The percentages 
of the residual elm population lost an- 
nually to each disease are illustrated 
in Fig. 11. 


Phloem Necrosis 


A rapid increase in the number of 
elms affected by phloem necrosis did 
not occur until 1949, 5 years after the 
disease had first appeared in two Ur- 
bana elms. The number of elms af- 
fected by phloem necrosis in Cham- 
paign-Urbana increased annually until 
1952, when 555 were affected. Follow- 
ing 1952 the number of phloem ne- 
crosis-diseased trees decreased annually 
until 1956, when only 60 elms were 
killed. This decrease was followed by 
an increase to 368 affected trees in 
1957. After 1957 the number of phloem 
necrosis-diseased trees decreased rap- 
idly until 1960, when only 12 elms were 


PERCENT 
@ 


———— Phloem necrosis 
Dutch elm disease 


0) 
1944 1946 1948 1950 1952 1954 1956 


Vol. 31, Art.4 
¥ 


affected. From 1961 through 1972 only — 
one to five elms were affected by 
phloem necrosis annually except in 
1969, when no phloem necrosis oc- — 
curred. From 1944 through 1972, a 
period of 29 years, 2,994 elms were 
killed by phloem necrosis. 

The two peak periods of elm deaths 
from phloem necrosis occurred in 1952 — 
and 1957 (Fig. 10). The cause for the — 
high death rate of elms in 1952 and — 
the subsequent decrease in the inci- — 
dence of phloem necrosis through 1956 — 
was not determined. However, four 
conditions that may have been involved — 
were (1) the rapid increase in the 
incidence of Dutch elm disease, (2) 
the reduction in the elm population, 
(3) the time required for symptoms 
of phloem necrosis to appear following 
infection, and (4) drought conditions 
from 1952 through 1955. 

The incidence of Dutch elm disease 
increased from 11 trees in 1952 to 1,805 
trees in 1955. During this period Dutch 
elm disease reduced the elm population 
by 2,674 trees, or 20 percent of the 
elm population of 1952. 

The time required for wilt symptoms 
to appear following infection is longer 


1958 1960 1962 1964 1966 1968 1970 1972 
R 


Fig. 10.—Annual percentages of the original population of 14,103 elms lost to phloem 
necrosis and Dutch elm disease in Champaign-Urbana, Illinois, 1944-1972. 


May, 1974 


for phloem necrosis than for Dutch elm 
disease. Phloem necrosis-affected elms 
do not show foliage symptoms for at 
least 1 year following infection, while 
many Dutch elm disease-affected elms 
show foliage symptoms in the year 
when infection occurs. 


Drought conditions that prevailed 
from 1952 through 1955, especially in 
1953 and 1954, may have caused a re- 
duction in the elm leafhopper popula- 
tion, since this insect is adversely af- 
fected by such conditions. Precipitation 
for the 5-month growing season, May 
through September, was 7.89 inches 
[200.41 mm] below normal in 1953 and 
4.19 inches [106.41 mm] below normal 
in 1954. Total precipitation was 10.34 
inches [262.64 mm] below normal in 
1953 and 6.73 inches [170.94 mm] be- 
low normal in 1954 (Carter 1955:40). 
Collection records of the Section of 
Faunistic Surveys and Insect Identifica- 
tion, Illinois Natural History Survey, 
show that the populations of leafhop- 
pers in general were drastically reduced 
during these drought years and that the 
elm leafhopper has never been col- 
lected in abundance in Illinois. 


The increase in the incidence of 


60 


50: —-—— Phloem necrosis 


Dutch elm disease 


PERCENT 


0 
1944 1946 1948 1950 1952 1954 1956 


1958 1960 1962 1964 1966 1968 
YEAR 


CarTER & CARTER: PHLOEM NECROSIS AND DuTcH ELM DISEASE 133 


phloem necrosis from 1957 through 
1959 may have been influenced mainly 
by an increase in the elm leafhopper 
population, which might be expected in 
years of near-normal rainfall. Following 
1959 the incidence of phloem necrosis 
decreased to very low levels during 
the period of rapid decline in the 
residual elm population. Only 7 per- 
cent of the original elm population 
remained by 1960. 


Dutch Elm Disease 


A rapid increase in the number of 
elms affected by Dutch elm disease 
started in 1953, 2 years after the disease 
first appeared in Urbana in a single 
elm. The number of elms affected an- 
nually by Dutch elm disease increased 
until 1957, when 2,116 trees were af- 
fected. During the 5-year period 1955- 
1959, the annual loss of elms to Dutch 
elm disease was 1,770-2,116, with a 
total loss of 9,331 trees. This number 
was 82.4 percent of the residual elm 
population of 11,324 trees in the spring 
of 1955, or 66.2 percent of the original 
elm population of 14,103 trees in 1944. 


The peak incidence of Dutch elm 
disease (Fig. 10) occurred in 1957, 6 


1970 1972 


Fig. 11.—Annual percentages of the residual population of elms lost to phloem necrosis 
and Dutch elm disease in Champaign-Urbana, Illinois, 1944-1972 


134 ILtINoIs NATURAL History SURVEY BULLETIN 


years after the disease first appeared 
in the area. The number of elms killed 
in 1957 was 2,116, or 15 percent of the 
original elm population and 28.21 per- 
cent of the residual elm population. 
Following 1959 the residual elm popula- 
tion decreased rapidly until 1963, when 
only 95 healthy elms remained and 
only 9 were killed by Dutch elm dis- 
ease. With the rapid decline in the 
elm population, the incidence of Dutch 
elm disease also declined drastically. 
From 1964 to 1969 only one to four 
trees (none in 1967) became affected 
by this disease annually. No elms were 
affected in 1970 and 1971, but five 
elms were affected in 1972. 


Figure 10 shows that the incidence 
of Dutch elm disease increased more 
rapidly, reached a much higher peak, 
and decreased more rapidly than did 
the incidence of phloem necrosis. Addi- 
tionally, only one peak period of elm 
deaths from Dutch elm disease oc- 
curred, while there were two peak 
periods of elm deaths from phloem 
necrosis. 


Effect of Each Disease on the 
Residual Population of Elms 


The annual loss of elms from each 
disease in relation to the residual elm 
population is illustrated in Fig. 11. The 
annual loss from phloem necrosis fluctu- 
ated from year to year, reaching five 
peaks of 5 percent or more of the 
residual elm population. The highest 
peak (9.43 percent) occurred in 1971, 
when the residual elm population was 
only 53 trees. The next highest peak 
occurred in 1958, when the residual elm 
population was 5,016 trees. 


Following the appearance of Dutch 
elm disease in 1951, the percentage of 
the residual elm population killed an- 
nually by this disease increased rapidly 
until 1960. In 1960 Dutch elm disease 
killed 689 trees (72.53 percent) of the 
residual elm population of 950 trees. 
Following 1960 the annual loss of trees 
in the residual elm population de- 
creased rapidly until 1964 when only 


Vol. 31, Art. 4 ; 


1 elm (1.22 percent) of the residual 
population of 82 was killed by Dutch 
elm disease. 


Accumulated Percentages of Elms 
Killed by Each Disease, 1944-1972 


The accumulated percentages of elms 
killed by each disease are given in 
Table 8 and illustrated in Fig. 12. The 
loss of elms from each disease follows 
a sigmoid curve. Less than 10 percent 
of the elm population was killed by 
both diseases from 1944 through 1952, 
a period of 9 years. Phloem necrosis 
was present throughout the 9-year pe- 
riod, but Dutch elm disease was present 
for only 2 years. During the second 
9-year period, 1953-1961, both diseases 
killed more than 89 percent of the 


Table 8.—Accumulated percentages of elms 
killed by phloem necrosis and Dutch elm dis- 
ease in Champaign-Urbana, Illinois, from 1944 
through 1972. 


Percent of Elms Killed by 


Year Phloem Dutch Elm 
Necrosis Disease 

1944 0.01 

1945 0.02 

1946 0.04 

1947 0.06 

1948 0.11 

1949 0.82 

1950 3.03 aan 
1951 5.58 0.01 
1952 9.52 0.09 
1953 12.27 1.25 
1954 13.54 6.17 
1955 14.41 18.97 
1956 14.83 31.99 
1957 17.44 46.99 
1958 19.88 59.54 
1959 20.93 72.33 
1960 21.02 17.22 
1961 21.04 78.06 
1962 21.05 78.28 
1963 21.07 78.34 
1964 21.10 78.35 
1965 21.11 78.38 
1966 21.12 78.39 
1967 21.14 78.39 
1968 21.15 78.39 
1969 21.15 78.40 
1970 21.17 78.40 
1971 21.21 78.40 
1972 21.23 78.44 


May, 1974 CarTER & CARTER: PHLOEM NEcROSIS AND DutcH ELM DISEASE 135 
100 -—  *———_ -——- - —— Phloem necrosis and 
ae Dutch elm disease 


90: 
80 
70 
60 


50: 


PERCENT 


40: 


0 
1944 


1948 1952 1956 


YEARS 


_-- 


1960 


Dutch elm disease 


Sieh See Se a= = — Phloem necrosis 


1964 1968 1972 


Fig. 12.—Accumulated percentages of the original population of 14,103 elms lost each 
year to phloem necrosis and Dutch elm disease in Champaign-Urbana, Illinois, 1944-1972. 


original elm population. With less than 
1 percent of the elms remaining after 
18 years, the annual loss became a few 
trees. Phloem necrosis killed 2,994 elms 
during the 29-year period, 1944-1972, 
while Dutch elm disease killed 11,062 
elms during the 22-year period, 1951— 
1972. Although no elms showed symp- 
toms of Dutch elm disease in 1970 or 
1971, five trees were killed by the dis- 
ease in 1972. Of the original popula- 
tion of 14,103 elms in 1944, all but 40 
had been killed by the fall of 1972, a 
period of 29 years. 


TIME OF YEAR IN WHICH ELMS 
DIED FROM PHLOEM NECROSIS 
AND DUTCH ELM DISEASE 


Starting in 1951 two surveys were 
made annually to record the number of 
elms affected by phloem necrosis and 
by Dutch elm disease. The first survey 
was made in the early part of the 
growing season, usually in June. The 
second survey was made in the late 
part of the growing season, usually in 
September. These surveys are referred 
to as the June and September surveys. 

Each year data were obtained on the 


Table 9.—Elms showing disease symptoms in June and September surveys, Champaign- 


Urbana, Illinois, 1955-1961. 


Elms Having Phloem Elms Having Dutch Elm Total Standing 

Necrosis Symptoms Disease Symptoms Wideawed Dead Elms 
Year “June September Total June September Total pines June September 

Survey Survey Survey Survey Survey Survey 

1955 40 81 121 1,059 746 1,805 1,926 86 133 
1956 21 39 60 1,162 674 1,836 1,896 160 528 
1957 44 324 368 1,159 957 2,116 2,484 255 520 
1958 217 127 344 1,049 721 1,770 2,114 447 1,323 
1959 58 90 148 1,116 688 1,804 1,952 1,064 1,509 
1960 4 8 12 496 193 689 701 608 341 
1961 2 1 3 92 27 119 122 ae age 
Total 386 670 1,056 6,133 4,006 10,139 = =11,195 2,620 4,354 


136 ILLINOIS NATURAL History SuRvEY BULLETIN Vol. 31, Art. 4 


number of elms affected by each disease 
in each of the two survey periods. No 
affected elm was included in the data 
of more than one survey. The num- 
bers of elms affected by each disease 
at the time of each June and September 
survey from 1955 through 1961 are 
recorded in Table 9. 


Of the 1,056 elms affected by phloem 
necrosis during this 7-year period, 386 
(36.6 percent) showed symptoms in 
June, and 670 (63.4 percent) showed 
symptoms in September. Of the 10,139 
elms affected by Dutch elm disease, 
6,133 (60.5 percent) showed symptoms 
in June, and 4,006 (39.5 percent) 
showed symptoms in September. Of 
the 6,974 dead elms standing at the 
time of the June and the September 
surveys during the years 1955-1960, 
2,620 (37.6 percent) were standing in 
June and 4,354 (62.4 percent) were 
standing in September. 


Without 
79 
66 
89 
169 
273 
144 
820 


Elms That Died 
in Winter 
tember Showing Wilt 


Sep- 


2 Years Later 


1 Year Later 


PERIOD OF TIME IN WHICH ELMS 
DIED FOLLOWING THE 
APPEARANCE OF FOLIAGE WILT 
OF PHLOEM NECROSIS OR 
DUTCH ELM DISEASE 


Not all elms affected by phloem ne- 
crosis or Dutch elm disease die in the 
same growing season in which foliage 
wilt first appears. Some elms wilt dur- 
ing two, and occasionally three, grow- 
ing seasons before they die; some elms 
that have shown no foliage wilt die 
during the winter. Data on 231 elms 
affected by phloem necrosis, 3,908 elms 
affected by Dutch elm disease, and 820 
elms that died but showed no foliage 
wilt are given in Table 10. The data 
cover the growing seasons of 1955 
through 1960. During this period most 
diseased elms were not removed until 
after they were dead. This practice 
made it possible to record the period 
of time in which the diseased trees died. 

Of the 231 phloem necrosis-affected 
elms, 143 (61.9 percent) died in the 
year following the appearance of wilt. 
Only 87 (37.7 percent) died in the year 
that wilt appeared, and only 1 (0.4 
percent) lived until the second year fol- 


Elms Showing Dutch Elm Disease Wilt That Died 


Same Year 


2 Years Later 


June 


1 Year Later 
Sep: 
tember 
19 
17 
1 
87 


in June 


Not 


Elms Showing Phloem Necrosis Wilt That Died 
Wilting June 


Same Year 


in June 
13 
11 
14 


Table 10.—The period of time jin which elms died following the appearance of wilt symptoms of phloem necrosis or Dutch elm disease, 
Wilting 


Champaign-Urbana, Illinois, 1955-1960. 


Year 


May, 1974 


lowing the appearance of wilt. Nearly 
half (45.9 percent) of the elms affected 
by phloem necrosis continued to live 
during the remainder of the summer in 
which wilt symptoms appeared but 
were dead by June of the next summer. 


Of the 3,908 Dutch elm disease- 
affected elms, 3,134 (80.2 percent) died 
in the year when wilt symptoms ap- 
peared. Only 753 (19.3 percent) died 
during the second year, and 21 (0.5 
percent) died during the third summer. 
These data indicate that, following the 
initial appearance of foliage wilt, elms 

having Dutch elm disease die more 
rapidly than do elms affected by 
phloem necrosis. 


Also of interest are the 820 elms 
that did not show foliage wilt in the 
September survey but died before the 
June survey of the following year. 
These elms represent 16.5 percent of 
the 4,959 elms that died during the 
6-year period. Although the cause of 
death was not determined, probably 
most, if not all, of these elms were 
killed by Dutch elm disease, since this 
disease usually causes elms to die more 
rapidly than does phloem necrosis. 


DISCUSSION 


Phloem necrosis was not known to 
occur in any areas close to Champaign- 
Urbana when the disease was discov- 
ered in two adjacent elms in Urbana in 
1944, The nearest area where the dis- 
ease had occurred was Danville, Ili- 
nois, 32 miles [51 km] east of Urbana. 
Beginning in 1935, elm plantings be- 
_ tween Urbana and Danville had been 
observed for disease symptoms fre- 
quently during the growing season of 
each year. 


In the course of this study careful 
_ examination of phloem samples from 
wilting elms showed that the charac- 
_ teristic butterscotch color usually was 
present only in the current phloem. 
However, samples from some wilting 
elms had butterscotch color in 1- and 
sometimes 2-year-old phloem. This 
condition occurred mainly in elms that 


CarTER & CARTER: PHLOEM NECROSIS AND DuTCH ELM DISEASE 137 


showed foliage symptoms during two 
or more growing seasons. Since the but- 
terscotch color in the current phloem 
indicates that the tree has been infected 
for about 1 year (Baker 1949:730), the 
butterscotch color in 1- and 2-year-old 
phloem indicates that the MLO is in 
some elms for 2-3 years before foliage 
wilt appears. 


Because phloem necrosis was present 
in the Champaign-Urbana area 7 years 
(1944-1950) before Dutch elm disease 
appeared there, the initial spread of 
phloem necrosis was not influenced by 
Dutch elm disease. In Urbana, phloem 
necrosis spread slowly for 4 years fol- 
lowing its appearance in 1944. The 
few affected trees were widely scattered 
at distances of approximately 300-2,000 
feet [100-600 m] from the two elms 
first attacked. Each affected tree rep- 
resented a new center of infection from 
which the disease continued to spread 
to nearby elms. This intitial spread 
resulted from transmission of the MLO 
by the elm leafhopper. 


The incidence of phloem necrosis in- 
creased rapidly in Urbana during 1948 
and 1949, and by 1950 over 300 trees 
were affected. The disease was con- 
centrated mainly within an area ap- 
proximately 1,400 feet [400 m] wide 
and 4,000 feet [1,200 m] long in the 
central part of the city, an area heavily 
populated with American elms. By 
1950 only a few scattered elms were 
affected beyond this area, and no af- 
fected trees were in the 3,600-foot-wide 
[1,100 m] area adjacent to Champaign. 

In Champaign phloem necrosis was 
not found until 1948, when it affected 
four elms. The trees were approxi- 
mately 5,000 feet [1,500 m] west of any 
affected trees in Urbana. Following 
1948 the number of elms affected by 
phloem necrosis increased rapidly in 
Champaign, and all but a few of the 
affected trees were concentrated in an 
area two blocks wide and four blocks 
long, centering around the 700 block 
of South Lynn Street. 

Of the 36 elms in the 700 block 
of South Lynn Street, 8 wilted in 1949. 


138 ILtiInoIs NATURAL History SURVEY BULLETIN 


The infection of these eight elms re- 
sulted from insect transmission of the 
MLO in 1948 or earlier. Because 28 
of the elms in this block wilted in the 
2 years following the initial appearance 
of the disease, it is possible that the 
MLO was spread by the elm leaf- 
hopper. However, all but three of these 
elms were within root-grafting distance 
of previously affected trees. Following 
1950 phloem necrosis spread rapidly 
throughout both cities, and the greatest 
loss of trees from this disease in any 
1 year occurred in 1952, 8 years after 
the disease first appeared. 


Phloem necrosis was widespread 
when Dutch elm disease was dis- 
covered in one elm in southwest Ur- 
bana in 1951. The infection of this 
elm resulted from insect transmission 
of the fungus. However, phloem ne- 
crosis-affected elms may have harbored 
the inoculum, because the Dutch elm 
disease fungus was isolated from 8 
of 40 elms that had phloem necrosis 
in 1952 (Campana 1954:358). There- 
fore, many of the hundreds of elms 
killed annually by phloem necrosis but 
not removed promptly served as col- 
onizing sites for the smaller European 
elm bark beetle, vector of the Dutch 
elm disease fungus. Millions of these 
insects were present in the Champaign- 
Urbana area as potential carriers of the 
Dutch elm disease fungus at the time 
the disease first appeared. 


Following the appearance of Dutch 
elm disease in 1951, the fungus was 
transmitted by insects in 1952 and 1953, 
for only 1 of 175 diseased elms was 
within root-grafting distance of a previ- 
ously diseased tree. Although only a 
few elms were affected in 1952, Dutch 
elm disease spread rapidly in the next 
3 years, and a 5-year peak period of 
elm deaths started in 1955. Dutch elm 
disease increased annually more rapidly 
than did phloem necrosis in the number 
of elms affected and in the number of 
infection centers. As the incidence of 
Dutch elm disease increased, the inci- 
dence of phloem necrosis decreased, 


Vol. 31, Art. 4 | 


and phloem necrosis failed to spread 
along some streets. 


During the peak years of loss from ) 
each disease, Dutch elm disease killed 
approximately four to five times as} 
many elms as did phloem necrosis. The » 
peak of elm deaths from Dutch elm) 
disease occurred over 5 years, while ° 
the peak of elm deaths from phloem 
necrosis occurred in two periods, the ° 
first lasting 4 years and the second 2 
years. Following these peak periods the » 
numbers of elms affected annually by | 
each disease decreased rapidly, for over 
90 percent of the 1944 elm population 
of Champaign-Urbana had been killed 
by 1960. 


In some blocks and along some 
streets all elms were killed within 
3-5 years by one or both diseases. All! 
36 elms in the 700 block of South Lynn } 
Street in Champaign were killed by’ 
phloem necrosis in 3 years, and 28 of 
29 elms in the 700 block of West Michi- - 
gan Avenue in Urbana were killed by. 
Dutch elm disease in 5 years. 


Where both diseases were present, | 
Dutch elm disease killed more trees 5 
than did phloem necrosis. Dutch elm} 
disease tends to kill trees more rapidly / 
than does phloem necrosis. Most elms 
affected by Dutch elm disease die in 
the same year in which foliage wilt| 
appears, but most elms affected by» 
phioem necrosis die in the year follow- 
ing the appearance of foliage wilt. Also, _ 
more phloem necrosis-affected elms * 
show wilt symptoms in September than } 
show them in June, while more Dutch / 
elm disease-affected trees show wilt | 
symptoms in June than show them in 
September. Some elms die during the: 
winter without any visible foliage wilt. 
While each disease may contribute to- 
these winter deaths, it seems likely that 
Dutch elm disease is mainly respon- 
sible. 

Some elms that first showed symp-~ 
toms of phloem necrosis subsequently | 
showed symptoms of Dutch elm dis- 
ease. As the number of elms killed by) 
Dutch elm disease increased, the num- 


May, 1974 


ber of phloem necrosis-affected elms 
subsequently affected by Dutch elm 
disease increased. This fact suggests 
that as the supply of Dutch elm disease 
inoculum increases, more phloem ne- 
crosis-affected elms are invaded by the 
Dutch elm disease fungus. Also, the 
greater the number of elms infested 
with bark beetles, the greater the 
chances for the spread of the Dutch 
elm disease fungus. 


Although some elms affected by 
phloem necrosis in June showed Dutch 
elm disease symptoms in September, 
in most cases Dutch elm disease symp- 
toms did not appear until the year 
following the appearance of phloem 
necrosis symptoms. Only phloem ne- 
crosis-affected elms that die slowly 
during one or more growing seasons 
can be subsequently affected by and 
show symptoms of Dutch elm disease. 
Phloem necrosis-affected elms that are 
subsequently affected by Dutch elm 
disease appear to be killed by Dutch 
elm disease and not by phloem necrosis 
(Campana & Carter 1955). 


The cycle of elm deaths from Dutch 
elm disease probably was affected only 
slightly, if at all, by the presence 
of phloem necrosis. This conclusion is 
based on the fact that elm deaths from 
Dutch elm disease built up to a peak 
more rapidly than did elm deaths from 
phloem necrosis. During the 5-year 
period 1955-1959 more than eight times 
as many elms were killed by Dutch 
elm disease as were killed by phloem 
necrosis. However, the cycle of elm 
deaths from phloem necrosis was 
greatly shortened by the presence of 
Dutch elm disease; Dutch elm disease 
killed 78.4 percent of the elms, while 
phloem necrosis killed only 21.2 per- 
cent. 


SUMMARY 


In the 29-year study reported here, 
data were recorded on the spread of 
and losses caused by elm phloem ne- 
crosis and Dutch elm disease in a mu- 


CARTER & CARTER: PHLOEM NECROSIS AND DuTcH ELM DISEASE 139 


nicipal area which had no community- 
wide control program for either disease. 


Phloem necrosis appeared in Urbana 
in 1944, when two trees were affected. 
Dutch elm disease did not appear until 
1951, when one tree was affected. The 
initial spread of phloem necrosis was 
not influenced by Dutch elm disease, 
since Dutch elm disease was not pres- 
ent during that period. Each of the 14 
elms that contracted phloem necrosis 
from 1945 through 1948 was scattered 
at random beyond root-grafting dis- 
tance of other diseased trees, and each 
tree represented a separate infection 
center. Phloem necrosis spread rapidly 
along some streets, killing all 36 elms 
in one block within 3 years. 


The early spread of Dutch elm dis- 
ease was influenced by phloem necrosis. 
Phloem necrosis-affected elms can har- 
bor the Dutch elm disease fungus, and 
the elms killed by phloem necrosis 
were heavily colonized by the smaller 
European elm bark beetle, vector of 
the Dutch elm disease fungus. Many of 
the phloem necrosis-affected elms were 
not removed before the bark beetles 
emerged. 


Dutch elm disease spread rapidly to 
elms in areas where phloem necrosis 
was abundant, and it also affected 
scattered elms located well away from 
phloem necrosis-affected elms. How- 
ever, of 164 elms having Dutch elm 
disease in 1953 only 41 were in scat- 
tered locations away from phloem ne- 
crosis-affected trees. 


Dutch elm disease, like phloem ne- 
crosis, spread rapidly to elms along 
some streets. Twenty-eight elms were 
killed by this disease in one block in 
5 years. Of 47 elms in six blocks of one 
street, 17 were killed by phloem ne- 
crosis in 2 years and 30 were killed by 
Dutch elm disease in 5 years. However, 
Dutch elm disease was present for 2 
years before phloem necrosis appeared. 
Phloem necrosis and Dutch elm disease 
were spread mainly by their respective 
insect vectors in this area, because 
39 of the 47 elms were beyond root- 


140 ILLINOIS NATURAL History SuRvEY BULLETIN 


grafting distance of previously diseased 
trees. 

Some phloem necrosis-affected elms 
subsequently became infected with the 
Dutch elm disease fungus and showed 
typical symptoms of Dutch elm disease 
before dying. The number of phloem 
necrosis-affected elms that subsequently 
became affected by Dutch elm disease 
increased as the incidence of Dutch 
elm disease increased. 

The greatest number of elms affected 
by phloem necrosis in 1 year was 555 
trees in 1952, 8 years after the disease 
was discovered in this area. The great- 
est number of elms affected by Dutch 
elm disease was 2,116 trees in 1957, 6 
years after the disease was discovered 
here. Of the original population of 
14,103 elms, 2,994, or 21.23 percent, 
were killed by phloem necrosis in 29 
years. Dutch elm disease killed 11,062, 


| 


| 
Vol. 31, Art. 4 _ 
or 78.44 percent, in 22 years. Both dis-— 
eases killed 14,056, or 99.67 percent, of 
the elms. Dutch elm disease had a 
greater effect on the residual elm popu- 
lation, since it killed more than three 
times as many elms as did phloem 
necrosis. 


More elms showed symptoms of 
phloem necrosis in the September sur-_ 
vey than showed such symptoms in the - 
June survey. The reverse was true of 
elms having Dutch elm disease. Fol-— 
lowing the appearance of wilt symp-— 
toms, elms affected by Dutch elm dis- 
ease tended to die more rapidly than | 
did elms affected by phloem necrosis. — 
Most elms that had Dutch elm disease : 
died in the growing season in which | 
foliage wilt appeared, while most elms ; 
that contracted phloem necrosis died | 
in the year following the appearance » 
of foliage wilt. : 


LITERATURE CITED 


Baker, W. L. 1948. Transmission by leaf 
hoppers of the virus causing phloem 
necrosis of American elm. Science 108: 
307-308. 


1949. Studies on the transmission 
of the virus causing phloem necrosis of 
American elm, with notes on the biology 
of its insect vector. Journal of Economic 
Entomology 42:729-732. 


BANFIELD, W. M., EB. G. Rex, and C. May. 
1947. Recurrence of Dutch elm disease in 
American elms in relation to tree stature. 
Phytopathology 37:1-2. 

Bretz, T. W., R. U. Swinetn, and D. E. 
ParKeR. 1945. Some recent observations 
on elm phloem necrosis and the Dutch 
elm disease. National Shade Tree Con- 
ference Proceedings 21:25-28. 


Campana, R. J. 1954. The present status 
of Dutch elm disease in Illinois. Plant 
Disease Reporter 38:356-358. 


1958. Dutch elm disease and elm 
phloem necrosis. Midwestern Shade Tree 
Conference Proceedings 13:17-25. 


, and J. C. Carrer. 1955. Spread of 
Dutch elm disease in Illinois in 1954. 
Plant Disease Reporter 39:245—-248. 


, and 1957. The current sta- 
tus of Dutch elm disease in Illinois. Plant 
Disease Reporter 41:636-639. 


‘Carter, J. C. 1945. Dying of elms in Illi- 
nois. Plant Disease Reporter 29:23-26. 


1950. Status of oak wilt and elm 
phloem necrosis in the Midwest. Arbor- 
ist’s News 15:45-51. 

1954. Elm phloem necrosis — re- 
sumé of the situation. Midwestern Shade 
Tree Conference Proceedings 9:14-16. 


1955. The Champaign-Urbana-Uni- 
versity of Illinois situation. Pages 36-42 
in Control of Dutch elm disease. Proceed- 
ings of a statewide conference on the con- 


141 


trol of Dutch elm disease. Illinois State 
Chamber of Commerce, Chicago. 


. 1961. Dutch elm disease up-to-date. 
Midwestern Shade Tree Conference Pro- 
ceedings 16:34-39. 

Forses, S. A. 1885. Insects injurious to the 
elm. Pages 112-115 in Fourteenth report 
of the state entomologist on the noxious 
and beneficial insects of the state of 
Illinois. 


1912. What is the matter with the 
elms in Illinois? Illinois Agricultural Ex- 
periment Station Bulletin 154. 22 pp. 

HIME ick, E. B., and D. Nreety. 1962. Root 
grafting of city-planted American elms. 
Plant Disease Reporter 46: 86-87. 

NeeEty, D. 1967. Dutch elm disease in Illi- 
nois cities. Plant Disease Reporter 51: 
511-514. 


, and J. C. Carrer. 1965. Species of 
elm on the University of Illinois campus 
resistant to Dutch elm disease. Plant Dis- 
ease Reporter 49:552. 


; , and R. J. Campana. 1960. 
The status of Dutch elm disease in Illi- 
nois. Plant Disease Reporter 44:163-166. 


SwIncte, R. U. 1938. A phloem necrosis of 
elm. Phytopathology 28:757-759. 

1940. Phloem necrosis in the Ohio 

River Valley. Phytopathology 30:23. 

1942. Phloem necrosis: a virus 
disease of the American elm. U.S. De- 
partment of Agriculture Circular 640. 
8 pp. 

VerRALL, A. F., and T. W. GRAHAM. 1935. 
The transmission of Ceratostomella ulmi 
through root grafts. Phytopathology 25: 
1039-1040. 


Witson, C. L., C. E. Seriskar, and C. R. 
Krause. 1972. Mycoplasmalike bodies 
associated with elm phloem necrosis. 
Phytopathology 62:140—-143. 


INDEX 


A 
Alton, 113 
American elm, 113-115, 117, 127, 129, 137 
Asiatic elm species, 115, 118 
Aurora, 115 
Automobile survey, 121 


B 
Bark beetles, 113 
Belleville, 113 
Bloomington, 113, 115 


Borers, 113 
Burlington, Iowa, 113 

C 
Cairo, 113 


Carbondale, 113 

Centralia, 113 

Ceratocystis ulmi, 114 

Champaign, 113, 116-117, 122-125, 127, 
129-130, 132, 137-138 

Charleston, 113 

Chebanse, 114 

Chenoa, 114 

Clayton, 113 

Coles County, 114 

‘Crystal Lake Park, 117 


D 
Danville, 113, 137 
Decatur, 113 
Du Quoin, 113 
Dutch elm disease fungus, 114, 116, 124, 127, 
138-140 
Dutch elm disease symptoms, 114-118, 130 
Dwight, 114 


E 
East-central Illinois, 113-114 
East Springfield Avenue, 128-129 
Edwardsville, 113 
Elgin, 115 
“Elm blight,” 113 
Elm leafhopper, 116, 123, 137 
English elm, 117 
European elm species, 115, 117 


F 
Fairfield, 113 
Faunistic Surveys and Insect Identification 
Section, 133 
First Street, 128 


G 
Galatia, 113 


H 
Hillsboro, 113 
Himelick, E. B., 124 
Hylurgopinus rufipes, 115 


J 
Joliet, 115 
June survey, 120, 135-137, 140 


L 


Leafhopper, 115, 133 
Lincoln, 113 


M 
Mattoon, 113, 115 
McLeansboro, 113 
Melvin, 113 
Midwest, 114 
MLO, 114-116, 123, 137-138 
Moline elm, 117, 129 
Mt. Pulaski, 115 
Mt. Vernon, 113 
Mycoplasmalike organism (MLO), 114 


N 
Normal, 113 
North-central Illinois, 113, 115 


O 
Ohio, 114 
Ohio Valley, 114 
Onarga, 114 

P 
Paris, 113 
Peoria, 113 


Phloem necrosis symptoms, 114, 117, 122, 
130 
Pittsfield, 113 


Q 
Quincy, 113 

R 
Robinson, 113 
Rockford, 114 

Ss 


Scaphoideus luteolus, 115-116 

Scolytus multistriatus, 116 

September survey, 120, 135-137, 140 

Shelbyville, 113 

Slippery elm, 117 

Smaller European elm bark beetle, 116, 124, 
138-139 

South-central Illinois, 113 

South Lynn Street, 122-123, 137-138 

Springfield, 113 

Sumner, 113 

Surveys, 120-121 


Tr 
Taylorville, 113 


U 
Ulmus alata, 115 
Ulmus americana, 115 
University of Illinois campus, 118 
Urbana, 113-114, 116-117, 122-125, 127, 130, 
132-133, 137-139 


Vandalia, 113 
Vector, 116, 123, 138-139 


142 


May, 1974 Carter & CARTER: PHLOEM Necrosis AND DurcH Ex_m DIsmAsE 143 


Virus, 114 


* 
WwW 
Waukegan, 115 
Wayne County, 113 
West-central Illinois, 113 
West Main Street, 122 


West Michigan Avenue, 127, 138 
West Pennsylvania Avenue, 124 
Winged elm, 115 

Wright Street, 128 


Zion, 115 


Mellue fin, 
MERE: iad 
Pd esd 
Wy 


bee 


BULLETIN 

Volume 30, Article 6—Comparative Uptake 
and Biodegradability of DDT and Meth- 
oxychlor by Aquatic Organisms. By Ke- 
turah A. Reinbold, Inder P. Kapoor, Wil- 
liam F. Childers, Willis N. Bruce, and 
Robert L. Metcalf. June, 1971. 12 p., in- 
dex. 


Volume 30, Article 7—A Comparative Study 
of Two Components of the Poinsettia Root 
Rot Complex. By Robert S. Perry. Au- 
gust, 1971. 35 p., index. 

Volume 30, Article 8—Dynamics of Condi- 
tion Parameters and Organ Measurements 
in Pheasants. By William L. Anderson. 
July, 1972. 44 p., index. 

Volume 31, Article 1—The Effects of Sup- 
plemental Feeding and Fall Drawdowns 
on the Largemouth Bass and Bluegills at 
Ridge Lake, Illinois. By George W. Ben- 
nett, H. Wickliffe Adkins, and William 
F. Childers. January, 1973. 28 p., index. 


Volume 31, Article 2—The Reproductive 
Cycle of the Raccoon in Illinois. By Glen 
C. Sanderson and A. V. Nalbandov. July, 
1973. 57 p., index. 


Volume 31, Article 3—Nutritional Respon- 
ses of Pheasants to Corn, with Special 
Reference to High-Lysine Corn. By Ron- 
ald F. Labisky and William L. Anderson. 
July, 1973. 26 p., index. 


BIOLOGICAL NOTES 


78—The Literature of Arthropods Associ- 
ated with Soybeans. II. A Bibliography 
of the Southern Green Stink Bug, Nezara 
viridula (Linneaus) (Hemiptera: Penta- 
tomidae). By N. B. DeWitt and G. L. 
Godfrey. March, 1972. 23 p. 


79.—Combined Culture of Channel Catfish 
and Golden Shiners in Wading Pools. By 
D. Homer Buck, Richard J. Baur, Charles 
F. Thoits III, and C. Russell Rose. April, 
1972. 12 p. 


80.—Illinois Birds: Hirundinidae. By Rich- 
ard R. Graber, Jean W. Graber, and 
Ethelyn L, Kirk. August, 1972. 36 p. 

81.—Annotated Checklist of the Butterflies 
of Illinois. By Roderick R. Irwin and 
John C. Downey. May, 1973. 60 p. 


List of available publications mailed on request 


No charge is made for publications of the ILt1nors Narurat History Survey. A 
copy of most publications will be sent free to anyone requesting it until the supply be 
low. Costly publications, more than one copy of a publication, and publications in § 
supply are subjects for special correspondence. Such correspondence should identi 
writer and explain the use to be made of the publication or publications. 


Address orders and correspondence to the Chief, 
Illinois Natural History Survey 
Natural Resources Building, Urbana, Illinois 61801 


Darters and the Inclusiveness 
Genus Percina, By Lawrence M 
Gregory S. Whitt. May, 1973. 7 p. 
83.—Illinois Birds: Laniidae. By Ric 
Graber, Jean W. Graber, and Ethe 
Kirk. June, 1973. 18 p. 


84.—Interactions of Intensive 
Channel Catfish with Largemouth 
1-Acre Ponds. By D. Homer Bu 
ard J. Baur, and C. Russell 
ruary, 1974. 8 p. ‘ 
85.—The Literature of Arthropo 
ated with Soybeans. III. A Bibliogs 
of the Bean Leaf Beetle, Ceroton 
furcata (Forster) and ©. ruficor 
vier) (Coleoptera: Chrysomelidae} 
P. Nichols, M. Kogan, and G. 
bauer. February, 1974. 16 p. 


86.—Illinois Birds: Tyrannidae. B 
ard R. Graber, Jean W. Grab 
Ethelyn L, Kirk. February, 1974. | 

87.—The Literature of Arthropods : 
ated with Alfalfa. I. A Bibliogra 
the Spotted Alfalfa Aphid, The 
maculata (Buckton) (Homopte 
dae). By D. W. Davis, M. P. Nie! 
E. J. Armbrust. February, 1974. 

88.—The Literature of Arthropods 
ated with Alfalfa. II. A Bibliogr: 
the Sitona Species (Coleoptera: C 
lionidae). By W. P. Morrison, B. 
M. P. Nichols, and E. J. Armbrust. 
ruary,, 1974. (24 ne 5 


CIRCULAR 

46.—Illinois Trees: Their Diseases, 
Cedric Carter. June, 1964. (Third 
ing, with alterations.) 96 p. 

47.—Illinois Trees and Shrubs: Their 
Enemies. By L, L. English. July, 1 
(Fifth printing, with revisions.) 91 p 

51.—Illinois Trees: Selection, Plantin 
Care. By J. Cedric Carter. August, 19 
123 p. me 

52.—Fertilizing and Watering Tree 
Dan-Neely and E. B. Himelick. 
ber, 1971. (Third printing.) 20 p. 

53.—Dutch Elm Disease in Illinois. 
Cedric Carter. October, 1967. 19 p. 


| ILLINOIS 
atural History Survey 


Larvae of the Sericothripini 
(Thysanoptera: Thripidae), 
with Reference to Other Larvae 
of the Terebrantia, of Illinois 


THE LIBRARY OF THE 


eG aS OT 


‘3 INOIS 
{TMENT OF REGISTRATION AND EDUCATION Sa eS NA-CHAMPAIGN 


VOLUME 31, ARTICLE 5 
AUGUST, 1974 


ILLINOIS 


‘atural History Survey 
BULLETIN 


Larvae of the Sericothripini 
(Thysanoptera: Thripidae), 
with Reference to Other Larvae 
of the Terebrantia, of Illinois 


mas C. Vance 


OF ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


URAL HISTORY SURVEY DIVISION 
ANA, ILLINOIS 


VOLUME 31, ARTICLE 5 
AUGUST, 1974 


STATE OF ILLINOIS 


DEAN BarriNGER, Ph.D., Chairman; 
Gurtowsky, Ph.D., Chemistry ; 
Forestry ; 


NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 
SCIENTIFIC AND TECHNICAL STAFF 
GeEorcE SpruGeL, Jr., Ph.D., Chief 
Auice K. Apams, Secretary to the Chief 


Section of Economic Entomology 


Witiram H. Luckmann, Ph.D., Entomologist and Head 
Wiis N. Bruce, Ph.D., Entomologist 
Wayne L. Howe, Ph.D., Entomologist 
STEVENSON Moore, III, Ph.D., Entomologist, Extension 
Howarp B. Perry, Ph.D., Entomologist, Extension 
James E. AppLesy, Ph.D., Associate Entomologist 
Epwarp J. ArmBrust, Ph.D., Associate Entomologist 
Marcos KoGan, Ph.D., Associate Entomologist 
JosepH V. Mappox, Ph.D., Associate Entomologist 
Ronatp H, Meyer, Ph.D., Associate Entomologist 
Rosert D. PauscH, Ph.D., Associate Entomologist 
Raupu E. Securiest, Ph.D., Associate Entomologist 
Joun K. Bouseman, M.S., Assistant Entomologist 
GeEorRGE L. Goprrey, Ph.D., Assistant Entomologist 
WiuiAm G. RuESINK, Ph.D., Assistant Entomologist 
James R. SANBORN, Ph.D., Assistant Entomologist 
DouGuas K. SELL, Ph.D., Assistant Entomologist 
Joun L. WepsBerG, Ph.D., Assistant Entomologist 
CLARENCE E. Wuire, B.S., Assistant Entomologist 
Keun S. Park, M.S., Assistant Chemist 
Sue E, Warkins, Supervisory Assistant 
Donaup E, KuHuMAN, Ph.D., Assistant Professor, 
Extension 
RoscoE RaNnDELL, Ph.D., Assistant Professor, Extension 
Tim Coouey, M.A., Assistant Specialist, Extension 
Kurt E, Repsore, M.S., Assistant Specialist 
Joun F. Watt, M.S., Assistant Specialist, Extension 
Jean G. Wiuson, B.A., Supervisory Assistant 
DanIEL P. Bartett, Ph.D., Research Associate 
Martua P. NicHous, M.S., Research Associate 
Rospert J, Barney, B.S., Research Assistant 
Tzu-Suan Cuu, M.S., Research Assistant 
SrerHen D. Cowan, B.S., Research Assistant 
STepuen K. Evrarp, B.S., Research Assistant 
BarBARA J. Forp, M.A., Research Assistant 
Raymonp A. Korex, M.Mus., Research Assistant 
Rose ANN Meccoul, B.S., Research Assistant 
BARBARA E. PETERSON, B.S., Research Assistant 
KETURAH REINBOLD, M.S., Research Assistant 
SrerHen Roserts, B.S., Junior Professional Scientist 
Joun T, SHaw, B.S., Junior Professional Scientist 
LowE.u Davis, Technical Assistant 
Cuarves G. Heum, M.S., Technical Assistant 
Linpa IsENHOWER, Technical Assistant 
Lu-pinc Len, M.S., Technical Assistant 


Section of Botany and Plant Pathology 


Rosert A. Evers, Ph.D., Botanist 

Eucene B. Himeuick, Ph.D., Plant Pathologist 

R. Dan Neety, Ph.D., Plant Pathologist 

D. F. ScHOENEWEIsS, Ph.D., Plant Pathologist 

J. Levann Crann, Ph.D., Associate Mycologist 
Water Hartstinn, Ph.D., Assistant Plant Pathologist 
Berry S. Neuson, Junior Professional Scientist 

Gene E. Reip, Technical Assistant 


Section of Aquatic Biology 


D. Homer Buck, Ph.D., Aquatic Biologist 

Witiram F, Curiupers, Ph.D., Aquatic Biologist 

R. Weipon Larimore, Ph.D., Aquatie Biologist 

Roper? C. HiLtTiBran, Ph.D., Biochemist 

ALLISON BrigHAM, Ph.D., Assistant Aquatic Biologist 
Warren U. BricHAaM, Ph.D., Assistant Aquatic Biologist 
Ricuarp E. Sparks, Ph.D., Assistant Aquatic Biologist 
Joun TRANQUILLI, M.S., Assistant Aquatic Biologist 
DonaLp W. Durrorp, M.S., Junior Professional Scientist 
Mary FRANCES Martin, Junior Professional Scientist 
Joun M. McNurney, M.S., Junior Professional Scientist 


CONSULTANTS AND RESEARCH AFFILIATES: 
life Research, Southern Illinois University ; 


Illinois; ENTOMOLOGY, Roper? L. Mercanr, Ph.D., 
partment of Zoology, University of Illinois; 
versity of Illinois ; 
of Illinois. 


DEPARTMENT OF REGISTRATION AND EDUCATION ¢ 
BOARD OF NATURAL RESOURCES AND CONSERVATION 
Tuomas Park, Ph.D., Biology; 
B.S.C.E., Engineering ; 
W. L. Everirt, E.E., Ph.D., Representing ‘the President of the University of Illinois ; 
Hapuey, Ph.D., Representing the President of Southern Illinois University. 


Ropert H. ANDERSON, 


Systematic ENTOMOLOGY, 
nois; WitLpuLirk ResEaRcH, WiLLarD D. Kuimstra, Ph.D., Professor of Zoology and Director of Cooperative Wi 
PARASITOLOGY, Norman D. Levine, Ph.D., Professor of Veterinary 
Parasitology, Veterinary Research and Zoology and Director of the Center for 
Professor of Zoology and of Entomology and Mead of the De- 
and GILBERT Ee 
Statistics, Horace W, Norton, Ph.D., Professor of Statistical Design and Analysis, University” 


| 


L. L. Stoss, Ph.D., Geology ; 
Cuaries E. 


Herperr §, 
Oumstep, Ph.D., 
Evserr HH, 


Tep W. Srorck, Ph.D., Junior Professional Scientist 
Ricuarp J. Baur, M.S., Research Assistant 

Tom Hi, M.S., Research Assistant 

RicHarD KocuHer, B.S., Research Assistant 

Ropert Moran, M.S., Research Assistant } 
C. Russet Rose, Field Assistant . 


Section of Faunistic Surveys and 

Insect Identification 

Puinie W. SmirH, Ph.D., Taxonomist and Mead 
WavuacE E. LaBerGe, Ph.D., Taxonomist 
Mitton W. SanpEerson, Ph.D., Taxonomist ‘ 
Lewis J, STANNARD, JR., Ph.D., Taxonomist 
Larry M. PaGeE, Ph.D., Assistant Taxonomist 
Joun D. Unazicker, Ph.D., Assistant Taxonomist 
DonaLp W. WEBB, M.S., Assistant Taxonomist 
BERNICE P. SWEENEY, Junior Professional Scientist 


Section of Wildlife Research it 


Gurn C. SanpERsoN, Ph.D., Wildlife Specialist and Head 
FRANK C. BELLROSE, B.S., Wildlife Specialist 
JEAN W. GRABER, Ph. D., Wildlife Specialist 
Ricwarp R. GRABER, Ph. D., Wildlife Specialist 
Haroxp C. Hanson, Ph.D., Wildlife Specialist 
Ronatp F. Lapisky, Ph.D., Wildlife Specialist 
Witiiam L. ANDERSON, MA. Associate Wildlife 
Specialist 
W. W. Cocuran, In., B.S., Associate Wildlife Specialist 
Wiuuiam R. Epw. ARDS, M.S. , Associate Wildlife Specialist 
G. Buair JOSELYN, MS. , Associate Wildlife Specialist — 
CHarues M. Nixon, M.S., Associate Wildlife Specialist — 
KENNETH E. Smitu, Ph. D. ., Associate Chemist ¢ 
Ronatp L. WesTeMEIER, M.S., Associate Wildlife 
Specialist 
SrepHEeN P. Havera, M.S., Assistant Wildlife Specialist 
Davin R. VANCE, M.S., Assistant Wildlife Specialist 
Ronaup E. Duzan, Junior Professional Scientist 
HELEN C. Scuvnrz, M.A., Junior Professional Scientist 
ELEANORE WILSON, Junior Professional Scientist ” 
SHARON FRADENBURGH, B.A., Laboratory Technician 
Rogsert D. CrRoMPTON, Field Assistant 
James W. SeeEts, Laboratory Assistant 


Section of Administrative Services 
Rosert 0. Watson, B.S., Administrator and Head ) 
Supporting Services . 

Vernon F, BrutMan, Maintenance Supervisor - 

Wiutma G. Dittman, Property Control and Trust ey 
Accounts 7 

Party L. Duzan, Technical Assistant 

Rosert O. Evis, Assistant for Operations 

Larry D, Gross, Maintenance Supervisor 

Luoyp E. HurrMan, Stockroom Manager 

J. Witu1aM Lusk, Mailing and Distribution Services 

MELVIN E. SCHWARTZ, Financial Records 

James E, SERGENT, Greenhouse Superintendent 


Publications and Public Relations } 

Owen F. Guissenporr, M.S., Technical Editor « 

Ropert M. ZEwaDskKI, MS. be "Associate Technical Editor 
SHirRLEY McCuLELLAN, Assistant Technical Editor 
LAWRENCE S. Fartow, Technical Photographer 

Lioyp LEMERE, Technical Illustrator 4 

* 


Technical Library 


Doris F. Dopps, M.S.L.S., Technical Librarian — 
Doris L, SuBLerre, M.S. LS. .» Assistant Teclaaas 
Librarian 


Roperick R. Irwin, Chicago, Il 


Human Ecology, University of 


WatpBAupr, Ph.D., Professor of Entomology, UI 


cad 


CONTENTS 


oy WE ELISIVERVTETS se eo eld odegi ney oo. 6c Ort Scene chem CnRERene Ceee 145 
PRATAP TESIPUN DIB NEEL ODS eerie eer anya) AN es oy oyay oc Success «ates ove Saracens 146 
MAS Gis) (CRTN INCHES gg do conte. Jo eto clo Seer 147 
(CIID -2b.5 oS aea cabs Sob e gage ad Ada ee ae on re 148 
ATGINEBIS 5.6 o'clp'a g BO oie aid. 6 © ahs Glo 6 ee ORS aan eee 148 
[Risavel arayal TPavosavelgiton) 6% s.qcerb aos. ded cue -Gin NR en eteee ne 149 
WenmninaleAbdominal Segments: 22.2... . 26s. cel aes scs ces eecsceaead 149 
SIBUDIRY se 0’ 9 Beet ele BAe: BOP CEI B CRDIIO Ee ee 149 
(ES opsieo ooh pod tie Mee eo athe Reve See ee 149 
SE OREHOSIS Me reas Mere tn Marana dette he sas cae Ses deh eieeeseceb’ies 149 
Lire History OF SERICOTHRIPS VARIABILIS (BEACH) ...............--0-0- 150 
WAlBRlNG@!S 2054.5 Sé.p cto Sse be BERIDD ODIO CIEE Ce ee ene 150 
[Passing aue ILRI El, 9444.08 Btho So bono 6 BOD ee ene eee eee 151 
SECT ILEINEL: Gelade go had oe Oo. or citea DEN e eee eee 152 
PESTS TOUIY) Meee se PI TA ol ox icv ees\ og etencie eh eqstoher of opes svar oi sbehonecaere; eran sys nip 152 
EI AMER pete ek RTS ric ese, «Teil ctaliayev atare o GremetesclnlGvevshnionavaueverevand “vtetevayedt 153 
EN GHEE. 5. Sdio 6 SSMS 8 OE SNL ERECT Ey re PP en 153 
Effect of Temperature and Photoperiod on Development ............. 153 
BLM OLM EAU a OMMerr sg tapers (eters cieie Sar stnh= s.0'e Gasapaueet shes vers © c+ dierescueuea aera 154 
IMUIBWHEOND “a2 5 cay dig Dee 0 Reuse TUE CRD ER REE pear ee aa 154 
TE CT LOTS eis hee B isso! j.<, = SPRINT  ieet eie S artie ns nyt Raat, ae 155 
EIGONOMMICPCASSESSINCIE 58.5.2 eee eet ecole re Paceteteyavaotuer cone eas 155 
DEE BRIEOHSTIN StS 5a tk ot, 0h tr. RRM Regal Sood Meena kate woe aaah 156 
ipbvlogeny) ofthe: Mhysanopteraa. cots ate -tabevenchatsrayc ee sein so avsichateveigeeie ote 157 
Phylogeny of the Tribes of the Thripidae ........................... 159 
Phylogeny of the Sericothripini 5 ne Conor OA DI EE SE A Eo 163 
MTEL SO ee crc aioe Ee 5h So 6. =o) RRR RT ECAP aaa ora Sa GL a Gre hes, eee eens, matien 166 
BEAUTE TIN CLUE A Bayay tht ee ae eae eBCn are Le its, wae 2 ove; ooh fast Moueieenseisi © 204 
TEESE yo. GSR neN Re Ween EI aR lee. te Ont on ate ea 207 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. 
It is a contribution from the Section of Faunistic Surveys and Insect Identification of 
the Illinois Natural History Survey. 

Thomas C. Vance is employed by the Illinois Department of Conservation as a Site 
Interpretive Specialist at Lincoln Log Cabin State Park, Lerna, Illinois. 


(58199—2M—8-74) 


Frontispiece.—Larva | (lower left) and larva Il (upper right) of Sericothrips pulchellus 
Hood on its host, wafer ash (Ptelea sp.). (Photographs by Lawrence S. Farlow) 


Larvae of the Sericothripini 
(Thysanoptera: Thripidae), with Reference 
to Other Larvae of the Terebrantia, of Illinois 


The morphology and taxonomy of the 
immature stages of the Thysanoptera 
have received minimum attention in 
North America. Significant contribu- 
tions on the larvae of thrips have been 
made in Europe, East Asia, and North 
Africa (Priesner 1926a, 1926b-1928, 
and 1960) and in India (Jagadish & 
Ananthakrishnan 1972), and_ these 
studies constitute the basis of our 
knowledge. In the United States most 
of the descriptions of the immature 
stages are found in accounts of the life 
histories of economically important 
thrips. 


This report deals mainly with the 
second-stage larvae, especially the 
known forms belonging to the tribe 
Sericothripini as represented in Illinois, 
and includes a comparison of the larval 
characteristics of many of the genera 
of the suborder Terebrantia that are 
found in the same region. Larval char- 
acteristics were used to substantiate the 
classification formerly based on adult 
features and to interpret the phylogeny 
of this insect order. A special study 
on the life history of Sericothrips vari- 
abilis (Beach) was included to_provide 
an example of the bionomics of a com- 
mon species. 


References to the literature, with few 
exceptions, terminated in 1971 when 
this report was submitted as.a Master 
of Science thesis to the Department of 
Entomology, University of Illinois, Ur- 
bana. 


ACKNOWLEDGMENTS 


Support for this work was provided 
by the Illinois Agricultural Experiment 
Station, Project S-74, Biology and Con- 


Thomas C. Vance 


trol of Arthropods on Soybeans, and 
by the Illinois Natural History Survey. 


I thank all those who have helped 
in the preparation of this report. I am 
particularly grateful to Dr. Lewis J. 
Stannard, Jr., for many suggestions and 
continued help throughout the course 
of this study and to Dr. Bruce S. 
Heming for additional advice and in- 
formation, especially on the tentorium 
and other morphological features. Ap- 
preciation is extended also to Dr. Wil- 
liam H. Luckmann, Illinois Natural 
History Survey, for arranging financial 
support; to Lloyd L. LeMere, Survey 
Technical Ilustrator, for drawing Fig. 
1-5; to Wilmer D. Zehr, former Tech- 
nical Photographer of the Natural His- 
tory Survey, and Lawrence S. Farlow, 
present Survey Photographer, for photo- 
graphic reproductions of the figures; 
and to many other staff members at the 
Survey for their assistance and kindness. 
Further, I wish to thank my former 
associate, Dr. Thomas H. Wilson, for 
help and consultation on many prob- 
lems, and my wife, Susan, for construc- 
tive criticism. 

Most of the material studied was 
from the collection of the Illinois Natu- 
ral History Survey. Additional speci- 
mens were lent to me through the gen- 
erous cooperation of Miss Kellie O'Neill, 
U.S. Department of Agriculture, and 
Dr. Tokuwo Kono, California Depart- 
ment of Agriculture. 

The manuscript was edited for pub- 
lication by Robert M. Zewadski, As- 
sociate Technical Editor, Illinois Natu- 
ral History Survey, and reviewed by 
Dr. Bruce $. Heming, Associate Profes- 
sor, University of Alberta, and Dr. 
Lewis J. Stannard, Jr., Illinois Natural 


145 


146 


History Survey Taxonomist and Pro- 
fessor of Entomology, University of Ili- 
nois. The typing and proofreading of 
the manuscript were done by Mrs. 
Bernice Sweeney and Mrs. Grace Fin- 
ger, Illinois Natural History Survey. 


MATERIALS AND METHODS 


During this study about 500 im- 
mature thrips were examined. In addi- 
tion, diagnostic features were analyzed 
from descriptions of immatures in the 
literature, the reference being cited in 
each case. Repositories and institutions 
are identified in the Material-Examined 
sections by these abbreviations: 
INHS =—lIllinois Natural History 
Survey collection 

USNM — United States National Mu- 
seum (National Museum 
of Natural History, Smith- 
sonian Institution) 


Three methods were used in collect- 
ing immature thrips. Large plants were 
sampled with a black sweep net (to 
make the light-colored immatures more 
visible), the thrips being recovered from 
the net with the aid of a hand lens 


and a small camel hair brush. Branches. 


were shaken over a piece of cardboard 
or other material from which the thrips 
were recovered. Small host plants were 
sampled by examining individual leaves, 
and the thrips were removed directly 
from the leaf surfaces. 


The preserving solution used was 
AGA (eight parts 95-percent ethanol, 
five parts distilled water, one part glyc- 
erine, and one part acetic acid), which 
kept the body soft and facilitated 
spreading of the appendages. For stor- 
age beyond 4 weeks, thrips were trans- 
ferred to 70-percent ethanol. 


Both Canada balsam and Hoyer’s 
medium were used in making whole 
mounts. Canada balsam is a permanent 
mounting medium (Hartwig 1952; Pries- 
ner 1960; Stannard 1968), which pre- 
serves the color and features of thrips 
well, but it is difficult to use and much 


Inuivois NaruraL History SURVEY BULLETIN 


time is required to make good prepara- 
tions. Further, because of dehydration 
and accompanying brittleness, speci- 
mens can be damaged during mount- 


ing in Canada balsam, and small setae, — 


microtrichia, cuticular sculpturing, and 


areas of light brown coloration often 


are obscured. 


Hoyer’s is easier to use and renders — 
visible many diagnostic features not 


usually seen on specimens mounted 


in balsam. Unfortunately, Hoyer’s, a 


water-base medium, usually crystallizes 
within a few years. Specimens for the 


permanent collection, therefore, were 


mounted in Canada balsam, but some 
of each series were mounted in Hoyer's 
medium for temporary study. 


Balsam mounts were prepared in a 
manner similar to that described by 
Heming (1969). Larvae and adults were 
transferred from AGA to 70-percent 
ethanol and were then passed succes- 
sively through 95-percent ethanol, ab- 
solute ethanol, and absolute ethanol 
and clove oil, remaining in each solu- 
tion for about one-half hour. Specimens 
were then placed in a small Syracuse 
watch glass containing pure clove oil; 
when each sank to the bottom, it was 


transferred to a slide. Clearing in 10- 


percent KOH or NaOH was usually un- 
necessary for immatures except to dis- 
solve the excessive amounts of fat body 
found in some larvae. 


In mounting, each thrips was placed 
ventral side up in a small drop of 
dilute balsam on a cover slip held in 
place on a small cardboard stage. The 
appendages were spread, and two chips 
of cover glass were added to the 
balsam. These chips prevent crushing 
of the specimen by the cover slip as 
the balsam dries. A small drop of 


balsam was placed in the center of a_ 


microscope slide, and the slide was in- 
verted and placed gently upon the 
cover slip. When the slide was lifted 
and turned right side up, the cover 
slip and specimen adhered to it. Slight 
pressure applied to the cover slip with 


Vol. 31, Art. 5 


August, 1974 


an insect pin spread the appendages 
farther. ; 

Whole mounts in Hoyer’s medium 
were prepared in the same way, but 
the dehydration schedule was omitted. 
Most Hoyer preparations in the Survey 
collection deteriorated after a few 
years, even when ringed with Zut Slide 
Ringing Compound (Bennett’s Paint 
Products, Salt Lake City) or clear fin- 
gernail polish. However, some prepara- 
tions ringed with fingernail polish have 
remained in good condition for more 
than 20 years, indicating that efficient 
ringing compounds might prove suc- 
cessful in preserving Hoyer mounts. 

Bright-light microscopes were used 
throughout this study except when 
minute structures, such as microtrichia, 
were being observed, for which work 
phase-contrast microscopes were em- 
ployed. 


ANALYSIS OF CHARACTERS 


According to Priesner (1960) “the 
shape of the antennal segments, the 
sculpture of the body cuticle, the 
chaetotaxy, and last but not least, the 
colour, are important” in taxonomic 
study of larval thrips. These characters 
and certain others were the principal 
ones used in this investigation. Many 
characters varied with the stage of 
larval development, particularly color, 
many body dimensions, and cuticular 


-sculpturing, which vary with growth 


and instar. Color also varies with the 
type of food consumed by the larvae. 
Different characters have been used in 
this study according to the taxonomic 
level concerned. In classifying thrips 
larvae at the family level, the form and 
shape of the antennal segments and the 
presence or absence of modified spines 
on the ninth abdominal tergite are im- 
portant in making distinctions. At the 
subfamily level, the form of certain 
antennal segments is important. Many 
characters at the tribal level were found 
to intergrade, but certain features could 
generally be assigned to each tribal 


VANCE: LARVAE OF THE SERICOTHRIPINI 


147 


group. Members of subtribal groups 
tended to exhibit a greater degree of 
similarity and could be assigned to the 
proper group with less difficulty. 


The greatest stabilization of charac- 
ters occurs at the genus level. Most 
genera are sharply delimited, and even 
closely related genera usually exhibit 
diagnostic differences. One exception 
occurred in the tribe Thripini in which 
the larvae of the Frankliniella-Thrips- 
Taeniothrips complex are quite similar. 
Important generic characters include 
cuticular sculpturing; microtrichia; setal 
type, length, and placement; coloration; 
and proportions and features of the 
antennal segments. 


Little distinction was found at the 
species level, closely related species 
often being nearly alike in form. Species 
differences that were found include the 
length and proportions of the body 
setae, brown sclerotized areas, setal 
basal rings, and cuticular and hypo- 
dermal coloration in mature larvae. 
Ward (1968) found that slight consist- 
ent differences are present in larvae 
of several closely related species of 
Thrips and that, despite their subtlety, 
these characteristics can be used to 
separate these species with confidence. 


Most of the characters mentioned 
above apply to second-instar larvae; 
in first-instar larvae few diagnostic 
characters occur at the generic level 
and none were detected at the spe- 
cific level. At the family and subfamily 
levels first-instar larvae may be recog- 
nized by the same antennal characters 
distinguishing second-instar larvae. At 
the tribal and subtribal levels the pat- 
tern of microtrichia on the antennae 
and general body and antennal features 
are useful in making distinctions. 


The prepupal and pupal instars show 
little interspecific variation. According 
to Priesner (1960), the only distinguish- 
ing characters are the presence or ab- 
sence of cuticular spines near the apex 
of the abdomen and the shape of the 
antennae. The taxonomic value of these 


148 


features above the species level may 
be questionable, since Priesner (1960) 
reported one species of Taeniothrips 
with spines and another species of the 
same genus without them. 


COLOR 

Four types of coloration occur in 
thrips larvae: (1) that of the internal 
organs and body contents, (2) that of 
the cuticle, (3) that of underlying hypo- 
dermal pigmentation, (4) and areas of 
brown sclerotization on the cuticle sur- 
face. Because color varies with the de- 
gree of larval development, it is best 
to deal only with fully mature larvae. 

The colors of internal organs and 
body contents depend upon the food 
ingested. Phytophagous larvae often 
appear green due to the ingestion of 
chlorophyl, and predacious larvae may 
assume the color of the prey ingested. 
Such colors are usually leached out dur- 
ing the mounting process and are prac- 
tically useless for taxonomic purposes. 

Cuticle color among specimens of 
the same species varies from white to 
yellow to orange. These pigments can 
be affected by the mounting media 
used, and are leached out with pro- 
longed storage in alcohol. 

Underlying hypodermal pigmenta- 
tion is usually not affected by mounting 
media but does vary greatly even in 
the members of a series of specimens. 
Some species never show hypodermal 
pigmentation, while in others it is usu- 
ally present in some members of a 
series of specimens. Hypodermal pig- 
mentation is susceptible to leaching 
with prolonged storage in alcohol al- 
though at a slower rate than is cuticular 
coloration. 


Brown sclerotized areas, such as cer- 
tain antennal segments, areas of the 
head and thorax, and areas of the 
terminal abdominal segments, are the 
most dependable color features. Dis- 
tinctive brown sclerotized areas are 
particularly valuable in the identifica- 
tion of many species of the Helio- 


Ittinois NAturAL History SuRvEY BULLETIN 


Vol. 31, Art. 5 


thripinae, Anaphothripini, and Chiro- — 
thripini. This brown color does not 
vary much within a species, is not 
leached with prolonged storage in al-— 
cohol, and is not affected by mounting — 
media although these light brown areas 
may be difficult to see in balsam. 


ANTENNAE a 


Antennal features are the most re- 
liable characters in the taxonomy of 
larval thrips. Lengths of segments and — 
the number of annulations present are — 
important at the family and subfamily — 
levels, whereas the shapes of the seg- — 
ments and the nature of their annula- 
tions and microtrichia can be diagnostic — 
of genera and higher groups. The 
microtrichia of antennal segments III — 
and IV and the shapes of the terminal — 
segments are often diagnostic in first-_ 
stage larvae of certain groups. Larval 
members of the Sericothripini, for ex-_ 
ample, tend to have narrowed, tapering, — 
seventh antennal segments and dense, 
random microtrichia on segment IV. 
However, members of some other tribes 
have broader seventh segments, and — 
few have microtrichia on com IV 
except on the annulations. 


Antennal sense cones are of diag-— 
nostic value at the generic and higher ~ 
levels. The length of sense cones in 
adult Thysanoptera often varies, but 
in the larvae it seems fairly stable. In” 
general, the primitive families (Aeolo- — 
thripidae, Merothripidae, and Hetero- — 
thripidae) and the tribes Chirothripini — 
and Thripini tend to have shorter sense 
cones, and the Anaphothripini, Serico- — 
thripini, and MDendrothripini have © 
longer ones. The sense cones on seg- — 
ments IV, V, and VI are the best de- 7 
veloped and therefore are used for — 
taxonomic analysis. g 

' 


The entire antennae of some genera 
are diagnostic (such as those of Chiro-— 
thrips, which has greatly reduced an-~ 
tennae); features of the entire anten-— 
nae, however, often show little dif- 
ferentiation at the generic level. fe 


zt 


August, 1974 


HEAD AND PRONOTUM 


The shape and size of the head and 
pronotum are distinctive and diagnostic 
of certain genera of thrips larvae. These 
features include the ratio of length to 
width, shape, size of eye facets, degree 
of bulging of the eyes, and degree of 
constriction at cheek margins. Small, 
nonbulging eye facets occur in the 
Chirothripini, and construction of the 
cheeks seems to be characteristic of 
the Heliothripinae and some Anapho- 
thripini. 

Problems associated with the head 
and pronotum include distortion due 
to pressure from the cover slip and dif- 
ferences in their degree of development 
within the larval stage. 


TERMINAL 
ABDOMINAL SEGMENTS 


The shape of the terminal abdominal 
segments differs between the suborders 
Terebrantia and Tubulifera. In the 
Thripini and in Anaphothrips a pos- 
terior comb is present on abdominal 
segment IX. According to Priesner 
(1960), each species has a characteristic 
form of this comb. 


SETAE 


The type and length of body setae 
are important features in larval dif- 
ferentiation. Setae vary in length and 
type above the generic level; however, 
they are useful in the diagnoses of 
genera. Setal types, as listed by Pries- 
ner (1960:66-67), are: pointed, lance- 
olate, blunt or rounded, knobbed, fun- 
nel-shaped, forked or fringed, and 
spoon-shaped or fanned. Their lengths 
may vary from less than 5 »m up to 
70 »m, and they may be slender or 
stout. Each genus has characteristic 
types and lengths of setae. 


Setae differ in their widths and 
lengths between species, and certain 
setae differ in their proportionate 
lengths. The degree of development of 
the brown rings at the bases of the 


VANCE: LARVAE OF THE SERICOTHRIPINI 


149 


setae can be important diagnostic fea- 
tures. Some variation in the setae oc- 
curs between individuals; the lengths, 
however, do not change with the degree 
of development. 


CUTICLE 


The presence and nature of cuticular 
pustules and cuticular microtrichia pro- 
vide good diagnostic characters at the 
generic and subtribal levels. Micro- 
trichia are long to short, depending on 
the species. Short microtrichia are al- 
most invisible when viewed through a 
light microscope and appear as a stip- 
pling effect. They are sparsely to 
densely scattered over the integument. 
Pustules are minute to large, depending 
on the species, and usually each pustule 
bears one microtrichium although the 
large pustules of the Anaphothripini 
and Heliothripinae lack microtrichia. 

Cuticular features which are stable 
at the generic level present some prob- 
lems. Small pustules and microtrichia 
are often difficult to see in balsam 
mounts and can be distorted by the 
mounting process. Also, cuticular sculp- 
turing varies with the degree of larval 
development and abdominal distension. 


METAMORPHOSIS 


In the Terebrantia there are usu- 
ally four immature stages, the first- 
and second-instar larvae, the prepupa 
(propupa), and the pupa. In the Tubu- 
lifera, by contrast, an additional pupal 
instar occurs, resulting in a total of five 
stages. Larval stages lack wings or 
wing pads and have free antennae, and 
active movements and feeding take 
place. The prepupal and pupal stages 
are quiescent and do not feed. Their 
antennae lack segmentation and are di- 
rectly forward in prepupae and are bent 
back dorsally (Terebrantia) or laterally 
(Tubulifera) along the head in pupae. 
Wing pads are usually present in 
prepupae and pupae of the Terebrantia 
but only in the pupal stages of the 
Tubulifera. Each stage is terminated 


150 


by a molt, with the exuviae usually left 
on the leaf surface. 


Thrips are usually recognized as 
exopterygote insects and are placed 
with the hemipteroid orders even 
though their postembryonic develop- 
ment more closely resembles. the holo- 
metabolous transformations found in 
the Endopterygota. This intermediate 
type of development in the Thysanop- 
tera has caused considerable contro- 
versy, some authors calling the im- 
matures nymphs and others calling 
them larvae and pupae. Takahashi 
(1921) even proposed the term “Reme- 
tabola” for thysanopteran metamorpho- 
sis. 


Recent histological studies on the 
postembryonic development of the 
Thysanoptera have provided insights 
into the problem. Davies (1961) found 
that the development and adult mor- 
phology of the female reproductive or- 
gans of Limothrips cerealium showed 
similarities to the exopterygote insects 
but that their delayed development 
recalled endopterygote morphogenesis. 
This conclusion is supported by Hem- 
ing (1970) in a similar study on Frank- 
liniella fusca (Hinds) and Haplothrips 
verbasci (Osborn ). Davies (1969) stud- 
ied the metamorphosis of the skeletal 
musculature of L. cerealium and found 
many details of myogenesis in the 
pupae of thrips to be similar to those 
in the pupae of endopterygote insects. 
He stated that “thysanopteran ontogeny 
shows histological changes at least as 
great as those in the holometabolous 
metamorphosis of many Endopterygota 
and these quiescent instars are per- 
fectly entitled to rank as pupal stages.” 
Davies further hypothesized that the 
holometabolous type of metamorphosis 
in the Thysanoptera developed inde- 
pendently of that of the Endopterygota 
and speculated about the selective 
value of two or three pupal stages in 
the Thysanoptera when only one is 
usually necessary for similar trans- 
formations in the Endopterygota. 


Tiuinois NATURAL History SurvEY BULLETIN 


Vol. 31, Art. 5 


LIFE HISTORY OF 
SERICOTHRIPS VARIABILIS 


(BEACH) 


S. variabilis was the first species of 


. 


Sericothrips described in North America 


(Beach 1896) and is one of the most 


common in the eastern states. It occurs — 
abundantly on soybeans and other 
legumes, but its life history and the — 


economic damage it causes are largely 


unknown. 


Life-history studies have been made — 


on several economically important 


thrips, the most complete being those 
of Horton (1918) on Scirtothrips citri— 


(Moulton), Bailey (1933) on Caliothrips 


fasciatus (Pergande), and Ghabn (1948) — 
on Thrips tabaci (Lindeman), Other — 


accounts by Bourne (1926), Davidson — 


& Bald (1930), Foster & Jones (1915), 
McKenzie (1935), Rivnay (1935), Rus- — 


sell (1912), Sakimura (1932), Schopp 
(1936), Watts (1934), and White (1916) 


are more brief. Bailey (1938) sum- 


marized and compared the life histories 


of several thrips of economic impor- — 
tance in California. Rearing methods ~ 
are described by Bailey (1932 and ~ 


1933), Rivnay (1935), and Callan (1947). 


The following data on S. variabilis 
are intended to provide information on — 


the 
stages, the effects of temperature and 


photoperiod, the site of pupation, — 
mating, and predators, and an assess- — 


ment of the economic importance of 
the species. 


METHODS 


development of the immature © 


Two types of rearing containers were — 
used. The first was a covered plastic — 


petri dish (85 mm in diameter and 10 


mm deep) set vertically in a wooden — 


a 


& 


rack. A soybean leaf was trimmed to — 


fit into the dish with its stem extending 


through a hole in one side of the dish — 


and into a vial of water below. 


The second rearing container was a — 


100- x 15-mm covered glass petri dish 


August, 1974 


containing two soybean leaves and a 
piece of filter paper which was moist- 
ened daily. Larvae and adults were 
collected from soybeans on the South 
Farm of the University of Illinois, 
Urbana. 


Rearing was done at controlled tem- 
peratures of 21.0°, 26.5°, and 32.0° C 
under constant light and at 22.0° C 
under an 8-hour-per-day light photo- 
period. Two cultures were confined 
at each temperature. One culture was 
started with eggs already present in 
the leaves. A second culture was 
started with 10 adults. The number of 
larvae at each stage of growth was 
recorded twice daily between 0800 and 
0900 hours and between 1600 and 1700 
hours. 


Data were tabulated and analyzed 
by recording the duration of each im- 
mature stage and computing each 
mean. Further analysis included the 
calculation of the standard error of the 
mean and ¢ tests at a significance level 
of 0.01. 


The site of pupal development was 
determined by examining for pupating 
thrips field samples of soil collected 
from beneath soybean plants at depths 
of 1 inch (25.4 mm) and at 4-5 inches 
(101.6-127.0 mm). Soil was placed in 
the lower end of a glass petri dish 
held at a 45° angle. Soybean leaves 
were set upright in the dish with the 
stems resting on the soil. Larvae pres- 
ent on the leaves could therefore drop 
or crawl to the soil when ready to 
pupate. 

Sticky traps were set in the field to 
determine how the second-stage larvae 
reach the ground. Tanglefoot (Tangle- 
foot Company, Grand Rapids, Michi- 
gan) was placed in 1-inch (25.4-mm) 
bands 6 inches (152.4 mm) from ground 
level around and directly on the stems 
of 12 soybean plants to trap any larvae 
crawling down the stems. Two 12-x 
18-inch (304.8- x 457.2-mm) cardboard 
sheets covered with Tanglefoot were 
placed on the ground beneath the 


VANCE: LARVAE OF THE SERICOTHRIPINI 


151 


plants at least 6 inches (152.4 mm) from 
the stems to catch any larvae dropping 
from the leaves. 


FIRST-INSTAR LARVA 


The mean duration of the first instar 
of S. variabilis larvae reared at 22.0° 
C with an 8-hour light photoperiod 
per day was 73.49 + 7.32 hours. The 
mean body length of the cultured im- 
matures of this instar varied from 
560 + 80 pm for the early larva I to 
720 + 70 um for the late larva I. The 
early larva I has a narrow, tapering 
abdomen and a_ disproportionately 
large head and legs (Fig. 7a). As feed- 
ing takes place, the body becomes dis- 
tended due to increases in the sizes 
of the internal organs, particularly the 
fat body (Fig. 7b). Cuticular color 
changes from white in the early larva 
I to yellow in the late larva I, and in- 
gested chlorophyl often gives the body 
a green color. 

The setae are short and narrowly 
fanned, setal pair P7 is lacking, and 
abdominal segment IX has three or four 
pairs of setae. (The setal and segmental 
numbering system used in this report 
is shown in Fig. 6.) Priesner (1958) 
speculated, but Ghabn (1948) had 
proved, that the male larva I has three 
pairs of setae on segment IX (two 
dorsally and one laterally), whereas 
the female has four pairs of setae on 
this segment (two dorsally, one later- 
ally, and one ventrally). The sexes can 
be determined by these setal arrange- 
ments. Antennal segment IV is covered 
with random microtrichia, and segment 
VII is tapered apically. 


Soon after hatching, the larva begins 
feeding, never moving far from the 
hatching site and often hiding in the 
angles of the larger veins on the lower 
leaf surface. The larvae are active and 
move about quickly when disturbed. 
In late larvae I the old cuticle becomes 
light gray. It splits dorsoventrally, the 
head and thorax are pushed out, and 
the antennae and legs are pulled free. 


152 


The exuviae is pushed partly down the 
abdomen by the hind feet, and the 
remainder of the abdomen is pulled 
free by forward pressure exerted on the 
leaf surface by the feet. About 4 
minutes are required for this process. 


SECOND-INSTAR LARVA 


The mean duration of the second 
instar of S. variabilis larvae reared at 
22.0° C under an 8-hour light photo- 
period per day was 91.30 + 10.44 
hours. The mean body lengths of the 
cultured immatures of this instar varied 
from 910 + 60 »m for the early larva 
II to 1,030 + 50 pm for the late larva 
II. A newly molted larva has a narrow 
abdomen and thorax and a dispropor- 
tionately large head and legs (Fig. 7c). 
As the larva feeds, the abdomen, par- 
ticularly, and the thorax become dis- 
tended (Fig. 7d). The cuticular color 
changes from white in the newly 
molted larva to orange, often with red 
hypodermal pigmentation, in the late 
larva II (although the red pigmentation 
was not observed in laboratory-reared 
larvae). Green body coloration due to 
ingested chlorophyl was predominant 
in many larvae. 


The setae are long and widely fanned, 
appearing proportionately longer in the 
early larva II because the lengths of the 
setae remain unchanged throughout the 
larval stage. Setal pair P7 is present, 
and abdominal segment IX has five or 
six pairs of setae. Sex was determined 
by following Priesner (1958) on the 
number of setae on segment IX. Those 
larvae with five pairs of setae (two 
dorsally, two laterally, and one ven- 
trally) were presumed to be females, 
and those with six pairs of setae (two 
dorsally, two laterally, and two ven- 
trally) were presumed to be males. 
Priesner ignored one pair of lateral 
setae (A3 in this study) because they 
were greatly reduced, and gave the 
setal counts as four and five pairs. 
However, A3 is not reduced in larvae 
of certain genera (e.g., Aeolothrips, 


Intivois NATuRAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


} 
Merothrips, and Heterothrips), and for : 
the sake of uniformity, this pair of — 
setae was included in all setal counts — 
here. The color of the setae are white — 
immediately following the molt (some- — 
times making newly molted second- — 
stage larvae easily confused with mid- 
first-stage larvae) but soon become — 
sclerotized and turn brown. Antennal 5 
segment IV has microtrichia only on the © 
annulations. 


$ 
Second-instar larvae feed on the leaf 
surface and occasionally hide in crey- 4 
ices. In nature they are almost always — 
found on the undersides of leaves, but — 
they also occur on the upper sides in — 
laboratory cultures. Near the end of — 
the larval stage, the larvae drop to — 
the ground and enter the soil for pupa- — 
tion. Hi 
} 

: 

d 


PREPUPA 


The mean duration of the pre-— 
pupal stage of S. variabilis reared at 
22.0° C under an 8-hour light photo- — 
period per day was 22.00 + 2.38 hours. — 
The mean body length was 1,180 + © 
80 »m. Changes in size are -imper- 
ceptible during the prepupal stage. The — 
color is predominantly orange; wing — 
pads are present, reaching posteriorly — 
to the second abdominal segment; the — 
antennae are indistinctly segmented, 
protruding anteriorly from the head; 
and the setae are simple and short. M 
Abdominal segment IX lacks the cuticu- — 
lar spines found in the prepupae of 
some genera (Fig. 7e). ; 


Female prepupae possess two pairs of — 
short lobes arising ventrally on abdomi- — 
nal segments VIII and IX; these are — 
the buds of the ovipositor valves. Male — 
prepupae lack these structures (Priesner 
1960). 5 


The prepupal period is normally 
passed in the soil, but in laboratory — 
cultures where soil was unavailable, . 
prepupation readily took place on the — 
leaf surface. Under laboratory rearing — 
conditions, the prepupae were quies-_ 


wy 
4 


| 


August, 1974 


cent and nonfeeding and were usually 
hidden in crevices between the large 
leaf veins; activity was observed only 
when the prepupae were disturbed or 
threatened. 


PUPA 


The mean duration of the pupa stage 
of S. variabilis reared at 22.0° C under 
an 8-hour light photoperiod per day 
was 74.00 + 2.83 hours. The mean 
body length was 1,040 + 60 »m. No 
change in size was noted during pupal 
development. The color is predomi- 
nantly orange during this stage. The 
wing pads reach the sixth abdominal 
segment, and the antennae are recurved 
along the dorsum of the head. The 
setae are simple and pointed and longer 
than in the prepupa. Abdominal seg- 
ment IX lacks the cuticular spines 
found in the pupae of some genera 
(Fig. 7f). 

The ventral lobes on segments VIII 
and IX in female pupae are longer and 
better developed than those found in 
female prepupae. Male pupae have a 
bluntly triangular production ventrally 
at the hind margin of segment IX. 


Pupal development took place in the 
upper inch (25.4 mm) of soil beneath 
soybean plants or in the soil provided 
in laboratory cultures. In cultures 
where soil was not available, pupation 
readily took place on the leaf surface, 
the quiescent, nonfeeding pupae being 
hidden between the larger leaf veins. 


ADULT 


Adults of S. variabilis may be dis- 
tinguished from those of other species 
of the genus in Illinois by the following 
combination of characteristics (Stan- 
nard 1968): each fore wing with two 
sharply defined crossbands; the pronotal 
blotch completely dark in contrast to 
the rest of the pronotum and deeply 
incised medially and posteriorly by 
yellow; anterior pronotal striations 
closely spaced; several abdominal seg- 
ments dark brown. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


153 


Adults are quite active and, when 
disturbed, dart about or jump rapidly. 
Adults seldom survive long in a cul- 
ture dish when transferred from field 
samples but remain alive for up to 4 
or 5 days when reared in the laboratory. 


EFFECT OF TEMPERATURE AND 
PHOTOPERIOD ON DEVELOPMENT 


Both temperature and photoperiod 
affected the durations of the immature 
stages of S. variabilis (Table 1 and 
Fig. 1). Under constant light the dura- 
tions of the stages were about 27 per- 
cent longer than that required at about 
the same temperature under an §-hour- 
per-day light photoperiod. Temperature 
and durations of the stages were in- 
versely correlated, the most rapid de- 


oe 
Ww 
a 
= 
< 
a 
Ww 
a 
= 
wi 
= 
4 Srl 2416 + 20) 524) 1528 
DAYS 
o 
w 
a 
=) 
— 
<x 
a 
Ww 
a 
= 
ui 
-— 
4 812° 16) 20) 24) 28 
DAYS 
Fig. 1.—Growth curves of immature stages 


of Sericothrips variabilis reared in the labora- 
tory. A, mean duration of the larva | and 
larva I] stages, combined, at various constant 
temperatures. B, mean duration of the pre- 
pupal and pupal stages, combined, at various 
constant temperatures, 


154 Inuinois NATURAL History SURVEY BULLETIN Vol. 31, Art. 5 
¥ 
Table 1.—Duration of immature stages of Sericothrips variabilis at different temperatures — 


and photoperiods. 


The numbers of insects observed are in parentheses. 


. 


y 
Temperature Mean Hours of Duration { 
(in Celsius) a 
and Photoperiod Larva I Larva II Prepupa Pupa j 
22.0°, 8-hour 73.49 = 7.328 91.30 + 10.44 22.00 + 2.38 74.00 + 2.83 ; 
photoperiod (23) (24) (2) (2) i 
21.0°, constant 97.33 + 4.68 107.33 + 6.99 35.00 + 7.57 3 
light (6) (8) (4) 
26.5°, constant 65.33 + 6.11 87.34 + 14.09 29.33 + 6.11 
light (5) (14) (3) . 
32.0°, constant 58.67 + 2.83 68.00 + 6.85 16.00 + 4.00 56.00 + 6.20 
light (7) (11) (7) (6) 4 
> 


4 Standard error. 


Larva I: 
Prepupa: 


22.0° and 26.5°, 26.4° and 32.0°. 


velopment taking place at 32.0° C. At 
26.5° C the mean duration of immature 
stages was increased by about 41 per- 
cent, and at 21.0° C the mean duration 
was increased by about 81 percent over 
the time required at 32.0° C. The least 
mortality occurred at 26.5° C, indicat- 
ing that this might be the optimum of 
the three temperatures for the develop- 
ment of the immature stages of this 
insect. 


S. variabilis requires more time for 
development of the immature stages 
than does Thrips tabaci Lindeman and 
Taeniothrips simplex Morison; the same 
time as Scirtothrips citri (Moulton), 
Caliothrips fasciatus (Pergande), and 
Frankliniella tritici (Fitch); and less 
time than Taeniothrips inconsequens 
(Uzel), Heliothrips haemorrhoidalis 
(Bouché), Liothrips vaneecki Priesner, 
and Hercinothrips femoralis (Reuter ) 
(Bailey 1938). 


SITE OF PUPATION 


Studies on the biology of many pest 
thrips indicate that late second-stage 
larvae drop to the ground and pupate 
in the soil. S. variabilis also pupates 
in the soil, as shown by the results of 
the field tests conducted during this 
study. Pupae were found 1 inch (25.4 
mm) below the surface in soil samples 
taken beneath soybean plants. Each 


Standard ¢ tests computed at a probability level of 0.01 showed all means 
to be significantly different except for phese pairs: 


22.0° and 21.0°, 22.0° and 26.55, 22.0° and 32.0°, 21.0° and 26.5°, f 


if 
‘4 
pupa was located in a small chamber in~ 
the middle of a dirt particle one-half 
inch (12.7 mm) in diameter. In the 
experiment designed to discover how 
the larvae reach the ground, no imal 
matures were caught in sticky traps” 
placed around the stems of the plants. — 
In contrast, on the sticky sheets beneath 
the plants 10 late second-stage larvae _ 
and three adults were found on one and 
5 late second-stage larvae and two 
adults on another, indicating that the 
larvae drop to the soil from the leaves 
before pupation begins. 


Information concerning the site andl 
conditions of pupation of some thrips” 
is given by Bailey (1933), and Parrot 
(1911) gives information on the use 
of sticky traps in locating pupation sites” 
of certain of the Terebrantia. ‘ 


MATING 

The complete mating process was ob- 
served in two adults that had emerged 
in a laboratory culture. Seemingly the » 
male first became aware of the female 
when he approached within about one 
half inch (12.7 mm) of her. He im- 
mediately ran, caught her, and mounted | 
her dorsally. The female began twist 
ing the abdomen about 2 seconds after 
the male had mounted. Three attempts” 


were then made to make genital con- 
tact, the third being successful. The 


August, 1974 


time lapse to this point from the initia! 
mounting was 22 seconds. Two seconds 
after making genital contact, the male 
dismounted while maintaining genital 
contact, and both male and female 
remained motionless for 51 seconds fac- 
ing in opposite directions. Contact was 
then broken, and each went in a sep- 
arate direction. 


PREDATORS 


Three predators were found in as- 
sociation with S. variabilis in the lab- 
oratory cultures: Aeolothrips fasciatus 
(Linneaus) (Thysanoptera: Aeolo- 
thripidae), Orius insidiosus (Say) 
(Heteroptera: Anthocoridae), and 
mites of the family Phytoseiidae 
(Acarina ). 


Several A. fasciatus immatures ap- 
peared in the cultures and developed 
along with S. variabilis. The Aeolo- 
thrips larvae were observed feeding on 
Sericothrips larvae on three occasions. 
One Aeolothrips reached maturity in 
the culture dish, as did others reported 
on by Robinson, Stannard, & Armbrust 
(1972). 


Phytoseiid mites were observed carry- 
ing dead Sericothrips larvae on two oc- 
casions but were not observed actually 
feeding. According to Chant (1958) 
and Chant & Fleschner (1960), phyto- 
seiid mites can be important predators 
of certain phytophagous mites, but little 
is known of their predation on thrips 
or other insects. In laboratory cultures 
these mites survived well and could be 
reared easily with thrips for study on 
the interaction between the two. 

Nymphs of O. insidiosus were ob- 
served in association with S. variabilis 
on many samples brought from the field 
and were found several times in the 
laboratory cultures. Although no preda- 
tion was observed, it is probable that 
these anthocorids were feeding on 
thrips larvae. Borror & DeLong (1964) 
reported O. insidiosus as predatory on 
various species of thrips and other in- 


VANCE: LARVAE OF THE SERICOTHRIPINI 


155 


sects, and Bailey (1933) showed that 
another species, O. tristicolor White, 
is a predator of the bean thrips, Calio- 
thrips fasciatus. The adults of tristi- 
color were observed to consume about 
one larva an hour, the nymphs appear- 
ing even more voracious. Both nymphs 
and adults preferred young larvae. 
O. indicus (Reuter) feeds extensively 
on Taeniothrips nigricornis (Schmutz ) 
(= T. distalis Karny) in India (Raja- 
sekhara & Chatterji 1970). 


Other predators reported by Bailey 
(1933) were larvae of Chrysopa cali- 
fornica (Coquillett), Hippodamia con- 
vergens (Guerin), Aeolothrips kuwanai 
(Moulton), and A. fasciatus. 


ECONOMIC ASSESSMENT 


Although S. variabilis is generally 
considered to be of minor economic 
importance, Bailey (1940) rated it as 
ninth in economic importance among 
thrips species of the conterminous 
United States. 


In laboratory cultures immature 
stages of S. variabilis apparently caused 
little damage to soybean leaves, even 
with a population of 8—10 thrips per 
leaf, despite the small amount of yel- 
lowing which was evident at times. 


During the latter part of the summer, 
many upper leaves on soybean plants in 
the field showed yellowing, browning, 
and other evidence of insect-feeding 
damage. This damage, however, can- 
not be directly attributed to thrips be- 
cause a variety of other insects also 
feed on soybeans. Furthermore, the 
population levels of S. variabilis in the 
field were estimated at an average of 
one or fewer thrips per leaflet at each 
observation. At this density level little 
economic damage results. However, 
thrips damage at levels of 30-60 insects 
per plant (number per leaflet not 
stated) was reported in Maryland in 
July 1971 in the Cooperative Economic 
Insect Report (U.S. Department of 
Agriculture 1971). So far as is known, 


156 


S. variabilis does not transmit plant 
viruses. 


Other Sericothripini of economic im- 
portance include the citrus thrips, 
Scirtothrips citri, ranked seventh among 
economic thrips species by Bailey 
(1940); the grape thrips, .Drepano- 
thrips reuteri Priesner, given a rating 
of 11 and considered of minor impor- 
tance; the long-winged thrips, Scirto- 
thrips longipennis (Bagnall), ranked 
number 20 and considered as rarely of 
importance; and Echinothrips ameri- 
canus Morgan, ranked 31 and also con- 
sidered rarely of economic importance. 


PHYLOGENY 


Interpretations of the phylogeny of 
the Sericothripini and the relationships 
of that tribe to some of the other groups 
in the Thysanoptera were made on the 
basis of larval characteristics, as pre- 
sented here. 


Larval characters used in assessing 
the relationships of the major groups of 
Thysanoptera were: (1) the degree 
of elongation of antennal segments III 
and IV, (2) the length of antennal 
segment V, (3) the presence or absence 
of antennal microtrichia, (4) the pres- 
ence or absence of antennal annula- 
tions, (5) the tendency toward fusion 
of antennal segments, (6) the degree 
of ornateness of the setae, (7) cuticular 
sculpturing, (8) the presence or absence 
of cuticular sclerotization, (9) the pig- 
mentation of the cuticle, (10) general 
body size, (11) the modification of 
setae into spines on abdominal segment 
IX, and (12) the presence or absence 
of a posterior comb on abdominal seg- 
ment IX. 


The characters used in assessing the 
phylogeny of the Sericothripini were: 
(1) the distinctness of the suture be- 
tween antennal segments IV and V, (2) 
the density of microtrichia on antennal 
segment IV in larva I, (3) body size, 
(4) the amount of cuticular pigmenta- 
tion, (5) the presence or absence of 


Iuuinois NATuRAL History SurRvEY BULLETIN 


Vol. 31, Art. 5 : 
hypodermal pigmentation, (6) the pres- 
ence or absence of brown sclerotized — 
body areas, (7) setal length, (8) the 
degree of setal ornateness, (9) the 
presence or absence and the position 
of setae, (10) the presence or ab- 
sence of setal basal rings, (11) the 
density of the cuticular microtrichia, 
and (12) the presence or absence of © 
cuticular pustules. 


4 


In selecting these characters and in 
determining their primitive and derived - 
states, it was assumed that: (1) charac- 
ters found mainly in primitive groups 
are primitive, and (2 ) characters re- 
garded as primitive in adult Thysanop- — 
tera (Stannard 1968; Gentile & Bailey 
1968 ) might be supposed, with reserva- _ 
tions, to be primitive in the larval 4 
stages also. Large body size, mod- 
erately ornate and long antennal seg- — 
ments, greater degrees of coloration, — 
moderately ornate setae, the presence — 
of cuticular microtrichia, lack of body — 
pustules, lack of a posterior comb on — 
abdominal tergite IX, and setae modi- — 
fied into spines on the terminal abdomi-_ 
nal segments were considered to be 
primitive features of the Sericothripini — 
and of some other tribes of the 
Thripidae. 


Each of the characters was assigned — 
a value from 0 to 2 for each Illinois — 
genus of the Terebrantia and for each ~ 
species of the Sericothripini found in” 
Illinois. A value of 0 indicates a plesio-— 
morph or primitive condition for the — 
character in the group or species; a 
value of 1, an intermediate or variable — 
condition; and a value of 2, the — 
apomorph or derived condition. The 
character states and values are sum- — 
marized in Tables 2 and 4, and scores — 
and sums are summarized for 29 genera — 
and one family in Table 3 and for Pel 
species of the Sericothripini in Table 5. — 
The sum of the values for the 12 
characters gives a measure of the de- 
gree of divergence of the taxon from — 
the primitive, ancestral stock. These — 
values are shown graphically in Figg 


CERRADO kien Fe fcr ge. 


ieee 


Pe 


August, 1974 


2 and 4, and the inferred phylogenies 
are represented in Fig. 3 and 5. 


PHYLOGENY 
OF THE THYSANOPTERA 


The Aeolothripidae have generally 
been accepted as representing the most 
primitive group because of their simi- 
larities to the more primitive Cor- 
rodentia (Psocoptera ) (Stannard 1957). 
According to Stannard (1968), the 
Merothripidae and Heterothripidae are 
of more recent origin, and the Thripidae 
the most recent of the Terebrantian 
families. The Tubulifera, according to 
Stannard, evolved from a phyletic line 
related to the Heliothripinae of the 
Thripidae, the evidence being the many 
similarities between certain members 
of the two groups and the many spe- 
cialized features of the Tubulifera. Gen- 
tile & Bailey (1968), however, believed 
that the Merothripidae and Thripidae 
evolved from the Heterothripidae, and 
Priesner (1926b-1928) felt that the 
Merothripidae represented a possible 
link between the Terebrantia and the 
Tubulifera because of certain inter- 
mediate features found in merothripids. 


The phylogenetic and _ systematic 
status of the tribes in the family 
Thripidae have been much debated 
because of the difficulty in delimiting 
groups at this level. Stannard (1968) 
recognized the Sericothripini, Dendro- 
thripini, and Thripini but did not sep- 
arate the Chirothripini and Anapho- 
thripini because they were difficult to 
categorize. Gentile & Bailey (1968) 
suggested a phylogenetic sequence for 
the tribes, from most primitive to most 
advanced: Heliothripini, Anaphothrip- 
ini, Chirothripini, Sericothripini, Den- 
drothripini, and Thripini. These au- 
thors indicated that the Thripini have 
become specialized by degeneracy. All 
of these phylogenetic arrangements 
were derived, primarily, from studies 
of adult characteristics. 


An interpretation of the higher Thy- 


VANCE: LARVAE OF THE SERICOTHRIPINI 


157 


sanoptera phylogeny, based on larval 
characters, can be depicted as in Fig. 3. 


Most larval features in aeolothripids 
were assumed to be primitive although 
lack of color and pigmentation seemed 
to be an advanced trait. This family 
is characterized by such primitive larval 
features as spines (modified Al and A2 
setae) on abdominal tergite IX; large 
body size; cuticular sculpturing lacking 
pustules; complete anterior and _ pos- 
terior tentorial arms (personal com- 
munication, B. $. Heming, 31 January 
1972); antennal segments III-V elon- 
gate and segments II-VII strongly an- 
nulated, with prominent microtrichia. 


From the Aecolothripidae two phyletic 
lines seem to have emerged: the mero- 
thripid-phlaeothripid (Tubulifera) line 
and the heterothripid-thripid line. 

The merothripid line is characterized 
by the retention of a smooth cuticle; 
long fifth antennal segment; large body 
size; the loss of antennal annulations 
and microtrichia; complete anterior and 
posterior tentorial arms (personal com- 
munication, B. $. Heming, 31 January 
1972): and a tendency toward the 
fusion of antennal segments VI and 
VII. The heterothripid line is charac- 
terized by the retention of antennal 
annulations and microtrichia; the de- 
velopment of cuticular pustules; small 
body size; the reduction of the fifth 
antennal segment; and, occasionally, 
the fusion of antennal segments VI and 
VII. 

The merothripids are more special- 
ized than are the Aeolothripidae in the 
elongation of antennal segments III—V 
(V remaining equal to IV), the reduc- 
tion of antennal annulations, the loss 
of annular microtrichia, the fusion of 
antennal segments VI and VII, and a 
partial reduction of the spines on ab- 
dominal tergite IX. Merothripids re- 
tain such aeolothripid features as ab- 
dominal spines, simple setae, smooth 
cuticle, large body size, and antennal 
segment V unreduced and equal to seg- 
ment LY. 


158 


Intivois NATURAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


Table 2.—Phylogenetically significant characters of the Thysanoptera and their charac- 
ter states and values. 
Character Number and State Value* 
; I Antennal sezments, cloneated —- coe sc co ieie en cers ote © vue wteyoueetsie she ee ree roatena 
Antennal sexments not Cloneated fei. << cect «0 o.elcie oe 'cre'h «0 ines 0:eceke areiei ene 2 
Il Antennal'sezment’ V long: oi. ooo oe cicada ans elajets eteisiel> afeve aie» cls terstalte eee Sa!) 
Antennal’ sepment V reduces es ores 5k oie cee» erenite: oc hoie lol'vin len eyernyee= ieee ee eee 2 
III Antennal segments with prominent microtrichia ...........-..2.eeeueeeeee 0 
Antennal segments with microtrichia reduced ..........-.-0 cece cece eee eee 2 
IV Antennal segments with prominent annulations ..............00.0-e eee eeee 0 
Antennal segments with annulations reduced .............++seeeeeeeeeeeee 2 
V_ No fusion) of antennal) Sez memts sete. of ace ohne ota e ja = oven ni Loe! =pnpol ese ekelel ye eee O” aa 
Fusion of certain antennal segments ............ cece ete eee e ee ee ne eee ene 2 
VE  Setae ornate OF: LOWS, jess sejecece serene Segrttee odin a wie at aderevege. cash cia chal es ates ee ee 0 
Setae simple ‘or SHOLE: . 6 sie cis ore cue a os unt ener n ale el ehsvese is clnieveln| ete eles tana 2 
VIE ‘Cuticle’ without pustules ooo c-c are co cin n'a) etetel ols) craletetel oleteratchetel erat <teteat stee tenet eaes 0 
Cuticle with pustules: 2. esc cece calc = cteletole nile letereayets siete fallelatS she a aeane nna 2 
VIII Brown sclerotized body areas present .......... cece eee eee eee eee teens 0 
Brown sclerotized body areas lacking .........-...+-.eesceeeecsccrencecees 2 
IX Prominent cuticular and hypodermal coloration ................0+eeseeeee 0 
Little cuticular and hypodermal coloration .......--......eee eee eee eee eeee 2 
DO Body Jar ee ah acéesniies ess aioce cteneo apeaetesevale:(o' 1" 'orevaste Nes aes fal EE ea ie RRO GR Cea 0 
Body SMA 1). . acetoccic 2.0 c adeno nceis aie. ne aoc obaniwteyhubve ehspetalepaete pele eles Hasek nee ea ae 2 
XI Setae on terminal abdominal segments modified into spines ................ 0 
Setae on terminal abdominal segments not modified into spines ............ 2 
XII Posterior comb lacking on abdominal segment IX .............--.-2.e00eee 0 
Posterior comb present on abdominal segment IX ...............-+-22+00-- 2 


8 An intermediate or variable state was given a value of 1. 


The Tubulifera are more specialized 
than the Merothripidae in the total 
loss of antennal annulations and of 
abdominal spines on tergite IX, but the 
two groups are similar in the retention 
of a smooth cuticle, large body size, 
a long fifth antennal segment, and oc- 
casional fusion of antennal segments 
VI and VII. 

Just as the merothripids are pos- 
sibly intermediate between the Aecolo- 
thripidae and the Phlaeothripidae, so 
the heterothripids may be intermediate 
between the Aeolothripidae and the 
Thripidae. The heterothripids retain 
such aeolothripid features as annula- 
tions on antennal segments IJ-VII and 
prominent spines on abdominal seg- 
ment IX but also have characteristic 
thripid features, such as reduced third 
and fourth antennal segments, the re- 
tention of antennal annulations and 
microtrichia, the reduction of the ten- 
torium, and the development of cuticu- 
Jar pustules. One feature that is ob- 


viously intermediate in the Hetero- 
thripidae is the length of antennal seg- 


ment V; in aeolothripids it is equal to — 


the length of segment IV, and in © 


thripids it is reduced to less than one- 
fourth the length of segment IV. In 
the heterothripids, however, antennal 
segment V is about one-half the length 
of segment IV. 

The Thripidae have retained antennal 
annulations and microtrichia, but the 
length of antennal segment V has been 
greatly reduced, body size has become 
smaller, and cuticular pustules have 
appeared. Many features vary from a 
primitive to an advanced state within 
the group. 

The family Thripidae shows con- 
siderable specialization and diversifica aa 


tion. As the Tubulifera became spe-— 


cialized into a fungus-eating niche, the 
Thripidae diversified into a phytopha- 
gous niche and tended toward an evolu- 
tionary degeneration or simplification of 
many characters. Since the Thripidae 


“artes 


Se ENN I 


\ 
y 
a 


S 


Z 


. : 


August, 1974 VaNcE: LARVAE OF THE SERICOTHRIPINI 159 
Table 3.—Character values for genera and one family of the Thysanoptera. The higher 
a group’s total of character values, the more advanced the group is interpreted to be. 
3 Character* nee 
ee Ri elias: VV VIL. IX ix, kl ckin a. 
Aeolothrips 0 0 0 0 0 1 0 2 a 0 0 0 4 
Franklinothrips 0 0 0 0 0 il 0 2 0 0 2 0 a) 
Phlaeothripidae iit 1 2 2 2 0 0 il 0 0 0 0 9 
Merothrips 2 il 2 i 2 1 0 i il 0 0 0 11 
Heterothrips 2 af i 0 0 0 2 2 2 i 0 0 lal 
Caliothrips nf i 2 0 0 0 2 1 0 2 2 0 11 
Heliothrips 1 1 2 0 0 2 2 2 1 2 2 0 15 
Hercinothrips 0 il 2 0 0 2 2 iL 2 2 2 0 14 
Parthenothrips 1 1 2 0 0 0 2 2 2 2 2 0 14 
Limothrips 2 1 2 2 0 i 1 0 1 2 2 0 14 
Chirothrips 2 2 2 2 2 2 1 ab 1 2 2 0 19 
Chilothrips 2 2 2 1 0 1 2 2 0 2 0 0 14 
Oxythrips 2 2 2 1 0 1 2 1 0 2 0 0 13 
Aptinothrips 2 2 2 il 0 2 2 ab 1 2 2 0 17 
Chaetanaphothrips 2 2 il 1 0 0 2 2 1 2 2 0 15 
Anaphothrips 2 2 1 1 0 2 2 0 1 2 2 1 16 
Echinothrips 2 2 0 1 0 0 1 2 2 0 2 0 12 
Sericothrips 2 2 0 1 al 0 1 1 1 2 2 0 13 
Zonothrips 2 2 0 1 1 0 1 2 1 2 2 0 14 
Drepanothrips 2 2 0 1 1 il 0 2 2 2 2 0 15 
Scirtothrips 2 2 0 1 1 1 0 2 2 2 2 0 15 
Dendrothrips 2 2 0 1 1 1 1 2 2 2 2 0 16 
Pseudodendrothrips 2 2 0 a il il 1 2 2 2 2 0 16 
Leucothrips 2 2 0 1 il 1 i 2 1 2 2 0 15 
Ctenothrips 2 2 it 1 2 2 2 2 2 2 2 2 22 
Scolothrips 2 2 uL al 2 0 2 2 2 2 2 0 18 
Thrips 2 2 a 1 2 2 2 1 1 2 2 2 20 
Frankliniella 2 2 1 1 2 2 2 2 2 2 2 2 22 
Taeniothrips 2 2 al 1 2 2 2 2 1 2 2 2 21 
Microcephalothrips 2 2 it 1 2 1 2 1 2 2 2 1 19 


"See Table 2 for characters and their states. 


show a more pronounced delimitation 
of groups, this phyletic line probably 
originated earlier than did the Phlaeo- 
thripidae. 


PHYLOGENY OF THE TRIBES 
OF THE THRIPIDAE 


As when using adult characters, de- 
limiting tribal groups of the Thripidae 
is also difficult when considering larval 
characters. As can be seen in Fig. 2, 
all tribal groups except the Thripini are 
derived to about the same extent from 
primitive stock. Since each group ex- 
hibits certain specializations and evolu- 
tionary advancements, it is possible that 


some of the tribes do not follow a 
single line of phyletic ascent but rather 
follow several separate lines, each being 
fairly independent of the others and of 
an origin ancient enough that many 
intermediate types have disappeared. 
Although easily distinguished, the 
Sericothripini and Dendrothripini show 
many similarities, possibly indicating a 
divergence of these groups later than 
the divergences of most others. One 
feature unique to this phyletic line is 
the pattern of microtrichia on antennal 
segment IV of the first-stage larvae. 
The microtrichia are found randomly 
placed between as well as on the an- 


160 Ixtuinois NaturaL History Survey BULLETIN 


nulations, whereas in other groups they dae the retention of annulations on ; 
occur mainly on the annulations and antennal segments V-—VII, less reduc- 
only sparsely between them. tion of segment V than occurs in most 
The most primitive subfamily in the — thripids, an elongation of segment VII, — 
Thripidae is the Heliothripinae. This and, in some genera, an elongation of 
subfamily shares with the Heterothripi- segments III and IV (this latter feature 


7 
\ 
4 


e 
e 
—-—~< 
Thripini 


e 
ss 
er Dendrothri 


pini 


e 
‘= 
ia 2 
tenes hg a 
Cost 
i ee 
- Ce o 
Ee Gane 
= 
g a 
Snagit < 
Ra a 
= e 
Qa e 
5 \ 
= ? \ 
2. Ne w 
aS ae 
3S “XN a 2 o 
= oO 
gc = 2.9 
— “= cw wo 
= ~~ (Ty Se 
>->~_ d = S28 
Se) eee 
ie ake 2 
pf ao) 
\- \22 3 Es 
my) = 5 
fo} 
i i e 
(e a; 
nw] a 
oh 
So) 


N Oo 
A eRe! SRB cacy Oh by eg rv ene ae 
a 
a) SANIWA YALIVUYVHI 40 1VLOL 
FO 
n= 
ox 
=> 
a 
<= 


“SdLayzoluael 

BL latulyuesy 
“sdrayy 

Sdt4yzO [00S 
‘Sdtayyoues9 
“‘sdtayzoone] 

Sd dy} 0Apuapopnesd 
“‘sdtayzospuag 
“SdL4YzO9 ALIS 
‘Sdiayzouedesg 
‘sditsyzouoz 
“sdtuyz0ol4as 
“sdiayzoydeuy 
‘sdluyzoydeurzery9 
“sdtuyzourzdy 
“sd Layzhxo 
“Sdrayzo4ryy 
SdLayzOwLy 
SdLayzouaYyzAed 
‘Sd LAyyOULIUSH 
“SdLAyzOL [eH 
sdpayyorleo 
Sdt4y}04979H 
‘sd1ayzO4aW 
aepidiuyzosel| yd 
Sdragyourlyuew4 
“Sdr 4470 [| 08y 


2 


MOST 
PRIMITIVE 


Vol. 31, Art. 5 _ 


wan 


Table 


ly of the Thysanoptera. These comparisons 


1 


in Table 2 and totaled i 


g 29 genera and one famil 
ru 


al 


he degree of derivation amon 


ig. 2.—Comparison of t 


Fi 


161 


: LARVAE OF THE SERICOTHRIPINI 


VANCE: 


August, 1974 


the Helio- 


with 


itive features 
thripinae. The genus Limothrips, like 


The tribe Chirothripini shares several 


are with prim 


reapproaching that in the Aeolothripi- 
the Thripinae a partial reduction of an- 


dae). The Heliothripinae sh 


the Heliothripinae, has a reduction in 


tennal segment V and the loss of spines 


on abdominal tergite IX. 


the number of annular microtrichia and 


brown sclerotized body areas. Segment 


The Heliothripinae have such primi- 


tive thripid features as large pustules, 
_ brown sclerotized body areas in many 


V in the antennae of first-stage larvae 


of Limothrips is elongate and has two 
annulations, a feature found only in 


luding the 


mc 


? 


more primitive groups 


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fo sulBii0 ajqeqoid ay} sayesipul AuabojAyd siy| “sa}deseyd jeAse] uo paseq ‘esa}douesAyy ay} Jo AuaBojAyg—'e “614 


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uo aezas $sajnysnd ynoyzim aloizng 


abptug LelLvoquaz yo ssol 
$paonpeau p pue ¢ sqUawbas |euUuaquy 


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SSO, £Salnzsnd uwe_ndi3nd 4o yUuawdolaaap 
fg quawbas ,euuazue 40 UOLZINpau [elLy4eg 


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{XI yuawbas Leutwopqe uo sautds 4o ssoq 


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a }euauabaqg 


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4OLuazsod 40 Sso| 
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$aqnz e OjUL X Yuaubas 
[eulwopge so jUuew 
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[euLwopqe uo sauids 
pue suo.zeinuue 
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LeuLwopqe uo ssa] a40W 
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UOLZPEILSLSUBALP Yeaub 
{Z quawbas jeuuazue 
$0 uoLzebuoly 


= i=] wn > ° = = ac) = > 
=x m m =z = m m cane ate m™m al 
partes! = a > - - a 4 pat oO 
Ne i] L=J uv =] Coal m c > oO = 
<eerct cia woo ox ano wo wn opm ouo +O 
worn - Oo = a) Sa a4 =) cwo Ries 4 
ws wa a4 ite == nx na ms nw —=x 
= 32x wx wx Ee] 2D wx ax Be Ce] 
Es) a =] = = a new uv = 

7 _ ry ~ ~ _ m _ - ~~ 

uv n! uv Lal _ ao] ~ (ee o _ 

~ Land cd = a Leal ens ad > i=] 

= = = i > i=] ea m > 

= al = m > > m 

m m 


eras) SLED 


AVGIdIYHL 


162 


Ituinois NaturAL History Survey BULLETIN 


Vol. 31, Art.5 — 


Table 4.—Phylogenetically significant characters of the Sericothripini and their character 


states and values. 


Character Number and State 


Value* 

I Antennal segment’V distinct in larva I ov .cc6. 2 cise oc ccc ee cee elena eee 0 
Antennal segment V partially fused to segment IV in larval ............... 2 
Il Microtrichia densely placed on antennal segment IV ....................-5- 0 

Microtrichia sparse on segment IV ........... ccc cecceccvcceccccceces 7 2 : 
TID Darge Dody wcrc): ci stetepateyete alahe o eterele le, « =. cleieiais aucjeiers aisvaieiele: cle ietele soit ea 0 
Small DOGG: sjericejart oi aye. o: -cavayeys oisdelaisueyayees, « abjrareyeus (sone ve lop sttege, oi ol uslels ber) OE 2 
IV Cuticle darkly pigmented (oo... 6 cicrsvc1e os) sixe c:e «//cle © wicte) oie le)oieielstats aye ieee 0 
Cuticle lightly pigmented co.cc sce cs cs wise wieianslcleyeite oly cia) el ole ei ccereieietel ene arate 2 

V Hypodermal pigment present .............0- cece cere cence o/s 6 oats ata epee eaneteiee Oo. 

Hypodermal pigment absent .... 2... sce sce fees cw snc sae oe ele ole)aleneeenn a ianeaaen 2 
VI Brown sclerotized body areas present ............00cscsceceeveceesserucven 0 
Brown sclerotized body areas not present ........... ese eee ee eee ce cnet tees 2 
WIL" Setac LOme 2 sicciec cseceieieiers sila’ cere a-0 eho oteveleiege © are jarw droves [eke ce evel ie CTera eae tiene ea 0 
Setae: SHOLE. s:.j5 5 sie a vraece'e wie so 0:0 leynie euslel ofane.crale) ote ie ce coe) oetaPe iene cist eae ae 2 
WIIL  Setae Ormate (oo. ecieseisreiaiscin pis iors atevsne sab cls feck crctetw s, wy ece)e-mpeiinge) eke sehr ea eee 0 
Setae not ornate (simple) ....... Pe ee ey ee re Pe SS oti ce camnn « 2 
IX Some setae reduced or lackimge 2. 3..).c.ci5 Me «ufc: rere vccie apaitvs 00 + 0 0) eee enone 0 
All setae normally present 2 o <5 60 cts cic ees cet o viet oils cleo vie crelens ele shel nee er enS 2 
X Setal basali.rings Present. 6 0... Wo dcece.c chs 2.» Scapa sioposeconndesert eittere eile eRe een en 0 
Setal basal rings: absent, ..6.5.:5 cicl0.s-Fesd ctcleieenetets ete ages 00 open eee 2 
XI Microtrichia dense on cuticle . 5.406 cd cos ce sues seis te.e classe eee ee 0 
Microtrichia ‘sparse on cuticle 3 .()6 012% fc «cj 2 nie o «cle oieiel = eialela)ee) ohio 2 
MIL“ Cuticle: with pustules. <1..0 5 sk cS eBew.« fee ad ols sca ceva ce ee oles eee ee 0 
Cuticle without pustules... dlc. sec c.e cic ecco wes 0 op 001s 6. «(0 ee eee 2 


. An intermediate or variable state was given a value of 1. 


Heliothripinae. The second-stage an- 
tennae of Limothrips are typical of 
those of the Thripinae. 


Within the Chirothripini, Limothrips, 
with normal antennae, long knobbed 
setae, and brown sclerotized body areas, 
is most primitive. Chirothrips exhibits 
an extreme evolutionary degeneracy 
and specialization. Some Chirothrips 
larvae, at least, spend their whole ex- 
istence within a grass floret (Watts 
1965); the antennae and legs of the 
larvae are greatly reduced, the setae 
are minute and pointed, and little 
brown sclerotization is present. 


The Anaphothripini also share many 
primitive traits with the Heliothripinae. 
Cuticular pustulation and the lack of 
microtrichia are very similar between 
the two groups, and many Anapho- 
thripini have brown sclerotized body 
areas similar to those in the Helio- 
thripinae. Genera such as Oxythrips 
and Chilothrips show a modification of 


certain setae on the terminal abdominal — 
segments into setaelike spines, a con- 
dition found only in the primitive fami- 
lies. 


The Anaphothripini have antennal 
segment V reduced in the first-stage — 
larva, and have shorter antennal seg- — 
ments, less ornate setae, shorter anal 
setae, and less diversification than have 
the Heliothripinae. 


The tribe Sericothripini is transi- 
tional between the more primitive and 
the more~ specialized tribes in the 
Thripidae. The Sericothripina share, 
in some species, several characteristics — 
with the Anaphothripini, and the Scirto- 
thripina with the Dendrothripini. The — 
annulipes group of Sericothrips have 
such seemingly primitive features as 
brown sclerotized body areas, hypo-— 
dermal pigmentation, large basal rings — 
on the setae, and in Sericothrips cingu-_ 
latus small cuticular pustules. Except 
for cuticular pustules these features are 


August, 1974 


Table 5.—Character values for members of the Sericothripini. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


163 


The higher a group’s 


total of character values, the more advanced the group is interpreted to be. 


= fer ur ae Vv 
Echinothrips 0 2 0 1 2 
Sericothrips 
cingulatus 2 0 1 0 2 
pulchellus 2 0 il 0 0 
annulipes 2 0 1 0 0 
variabilis 2 0 0 0 0 
baptisiae 2 0 1 2 2 
campestris 2 0 i 0 2 
beachae 2 0 af 2 2 
sambuci 2 0 1 aft 2 
tiliae 2 0 1 1 2 
nudilipennis 2 0 a 2 2 
langei 2 0 1 1 2 
Drepanothrips 
reuteri 2 ? 2 2 2 
Scirtothrips 
niveus 2 0 2 2 2 
taxodii 2 0 2 1 2 
brevipennis 2 0 2 1 2 


Character* 


VI Vil VII Heke! 


ES XS XT IE 


2 0 0 0 2 2 al 12 


NNYONNNNNNOOS 
ee Oe 
oe — i — i — i — 
pwywnwmymnwhyh bp 
PHENO rP HH OOOO 
DNONNNNHNYNNND 
a 

= 

~ 


bo 
bo 
bo 
bo 
bo 
io 
bo 
bo 
rare 


oS 


bo bo po 
bo po pb 
bo bo bo 
ee ) 
bo bo be 
i) 
bo bo bo 
be 
=) 


®See Table 4 for characters and their states. 


usually not found in the more advanced 


'Thripinae. The tiliae group of Serico- 


thrips also lacks these features. The 
Sericothripina and Echinothripina are 
plesiomorphic in their ornate, fringed 
setae and in certain other characteris- 
tics; in the Scirtothripina, however, the 
setae are reduced. The Scirtothripina 
are smaller, lack body coloration, and 
have shorter setae that are only termi- 
nally funneled, all seemingly indicating 
a more derived condition than those 
found in the Sericothripina and Echino- 
thripina. 

The Dendrothripini are similar in 
their morphology to the Scirtothripina, 
indicating close phyletic relationships. 


Of the tribes in the subfamily Thrip- 
inae, the Thripini is the most spe- 
cialized, lacking body coloration and 
brown sclerotization, having reduced 
setal ornateness, and exhibiting a gen- 
eral lack of diversification. The pos- 
terior comb of abdominal tergite IX 
is an advanced characteristic found in 
several genera in this group. Cuticular 
pustules and microtrichia are often re- 


duced, and certain genera (e.g., the 
Frankliniella-Taeniothrips-Thrips com- 
plex) lack divergence in their larval 
stages, indicating close relationships be- 
tween them. 


PHYLOGENY 
OF THE SERICOTHRIPINI 


There appear to be at least three 
subtribal groups in the tribe Serico- 
thripini (Fig. 5). The Scirtothripina is 
seemingly the most specialized, and the 
Sericothripina is the most primitive. 
The Scirtothripina have reduced body 
size, reduced body coloration, and re- 
duced setal ornateness, and lack sclero- 
tized body areas, all of which are de- 
rived characters. Lack of cuticular 
pustules and proliferation of cuticular 
microtrichia may be specializations in 
this group. 

Of the Scirtothripina genera, Dre- 
panothrips is here considered the more 
primitive because most body setae in 
the members of this genus are termi- 
nally funneled, whereas only certain 
ones are so in Scirtothrips. Most body 


" 
164 Inuinois NAturAL History SurvEY BULLETIN Vol. 31, Art.5 — 


setae in Scirtothrips are short and In the Sericothripina the genus Seri- — 
pointed, and cuticular pustules are cothrips contains two subgroups, the 
totally lacking, these two features con- annulipes and tiliae groups. The tiliae 
sidered here to be derived. group is characterized by light body 
The Scirtothrips species show a pat- coloration, a lack of hypodermal pig- 
tern of setal simplification. S. niveus mentation, reduced brown sclerotized 
has four pairs of setae terminally fun- areas, and reduced rings at the bases — 
neled, and S. taxodii and S. brevipennis of the setae, all considered here to 
have only two pairs of funneled setae, be derived conditions (Fig. 5). The 
the latter situation being the derived annulipes group is characterized by the ~ 
state. presence of cuticular and hypodermal 


: oe 
x = :@ 
- =) o3 
. ° o's 
& © ae 
~ D i= ae 
° Fo, 
Pp w - a 
S o = a 
a 0 = at p 
= ”n = toe 
= eS 52. 
= & 
~ cn a 
3] ey - > StuuadlvAadg Sck 
o| ff Pe ae = LLpoxeq ae 
2 Sea 
o levy £ = SnaALu 3o2 
a; NN } 
o wn SdLUY}OFULIS Gua 
2 ow — he 
AN ‘< —=—_—— afe 
Ve) = = Lua znead oor 
nn r= t 
s sdiayjzouedesq eg 
S aoe 
—= = o— o 
a> 
{2 @ rabue, Beg 
o-— 
/ e\ wo Stuuedi{iqnu 9.8 
/ oN = ———— oe 
\ a eetll? =e 
! » rid 
\ 2 ponques eg* 
| oO a Sod 
} Gd aeyoeeq pes 
\ i s o*o 
: 2 2 Stujzsadwes S67 
ate arg ae 
Roe sets tadeq a 
BN StLiqelueA Sage 
n we] 
2 sadi|nuue 20% 
£ Sn[[eyoind 2G 
& EEN £25 
» oO 
5 Sn}e|[NbuULo re | 
- ~ 
= Sdtayz0otdas 48 
oO ces 
wl te Sdiayzouryoq gee 
=. 
ae) 
& o aya 
ban peed RE Ba DR eae EQS 
5 ar 
a Lu 159 
an SANTWA YALIVYVHD 40 TWLOL z OF 
ez —e v6 f 
x Ne 2D ain 
=> o= “ws 
<x a oa 
a abe 


August, 1974 


pigmentation, brown sclerotized body 
areas, occasional enlarged rings at the 
setal bases, and, in one species, by 
small cuticular pustules, all suggested 
here to be the primitive state. 

The tiliae group includes campestris, 
beachae, sambuci, tiliae, nubilipennis, 
baptisiae, and langei (Fig. 5). The first 


VANCE: LARVAE OF THE SERICOTHRIPINI 


165 


five species all have long body setae, 
and the other two, short setae. S. 
campestris resembles members of the 
annulipes group, having orange cuticu- 
lar pigmentation and generally long, 
wide body setae. For these reasons 
campestris is considered the most primi- 
tive of the tiliae group. S. beachae, S. 


~ ye we 
a _ Oo 
ow wo 
=) mr iS 
Stuuadtasaq c ied = 
StuuedtAaag S ey fe) 
n = = 
Laan oa sf o 
= ee EE — +. 
TLpoxez = = ° 2 
TEP : aS & oe 2 
a Le ou 0 
ne ec Oo a cen cs & 
Ba a EE wEo [e) 
- «6 n & [a4 ” — = 
= = £@o a 
Tey 2 oe 
” u-w o 
=e -x< 0 osu acs 
sn3sAltu~—= = ow a0 ~ 
Sn rr e 
eo<cto 4 spe o 3 
oH = = esa oF ae 
eve mos vv a 
Ch) =< “oo Oo 
no 8 x= od = 
qasara 5 22 2 
c 
t4ezned z oc ra 
a 
o a rr = 
o rv) » oO 
es) 4 a ro 
Tobuel a a 3 8 
ie = = 
» = we 
Beisizdeq eq 5 o [au 
» 
= ou <= 2 
Spousdtprqna n nou = = 
uUSdL[L ow 
St Liqnu een a = 
outs = m 
oe ouvuo-™ nw 
aPlLt} g Sere = D 
s a = = 
o wuve So rs} 
a = soos 5 o 
Lonques c2aD = pe 
o on 
c Lunvy o 2e 
= ° ood ” Vv 
aeyoeaq 2a — ON =a 
Orr o 
OLer = 
o+| 
Stajysodwey yet _— 
Sluzsadwed A ae 
oun 
ee) s 6 
SULLqeLdeA ae = = = 
| a4 = 5} 
=x o hr 
SS S E g 
- 
SM[Leyo[n = = = He) 
= = x pan 
; _ o 
Sadt[nuue Re ae, 2 a 2 £ 
3 Sn ==! a — a 
sod ciao o ew = 
o —- One =x os 6 
ca Le OD = ANA ~ 
= w as “ ° ov ° 
oO uo al a a = Pa 2 
“ OYE © 
p Ns poe = co EU o 
au oY, a ae =x 7 or ao n 
eoo re 
SN7e[NBULS OY HOO noses z =o o 
-3 suo wuac = as re oc £ 
crv eos morc oO —s = > 
oun —-ow sSacnd N oo o ee ue 
Es OL o> uo jo sE£ ° 
na noo SSE o rr ==. 
= ay ee = = 
aepunuso = we Ee ee = c 
=x Oc wo -> = i=) 
wn” = mo YQ ww = me} 
— a = = su Ww 
Snuedlaowe Se a ia Oo” 8 = ze 
= = oo shake e— oO 
= i-3 > eow Oo 
FS oe orn Pz 
So = ae £ = s 
= So 2 cow a n 
= 4 = orp ay 
——— —————__ al ” ” ~ 
snaelyqns aS = —'m ae 7) fe) 
tu oO vt vos iL 
tu ow 3sf+o 
a Pr AEN 
oo Vere 
nu ea} = S) 


the taxa. 


166 


sambuci, S. tiliae, and S. nubilipennis 
are light colored and have long setae: 
beachae and sambuci have wide setae, 
and tiliae and nubilipennis narrow 
setae. The larvae of the latter two 
species are seemingly indistinguishable, 
suggesting that they separated rela- 
tively recently. S. baptisiae and langei 
are considered to be the most derived 
of the tiliae group because of their 
setal reduction, those of langei being 
so narrow as to approach the condition 
found in the Scirtothripina. 

The annulipes group includes cingu- 
latus, annulipes, pulchellus, and pos- 
sibly variabilis (Fig. 5). The most 
primitive species is cingulatus, as evi- 
denced by its brown body areas, the 
presence of small cuticular pustules 
similar to those of the Chirothripini, its 
pteronotal sclerotized plates, and _ its 
enlarged setal basal rings. S. annulipes 
and pulchellus have a reduced amount 
of brown coloration and fewer pustules, 
but possess pteronotal sclerotized plates, 
setal basal rings, and hypodermal pig- 
mentation. S. variabilis lacks pteronotal 
sclerotized plates, but has hypodermal 
pigmentation and somewhat enlarged 
setal basal rings. 


The evolutionary status of the Echi- 
nothripina is uncertain. Echinothrips 
species have large body size and ornate 
setae, both primitive features, but also 
show derived features, such as weak 
body coloration and sclerotization and 
a reduction in certain setae. 


SYSTEMATICS 


The known larvae of the Thysanop- 
tera of Illinois are described here at 
the family level and for the suborder 
Terebrantia at the subfamily and tribal 
levels. Genera and species larvae are 
described only for the tribe Serico- 
thripini. Larval descriptions pertain to 
the second-instar larya unless otherwise 
stated and include as little repetition 
as possible from higher to lower groups. 
A key is included to the major groups 
and to many genera of the Illinois 


Inuinois NaTuRAL History SuRvEY BULLETIN 


Vol. 31, Art. 5 


thrips fauna, and keys to the species 
of some genera of the Sericothripini are 
given. 

Measurements, taken with a cali- 
brated ocular micrometer, are ex- 
pressed in microns. They include 
lengths and widths of antennal seg- 


ments, antennal length, body length 


(excluding antennae), head and prono- 
tal length and width, and lengths of 
certain body setae, the particular setae 
measured depending on the genus con- 
sidered. The setal numbering system 
used in this report is given in Fig. 6. 


KEY TO IMMATURES 
OF THE THYSANOPTERA 


1. Antennae projecting forward, with 
distinct segmentation; wing pads 
absent (LARVA) 


Antennae short, projecting back over 
head or to side of head and indis- 
tinctly segmented; if antennae pro- 
ject forward, they are indistinctly 
segmented 


2. Antennae directed forward and in- 
distinctly segmented or short and 
directed laterally or directed poste- 
riorly along sides of head, not reach- 
ing anterior margin of prothorax 
(If antennae are recurved over head, 
wing pads reach only to second or 
third abdominal segment.) (PRE- 
PUPA) 

Antennae are recurved posteriorly 
over dorsum of head or along sides 
of head, reaching or surpassing an- 
terior margin of pronotum (PUPA) 4 


3. Antennae long and directed forward 
(recurved over head in Aeolo- 
thrips); wing pads, if present, ex- 
tending posteriorly only to second 
or third abdominal segment; ab- 
dominal segment X not tubelike 
COI eae Ora OIG Terebrantia Prepupa 


Antennae short and directed to side or 
if long, posteriorly directed along 
sides of head; wing pads absent; 
abdominal segment X tubelike or 
elongately conical. . Tubulifera Prepupa 

4, Antennae directed posteriorly over 
dorsum of head; wing pads, if pres- 
ent, reaching abdominal segment VI 
or VII; abdominal segment X never 
tubelike ............ Terebrantia Pupa 

Antennae directed posteriorly along 
sides of head; wing pads, if pres- 
ent, reaching to abdominal segment 


at Qe 


3 
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3 
we 
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August, 1974 VANCE: LARVAE OF THE SERICOTHRIPINI 167 


Il or III; abdominal segment X GENERIC KEY TO LARVAE 
tubelike or elongately conical .... OF THE TEREBRANTIA OF ILLINOIS 
\ 3 lo4 Se aoe doar Tubulifera Pupa 1. Prothorax usually with six pairs of 
5. Abdominal segment X never tube- setae; abdominal segment IX with 
like, usually broader than long; mid- three or four pairs of setae...Larva | 
dle antennal segments with micro- Prothorax usually with seven pairs of 
trichia-bearing annulations ....... setae; abdominal segment IX with 
NO etn afokniss<yeie a ae ij0'e Terebrantia Larva five or six pairs of setae (LARVA 
Abdominal segment X tubelike or HD) So ommocehnccoo a fed Oo Gor or dee 2 
elongately conical, usually longer 2. Antennal segment V from one-half 
than wide; middle antennal seg- to equal to the length of antennal 
ments without annulations (Fig. 21, segment IV (PRIMITIVE FAMI- 
1 SE hs (LC) Eee Tubulifera Larva TS Peet aes bee sels eee tenets 3 
sense cones 
___——antennal annulations . 
and microtrichia 
———Antenna H 
E 
A 
D 
___—eye_ facets 
cheek margin 
————Pronotum 
7 
___—mesothoracic spiracle 5 
Mesonotum R 
A 
X 
>__Metanotum 
SSS femur 
——_tibia A 
B 
D 
—tarsus 
O 
M 
Abdominal segments ; 
ery vi ______+——qbdominal spiracle 
d S 


32, = anal setae 


Fig. 6.—The external morphology of a Terebrantian larva (Sericotnrips [aiigei). 


168 


Antennal segment V much less than 
one-half the length of segment IV 
(usually about one-fifth as long) 
(THRIPIDAE) 

3. Antennal segment V one-half the 
length of segment IV (Fig. 22); seg- 
ments VI and VII with visible an- 
nulations 


Inxtinois NaTuRAL History SuRVEY BULLETIN 


aiets eiaisietpterete Heterothripidae 


(genus Heterothrips ) 


Antennal segment V equal or subequal 
to length of segment IV; segments 
V, VI, and VII with or without an- 
PULA RES | atta re ates, deste) esse al etal eter 

4. Antennal segments VI and VII 
fused; segments V—VII without an- 
nular rings, and segments III and 
IV not greatly elongated (Fig. 20) 


Merothripidae 


(genus Merothrips ) 


Antennal segments VI and VII not 
fused; segments V—VII with annu- 
lations, and segments III and IV 
elongated (Fig. 18) (AHOLOTHRIP- 


TTA) Me FSB cok ane eases eect ciate 5 
5. Four median dorsal setae on ab- 

dominal segment IX thornlike (Fig. 

Buiilics setae tires: pms stce sitass (seen aenoeae Aeolothrips 


Four posterior setae on abdominal 
segment IX not thornlike; not yet 
found in Illinois 

6. Antennal segment VII greatly elon- 
gated, length seven to eight times 
the greatest width (Fig. 25) (HE- 


Byercas Franklinothrips 


LIO TERIPINAE) ianiekee cele allan 7 
Antennal segment VII not greatly 
elongated, length only two to three 
times the greatest width (THRIP- 
PIA conte. ore Rie cunt hoe erates ea cera Ts 10 
7. Abdominal segment X with six anal 
BELAS Ei eycge tere sts nie eteieters oteeRs tee re eee ae 8 
Abdominal segment X with less than 
Six ANMASStAS Bike nis, cepereeeeeiaacshereces 9 
8. Body setae somewhat fanned for 
entire length (Fig. 46) ....Caliothrips 
Body setae moderately long and sim- 
ple, with hyaline terminal knob 
(Wig t43)) eerie h eke teins Heliothrips 


9. Body setae moderately long and 


widely funneled at tip. .Parthenothrips 


Body setae very short and simple 
(Fig. 65) 
Body cuticle generally smooth, with 
minute pustules or a stippling of 
fine microtrichia; abdominal seg- 
ment IX never with a _ posterior 
comb (CHIROTHRIPINI, SERICO- 
THRIPINI, DENDROTHRIPINI)... 


Body cuticle generally with raised pro- 
tuberances or pustules, with or 


10. 


sata lata cubagekoraie Hercinothrips 


11 


without microtrichia; with or with- 
out a posterior comb on abdominal 
segment IX (ANAPHOTHRIPINI, 
THRIPIND) +... aronscseere oe eee: 


All major dorsal body setae ex- 
panded and fimbriate or quite long 
and fimbriate, or if most dorsal 
body setae are small and simple, 
cuticle densely covered with fine 
but obvious (under high-power mag- 
nification) microtrichia (SERICO- 
THRIPINI)  ..:<.:.,a.. 901: 


Setae mostly small and simple and 
cuticle with stippling pattern devoid 
of obvious microtrichia (DENDRO- 
THRIPINI, CHIROTHRIPINI)..... 


Setae only terminally funneled, not 
greatly fimbriate (SCIRTOTHRIP- 
TINIAS) oon ce Sidi ee 


Setae fanned and fimbriate for most 
of length (except in Sericothrips 
langei) (SERICOTHRIPINA) 


Setae long, unexpanded, and fimbriate 
for most of length (Fig. 54) (HCHI- 
NOTHRIPINA) 


Cuticle with closely set microtri- 
chia; a maximum of four or five 
pairs of setae expanded, the re- 
mainder small and simple (Fig. 49 
and 50) 
Cuticle with prominent stippling and 
with less dense and less obvious 
microtrichia; all major dorsal body 
setae terminally funneled (Fig. 52) 


11. 


12. 


13. 


14. Associated adults with eight anten- 
nal segments; found statewide in 
Illinois 

Associated adults with seven antennal 
segments; found only in Volo Bog 


oeateceeee Echinothrips 


¢. ana arel sees aee e Scirtothrips 


Drepanothrips 


Vol. 31, Art.5 


19 


12 


15 


13 


14 


ays eps soot Gee es ee Sericothrips — 


in Lake County, Illinois....Zonothrips 


15. Eye facets large and eyes bulging 
at sides of head; brown sclerotized 
body areas lacking (DENDRO- 


THRIPINI)) 2 .2-46.)).62ee eee 


Eye facets reduced and eyes not bulg- 
ing at sides of head; brown sclero- 
tized areas present or if absent, 
antennae and legs greatly reduced 


(CHIROTHRIPINI) .............. 18 
16. Lateral abdominal setae with mi- 
nute terminal knobs ..... Leucothrips 


Certain dorsal body setae terminally 
funneled 


17. Lateral setae expanded only on ab- 
dominal segment IX (Fig. 71) 


Lateral setae expanded on abdominal 
segments II-IX and posteroangular 


16 


Dendrothrips 


$e LOIS; CLR AEP ELIOT SBME I 5 PNB, ete sitet a bch 


ai 


BESTS a 


August, 1974 


18. 


19. 


20. 


21. 


22. 


23. 


24. 


25. 


setae also expanded (Fig. 74) 
Pseudodendrothrips 


Antennae and legs greatly reduced; 
all body setae reduced and pointed 
AONE AAS) pia Mord ctaccte late c's 'etesare ba Chirothrips 


Antennae and legs not reduced; cer- 
tain body setae on posterior abdom- 
inal segments long and knobbed 
GAA) ce ate aig aicse mpelete. el ee Limothrips 


Cuticular protuberances without mi- 
crotrichia; abdominal segment IX 
without posterior comb (except in 
some Anaphothrips); body often 
with brown sclerotized areas; all 
or certain setae often knobbed or 
blunted (ANAPHOTHRIPINI) . 20 


Cuticular protuberances with or with- 
out microtrichia; abdominal seg- 
ment IX with a posterior comb (ex- 
cept in Scolothrips); body without 
brown sclerotized areas; setae usu- 
ally pointed (THRIPINI) 
Median and lateral dorsal setae on 
abdominal segment IX all nearly 
equal in length and width 
Median setae significantly shorter or 

thicker than lateral dorsal setae on 

abdominal segment IX 


Dorsal setae on abdominal segment 
IX pointed, with prominent rings at 
pases! (HiSH 6M) o.oc4 wc nr Anaphothrips 
Dorsal setae narrowly fanned, with- 
out fimbriation and without basal 
TUTE em A vonmee er enaitoie: aget Chaetanaphothrips 
Most body setae roundly blunt; mid- 
dorsal setae of abdominal segment 
IX shorter and much thicker than 
lateral setae and almost thornlike 
CES Es Gy) cee terete ayate ect ac, stern, arahbre¥oronye fol ets 
All body setae pointed; mid-dorsal 
setae of abdominal segment IX not 
thornlike; lateral setae long and 
wiih) @eeoanene see Aptinothrips 
Abdominal tergal sculpture in form 
of wavy, thickened, raised, trans- 
WHLSS Stlige | hack ca eee we Chilothrips 
Abdominal tergal sculpture in form of 
raised pustules arranged in trans- 
MENG PLOWS cr chersherte rete focie/=.c¥e Oxythrips 


24 


21 


23 


All body setae quite long (each 
epimeral seta—60um); posterior 
comb absent on abdominal segment 
eae (re. 56) ANG KU) tei a Scolothrips 
Body setae much shorter, normally 
less than 30-40 um; posterior comb 
present on abdominal segment IX.. 25 
Antennal segment IV reduced and 


shorter than the combined length 
of segments V—VII (IV about two- 


VANCE: LARVAE OF THE SERICOTHRIPINI 


169 


thirds the length of V-—VII) (Fig. 
il) Doefordids, srateretel nels mrs wee) oe Ctenothrips 


Antennal segment IV not reduced and 
equal to or longer than the com- 
bined length of segments V—VII.... 26 


Other Thripini genera (Baliothrips, 
Dorcadothrips, Frankliniella, Irido- 
thrips, Microcephalothrips, Odonto- 
thrips, Plesiothrips, Rhaphidothrips, 
Taeniothrips, and Thrips) larvae 
cannot be keyed at this time. 


AEOLOTHRIPIDAE Uzel (1895) 


Larva—Antennae (Fig. 8, 18, and 
19) each seven segmented; segments 
IlI-V elongate, II-VII with well- 
developed, numerous annular rings; 
microtrichia present on most annuli. 
Antennal segment V as long as or 
longer than IV. Sense cones (segments 
IV-VI) generally long and pointed. 


Head (Fig. 39) usually rounded from 
the dorsal aspect, with well-developed 
tentorium, mouth cone short and hy- 
pognathous; body elongate and cylin- 
drical. Setae usually long, moderately 
stout, and pointed or knobbed. Ab- 
dominal tergite IX (Fig. 57) with two 
median pairs of setae modified into 
stout spines (not modified in Frank- 
linothrips). Cuticle with fine micro- 
trichia producing a stippled pattern. 


Larva I lacking stout spines on ab- 
dominal segment IX. 


Diagnosis.—Larvae of the Aeolo- 
thripidae are easily distinguished by 
antennal features: segments III-V are 
elongate, annular rings are numerous 
on segments III-VII, and segment V 
is as long as or longer than segment 
IV. In heterothripids, antennal seg- 
ment V is about one-half the length of 
segment IV, and in the Thripidae, seg- 
ment V is greatly reduced and less than 
one-fourth the length of IV. 


In the Merothripidae, antennal seg- 
ment V is as long as IV, but both are 
relatively short, fewer annulations oc- 
cur on the antennal segments, and seg- 
ments VI and VII in Merothrips are 
fused. 


170 


Descriptions of Aeolothrips, Melan- 
thrips, and Ankothrips larvae and a 
key to species of Aeolothrips larvae 
were given by Priesner (1926b-1928). 
In 1960 Priesner presented a key to the 
genera of larval Aeolothripidae, includ- 


Fig. 7.—Immature stages of Sericothrips variabilis. 
first-instar larva. c, Early second-instar larva. d, Late second-instar larva. e, Prepupa. 


Intrvois NATuRAL History SurRvEY BULLETIN 


Vol. 31, Art. 5 


ing Franklinothrips and Rhaphidothrips 
in addition to those mentioned above, 
and gave descriptions of the larvae of 
Melanthrips and Rhaphidothrips and 
some larval characters of Aeolothrips 
and Franklinothrips. 


a, Early first-instar larva. 


August, 1974 


Melis (1959) published descriptions 
(in Italian) and illustrations of Aeolo- 
thrips and Melanthrips immatures. 

Material Examined.—INHS: Aeolo- 
thrips bicolor Hinds; 1 larva I; June; 
on grasses; Vermilion County, Illinois. 

A. fasciatus (Linneaus); 5 larvae I, 
2 larvae II; August and September; on 
soybeans; Champaign County, Illinois. 

A. vittipennis Hood; 3 larvae I, 3 lar- 
vae II; June and July; on black locust; 
Johnson and Union counties, Illinois. 


MEROTHRIPIDAE Hood (1914) 


Larva.—Antennae (Fig. 20) each six 
segmented, segments VI and VII usu- 
ally fused; segments not elongate, hav- 
ing faint annular rings and _ lacking 
microtrichia; segment V as long as 
segment IV. 

Head and pronotum (Fig. 40) small 
and tapering anteriorly, posterior and 
anterior tentorial arms joined. Body 
cylindrical. Setae generally long and 
pointed. Abdominal tergite IX (Fig. 
58) with two median pairs of setae 
modified into stout spines. Cuticle 
with very fine microtrichia on abdomen 
and pteronotum, producing a stippled 
pattern. 

Diagnosis.—Merothripid larvae can 
be easily distinguished by the fusion 
of antennal segments VI and VII and 
by the reduction of annular rings and 
absence of the annular microtrichia 
found in other families of the Tere- 
brantia. 

The larvae of the Merothripidae ap- 
pear to be transitional between those 
of the Aeolothripidae and Phlaeothripi- 
dae (suborder Tubulifera). Antennal 
segment V is relatively long in Mero- 
thrips, as in the Aeolothripidae and 
Phlaeothripidae, but it is reduced in the 
Heterothripidae and Thripidae. Mero- 
thripids have the median setae of ab- 
dominal tergite IX modified into spines, 
as in the Aeolothripidae, and a reduced 
number of annular rings and a lack of 
microtrichia on the antennal segments, 
as in the Phlaeothripidae. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


171 


Material Examined.—INHS: Mero- 
thrips morgani Hood; larva Il, 14; 
28 September 1952; on ground cover; 
Key West, Florida. 


HETEROTHRIPIDAE Bagnall (1912) 


Larva.—Antennae (Fig. 9 and 22) 
each seven segmented, segments II-V 
with four or five annular rings, seg- 
ments VI and VII with two or three 
annular rings; microtrichia present on 
most rings; segment V about half the 
length of IV. 


Setae short to long and blunt to 
terminally funneled. Abdominal tergite 
IX (Fig. 59) with two median pairs 
of setae modified into stout spines. 
Cuticle with prominent pustules bear- 
ing fine microtrichia (Fig. 41). 

Larva I with stout spines on abdomi- 
nal segment IX. 

Diagnosis.—Larvae of the Illinois 
Heterothripidae can be distinguished 
by the length of antennal segment V 
and by a combination of many features 
which they share with aeolothripid and 
thripid larvae. Heterothripid larvae ap- 
pear transitional between Aeolothripi- 
dae and Thripidae laryae. Antennal 
segment V is relatively long, annular 
rings are present on segments VI and 
VII, and two pairs of setae on abdomi- 
nal tergite IX are modified into stout 
spines, characteristics also present in 
aeolothripids. The shorter antennal seg- 
ments with fewer annular rings and the 
presence of cuticular pustules are 
thripid characteristics. 

Material Examined.—INHS: Hetero- 
thrips arisaemae Hood; 2 larvae I, 10 
larvae II; May-June; on jack-in-the- 
pulpit (Arisaema sp.); La Salle and 
Carroll counties, Illinois, and Raleigh, 
North Carolina. 


THRIPIDAE Stephens (1829) 


Larva—Antennae each seven seg- 
mented, usually only segments IIT and 
IV have annular rings (also segments 
V-VII in the Heliothripinae); micro- 


172 Inyinois NATURAL History SuRVEY BULLETIN Vol. 31, Art. 5 


trichia often but not always present on than one-fourth the length of segment 
annulations; segment V reduced to less__ IV. 


d 
$ 
a 
oe 
4 
> 
a 

J 


Mer 


Fig. 8-17.—Right antenna (except where indicated) of the first-instar larva. 8.— 
Aeolothrips vittipennis. 9.—Heterothrips arisaemae. 1|0:—Heliothrips haemorrhoidalis, 1].— 
Limothrips denticornis. |12.—Anaphothrips secticornis, 13.—Dendrothrips ornatus. 14.— 
Sericothrips variabilis. 15.—Scirtothrips taxodii. 16.—Echinothrips americanus, left antenna. 
1 7.—Frankliniella tritici. 


August, 1974 


Abdominal tergite IX without spines 
(a posterior comb usually present on 
tergite IX in the Thripini). Setal 
features and cuticular sculpturing vari- 


able. 


Diagnosis.—Larvae of the Thripidae 
can be recognized by the great re- 
duction of antennal segment V and by 
the lack of spines on abdominal tergite 
IX. 

The Thripidae include the  sub- 
families Heliothripinae and Thripinae. 
These subfamilies and their tribes are 
distinguished by antennal features, 
cuticular sculpturing, and other charac- 
teristics. 


Subfamily HELIOTHRIPINAE 
Kamy (1921) 


Larva.—Terminal antennal segment 
greatly elongate (length seven to eight 
times the greatest width, as in Fig. 10, 
25, 26, and 27); antennae with annula- 
tions on segments V—VII, annulations 
often with no or few microtrichia; sense 
cones fairly short. Body often with 
prominent areas of brown coloration. 
Cuticle usually with small to large 
pustules, which generally lack micro- 
trichia. Head usually constricted be- 
hind the eyes (Fig. 43 and 46). Ab- 
dominal segment IX (Fig. 60, 63, and 
65) lacks a posterior comb; segment X 
sometimes with long anal setae. Body 
setae variable, often ornate. 


Diagnosis.—Larvae of the  Helio- 
thripinae are easily recognized by the 
elongate terminal antennal segments 
and by the combination of features 
mentioned above. The Heliothripinae 
may be considered the most primitive 
subfamily in the Thripidae. Primitive 
features are the elongate antennal seg- 
ments, the presence of annulations on 
the terminal segments, and the shorter 
sense cones found also in the Aecolo- 
thripidae. The Heliothripinae resemble 
the Anaphothripini in having cuticular 
pustulation, cheek constrictions, and 
body areas of brown  sclerotization. 
They resemble the Chirothripini in 


VANCE: LARVAE OF THE SERICOTHRIPINI 


173 


having a reduced number of annular 
microtrichia, an enlarged antennal seg- 
ment V (in the first-stage larva of 
Limothrips), and body areas of brown 
sclerotization. 

The only native genus of this sub- 
family in Illinois is Caliothrips; exotic 
genera occurring in greenhouses and 
homes are Heliothrips, Parthenothrips, 
and Hercinothrips. 

Material Examined.—INHS: Calio- 
thrips indicus (Bagnall); 2 larvae I, 1 
larva II; 3 March 1970; reared from 
soybeans; Jabalpur, M.P., India. 

Heliothrips haemorrhoidalis var. an- 
gustior Priesner; 1 larva I, 3 larvae II, 
prepupa (on slide with @ lectotype); 
on plants of virgin forest; Paramaribo, 
Surinam, S.A. 

Hercinothrips femoralis (Reuter); 1 
larva II; 30 April 1953; on African 
violet; St. Louis, Missouri. 

Parthenothrips dracaenae (Heeger); 1 


larva II; March 1952; on Cordyline 
feminalis leaves; Wahiawa, Oahu, 
Hawaii. 


Subfamily THRIPINAE 
Stephens (1829) 


Larva.—Terminal antennal segments 
not greatly elongated (length only two 
to three times the greatest width), 
antennae without annulations on seg- 
ments V—VII, antennal microtrichia and 
sense cones variable. Body coloration, 
cuticular sculpturing, head shape, and 
abdominal segment IX variable. Ab- 
dominal segment X lacks long anal 
setae. 

Diagnosis.—Larvae of the subfamily 
Thripinae can be distinguished from 
those of the Heliothripinae by the 
short, terminal antennal segments and 
by other features not usually occurring 
in the Heliothripinae. 

Priesner (1957) recognized the tribes 
Dendrothripini, Sericothripini, Thripini, 
and Chirothripini in the Thripinae, and 
included the Anaphothripini under the 
Thripini as a subtribe. Stannard (1968 ) 
recognized these tribes, too, but tenta- 


174 TIuurwworis Natura Hisrory SuRvEY BULLETIN Vol. 31, Art. 5 


Fig. 18-27.—Right antenna of the second-instar larva. 18.—Aeolothrips vittipennis. 
19.—Franklinothrips sp. 20.—Merothrips morgani. 2].—Phlaeothripid (Tubulifera), 22.— 
Heterothrips arisaemae. 23.—Chirothrips simplex. 24.—Limothrips cerealium. 25.—Calio- 
thrips indicus. 26.—Heliothrips haemorrhoidalis. 27.—Hercinothrips femoralis. : 


August, 1974 


tively included the Chirothripini under 
the Thripini because of difficulties in 
their categorization. 


Here, five tribes are tentatively rec- 
ognized in the Thripinae: the Chiro- 
thripini, Anaphothripini, Dendrothrip- 
ini, Thripini, and Sericothripini. Cer- 
tain combinations of larval features 
have been found to be characteristic of 
each tribe. Important characters at the 
tribal level include cuticular sculptur- 
ing; features of the antennal annula- 
tions, microtrichia, and sense cones; 
presence or absence of a posterior comb 
on abdominal tergite IX; and to a 
lesser extent the setal types and brown 
sclerotized body areas. 


Tribe CHrRoTHRIPINI Priesner (1949) 


The diagnostic features of Chiro- 
thripine larvae are: (1) cuticle with 
small pustules bearing minute micro- 
trichia, (2) body setae simple with 
certain ones knobbed (Fig. 64) (all 
reduced and pointed in Chirothrips) 
(Fig. 45 and 62), (3) head and 
pronotum often with brown sclerotized 
areas (Fig. 44) (reduced in Chiro- 
thrips) (Fig. 45), (4) antennal micro- 
trichia greatly reduced, (5) antennal 
sense cones short to moderately long 
(Fig. 23 and 24), and (6) eye facets 
small and not bulging at sides of head. 


The Chirothripini larvae resemble 
the Heliothripinae larvae in the reduc- 
tion of the annular microtrichia on the 
antennae, segment V in the first-stage 
larva being longer and having two an- 
nulations (Fig. 11) (a trait found in 
most Heliothripinae but in no other 
Thripinae). The brown sclerotized 
areas of Limothrips resemble those 
found in many Heliothripinae. 


Two genera in this tribe occur in 
Illinois, Chirothrips and Limothrips. 
Both contain species that are grain 
feeders and can be serious pests. 


Material Examined.—INHS: Limo- 
thrips cerealium (Haliday); 1 larva 
II; 24 June 1953; Kenney, Illinois. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


175 


USNM: Chirothrips simplex Hood; 
10 larvae II, 1 prepupa; 21 October 
1961; reared from Bouteloua eriopoda; 
Las Cruces, New Mexico. 


Limothrips denticornis Haliday; 2 
larvae I, 3 larvae II, 2 prepupae; 10 
July 1959; on barley; Northwood, 
North Dakota. 


Tribe ANAPHOTHRIPINI Priesner (1949) 


The diagnostic features of anapho- 
thripine larvae are: (1) cuticle cov- 
ered with large pustules and usually 
lacking microtrichia; (2) dorsal body 
setae pointed, knobbed, or blunt; (3) 
brown sclerotized body areas present 
in some species (Fig. 47 and 48); (4) 
posterior comb on abdominal tergite 
IX usually lacking (Fig. 66 and 67) 
(sometimes present in Anaphothrips ); 
(5) annular microtrichia on antennae 
generally reduced (Fig. 12, 28, and 
29); and (6) antennal sense cones 
moderately long to long. 


The status of the Anaphothripini has 
long been variously interpreted. Pries- 
ner (1957) included the members of 
this tribe within the Thripini. Gentile 
& Bailey (1968) considered the Ana- 
phothripini to be the most primitive 
of all thripine tribes, and Stannard 
(personal communication) is of the 
opinion that the Anaphothripini are 
close to the Heliothripinae. 


The larvae of the Anaphothripini re- 
semble those of the Heliothripinae in 
cuticular sculpturing, the presence of 
brown coloration in some species, a 
reduction of antennal microtrichia, and 
other features. They resemble the 
Thripini larvae in cuticular sculpturing 
and in the posterior comb that is some- 
times present in Anaphothrips. Ana- 
phothrips secticornis has brown sclero- 
tized areas on the pteronotum similar 
to those in the annulipes group of Seri- 
cothrips. Chaetanaphothrips possesses 
expanded setae similar to those of 
Sericothrips. 


In Illinois the tribe Anaphothripini 


176 


is represented by Anaphothrips, Ap- 
tinothrips, Bregmatothrips, Chaetana- 
phothrips, Chilothrips, Oxythrips, and 
Prosopothrips. 


Material Examined.—INHS: Ana- 
phothrips secticornis Karny; 3 larvae 
I, 6 larvae II; 21 January 1964; on 
short grasses; Barff Peninsula, Sérling 
Valley, South Georgia Island. 


Aptinothrips rufus (Gmelin); 8 larvae 
I, 4 larvae II; 23 June 1933; on timothy 
heads; Champaign County, Illinois. 


Oxythrips cannabensis Knechtel; 13 
larvae I, 20 larvae II; August; on mari- 
juana; Henry and Morgan counties, 
Illinois. 


Chilothrips pini Hood; 5 larvae II; 
on cottonwood; 15 October 1959; Park- 
land, Adams County, Wisconsin. 


Chilothrips sp.; 4 larvae II; on rot- 
ten wood and pigmy cypress duff; 
Deschutes County, Oregon, and Men- 
docino County, California. 


Tribe DENDROTHRIPINI 
Priesner (1926b-1928 ) 


The diagnostic features of dendro- 
thripine larvae are: (1) cuticle covered 
with minute microtrichia, resulting in 
a stippled pattern and forming larger 
pustules in transverse rows on the ter- 
minal abdominal segments (Fig. 71); 
(2) body setae generally simple but 
with certain ones terminally knobbed 
(Fig. 74); (3) brown sclerotized areas 
lacking (Fig. 51); (4) antennal sense 
cones long (Fig. 35 and 36); (5) an- 
tennal microtrichia prominent and lo- 
cated between the annulations on seg- 
ment IV (larva I) (Fig. 13), as in the 
Sericothripini. 


Larval characters of the Dendro- 
thripini are well defined and easily 
delineated; the larvae are very similar 
to those of the Sericothripini. Cuticular 
sculpturing is similar to that of the 
Sericothripina, and setae are similar 
to those of the Scirtothripina. Random 
microtrichia on antennal segment IV 


Intivois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 5 


are also diagnostic for larvae of the 
Dendrothripini and Sericothripini. 

The tribe Dendrothripini in Illinois 
includes one native genus, Leucothrips, 
and two genera introduced from 
Europe and Japan, Dendrothrips and 
Pseudodendrothrips, respectively. 

Material Examined.—INHS: Dendro- 
thrips ornatus (Jablonowski); 5 larvae 
I, 3 larvae II; 23 August 1955; on 
privet; Champaign County, Illinois. 

Leucothrips piercei 
larva I, 3 larvae II; 20 June 1967; on 
redbud leaves; Montgomery County, 
Illinois. 


Pseudodendrothrips mori (Niwa); 1 
larva I, 4 larvae II; 25 October 1961; 
on Japanese mulberry leaves; McLean 
County, Illinois. 


Tribe Turipmi Stephens (1829) 


The diagnostic features of thripine 
larvae are: (1) cuticle covered with 
small to large pustules (Fig. 55), often 
with microtrichia present; (2) dorsal 
body setae pointed (Fig. 56), knobbed, 
blunt, or terminally funneled; (3) 
brown sclerotized body areas usually 
absent; (4) posterior comb or teeth 
usually present on abdominal tergite 
IX except in Scolothrips (Fig. 75, 76, 
and 77); (5) antennal annular micro- 
trichia not reduced; and (6) antennal 
sense cones short to moderately long 
(Fig. 17, 37, and 38). 

The Thripini larvae resemble the 
Anaphothripini larvae in cuticular 
sculpturing and general body appear- 
ance. Generally, less diversification is 
found among closely related members 
of the Thripini than is usual among 
the members of other tribes. 


The tribe Thripini in Illinois includes 


Baliothrips, Ctenothrips, Dorcadothrips, 


Frankliniella, Iridothrips, Microcephalo- 
thrips, Odonotothrips, Plesiothrips, 
Rhaphidothrips, Scolothrips, Taenio- 
thrips, and Thrips. 

Material Examined.—INHS: Cteno- 
thrips bridwelli Franklin; 1 larva II; 11 


(Morgan); 1° 


; 
| 
i 
{ 
q 
i 


August, 1974 VaANcE: LARVAE OF THE SERICOTHRIPINI 177 


Fig. 28-38.—Right antenna (except where indicated) of the second-instar larva. 28.— 
Anaphothrips secticornis. 29.—Oxythrips cannabensis. 30,—chinothrips americanus. 31].— 
Sericothrips annulipes. 32.—Sericothrips variabilis. 33.—Scirtothrips taxodii. 34.—Drepano- 
thrips reuteri. 35.—Dendrothrips ornatus. 36.—Pseudodendrothrips mori, left antenna. 37.— 
Ctenothrips bridwelli. 38.—Taeniothrips simplex. 


178 


July 1947; on Arisaema dracontium; 
La Salle County, Illinois. 

Frankliniella fusca (Hinds); 3 larvae 
II; 9 June 1949; Berlese collecting 
method; Mercer County, Illinois. 

F. parvula Hood; 1 larva II; 20 June 
1970; on bananas; Ciudad Chontalpa, 
Tabasco, Mexico. 

F. tritici (Fitch); 1 larva I, 9 larvae 
II; on flowers of yarrow and Culver’s 
root flowers; Lake, Livingston, and 
Massac counties, Illinois. 

Frankliniella sp.; 12 larvae II, 3 pre- 
pupae, 2 pupae; May-July; on oats and 
from Berlese collecting method; Jack- 
son County, Illinois, and Friday Har- 
bor, Washington. 

Microcephalothrips sp.; 5 larvae II; 
16 December 1949; on Spanish moss; 
Chiefland, Florida. 

Scolothrips pallidus (Beach); 1 larva 
II; 28 July 1964; on cotton; Kewanee, 
Missouri. 

Taeniothrips simplex (Morison); 10 
larvae II, 7 prepupae, 11 pupae; July— 
August; on gladiolus; Champaign and 
Will counties, Illinois. 

Thrips impar Hood; 1 larva I, 5 
larvae II; 16 July 1969; on jewelweed; 
Edward, Henry, and McLean counties, 
Illinois. é 

Thrips physapus Linneaus; 1 larva 
II; December 1959; Recoaro, Italy. 

Thrips tabaci Lindeman; 1 larva I, 
1 larva II; 25 April 1968; on clover; 
Champaign County, Illinois. 


Tribe SERICOTHRIPINI 
Priesner (1926b-1928 ) 


The diagnostic features of serico- 
thripine larvae are: (1) cuticle covered 
with fine microtrichia, resulting in a 
stippled pattern over the abdomen and 
pteronotum; (2) cuticular pustules ab- 
sent or very reduced; (3) all or some 
dorsal body setae expanded and/or 
fimbriate; (4) brown sclerotization lack- 
ing; (5) fourth antennal segment of 
first-instar larva densely covered with 
microtrichia. 


Tttinois NATURAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


The tribe Sericothripini is divided 
into the subtribes Sericothripina, Scirto- 
thripina, and a new subtribe, the 
Echinothripina. Each of these groups 
is distinctive in certain larval charac- 
teristics, and each shows certain simi- 
larities with other tribes, indicating 
possible lines of relationship. 

Sericothrips, particularly the an- 
nulipes group, resembles the Anapho- 
thripini in such genera as Anaphothrips 
and Chaetanaphothrips. Most mem- ~ 
bers of the annulipes group possess 
pteronotal sclerotized areas similar to 
those found in Anaphothrips  secti- 
cornis. Sericothrips cingulatus has 
darker brown markings and small 
pustules reminiscent of those in the 
Anaphothripini and Chirothripini. 
Chaetanaphothrips has fanned (but not 
fimbriate) setae similar to those in 
Sericothrips. 


Scirtothrips and Drepanothrips show 
similarities to the Dendrothripini in 
setal features, coloration, and cuticular 
sculpturing. 


Subtribe SrRicoTHRIPINA 
Priesner (1957) 


Larvae of the Sericothripina are 
characterized by fan-shaped, fimbriate 
setae; minute cuticular microtrichia set 
on pustulelike bases on the terminal 
abdominal segments; and the absence 
of pustules elsewhere. An exception is 
Sericothrips cingulatus, in which all 
microtrichia are set on small pustules. 
Microtrichia in all Sericothrips species 
form transverse rows on the abdomen, 
especially on the terminal abdominal 
segments, similar to the rowed ordering 
of the larger pustules in other groups. 

Setal form in this group is unique 
among all thrips larvae and is an 
easily recognized diagnostic character. 
Caliothrips and possibly Chaetanapho- 
thrips have fan-shaped setae somewhat 
like those of the Sericothripina, but the 
setae of Caliothrips and Chaetanapho- 
thrips are smooth rather than fimbriate. 
The only genera in the Sericothripina 


August, 1974 


in Illinois are Sericothrips Haliday and 
Zonothrips Priesner. 


Subtribe ScmrroTHRIPINA 
Priesner (1957) 


Larvae of the Scirtothripina are char- 
acterized by setae expanded or fun- 
neled terminally only and by long, 
dense cuticular microtrichia in the ab- 
sence of cuticular pustules. Larvae of 
this subtribe are smaller than those of 
the Sericothripina and tend to have 
less ornamentation and less interspe- 
cific variation. Larval Scirtothripina 
have no hypodermal pigmentation, 
brown sclerotized areas, or setal basal 
rings; the setae are much simpler, and 
no cuticular pustulation is evident ex- 
cept for transverse rows formed by 
stippling on abdominal segments IX 
and X. Scirtothripina larvae resemble 
those of the Dendrothripini in their 
setae, both having combinations of 
long, terminally-funneled and_ short, 
pointed setae. Larvae of the Scirto- 
thripina can be easily identified (par- 
ticularly Scirtothrips) by their dense, 
long cuticular microtrichia and_ their 
lack of cuticular pustules. The genera 
included in this subtribe, according to 
Priesner (1957), are Charassothrips 
Hood, Drepanothrips Uzel, Enneo- 
thrips Hood, Ensiferothrips Bianche, 
Octothrips Moulton, Scirtodothrips 
Hood, Scirtothrips Shull, and Sericopso- 
thrips Hood. 


Subtribe EcHINOTHRIPINA, 
New Subtribe 


The proper placement of the genus 
Echinothrips in higher categories has 
long been in question. Moulton (1911) 
placed Echinothrips in the Helio- 
thripinae, and Medina as late as 1961 
still considered this to be the best 
placement. Priesner (1957), however, 
considered this genus to be in the 
Thripini because of imaginal endo- 
thoracic morphology, and Stannard 
(1968) transferred Echinothrips into 


VANCE: LARVAE OF THE SERICOTHRIPINI 


179 


the Sericothripini because of the pres- 
ence of abdominal microtrichia and the 
lack of fusion of the fore vein to the 
costa in the fore wing of the adults. 


The larval characters of Echinothrips 
support Stannard’s placement of the 
genus. Similarities of Echinothrips 
larvae to the larvae of other Serico- 
thripini genera can be seen in cuticular 
microtrichia, fimbriate setae, antennal 
shape and sense cones, and extra micro- 
trichia on antennal segment IV of first- 
stage larvae. Echinothrips differs in its 
unexpanded and more elongate body 
setae, larger and more elongate body 
size, positioning of head setae H1, and 
loss of pronotal setae P3. 


Generally, Echinothrips most closely 
resembles the Sericothripina, but be- 
cause of the differences described, the 
genus has been placed in its own sub- 
tribe. Wilson (1971) delimits a group 
of closely related genera that he calls 
the Echinothrips complex, including 
Cercyothrips Morgan, Echinothrips 
Moulton, Enneothrips Hood, Plesiopso- 
thrips Hood, Plesiothrips Hood, and 
Pteridothrips Priesner. Some of these 
he placed with the Thripini and others 
with the Sericothripini; Wilson feels 
that this group is transitional between 
the Sericothripini and the Thripini and 
that possibly it merits tribal status. The 
only genus included here in this sub- 
tribe is Echinothrips Moulton. 


Drepanothrips Uzel (1895) 


Larva Il—Body color yellow. An- 
tennal segments, tibiae, bases of femora, 
setae, and setal basal rings light brown 
to brown. Apices of antennal segments 
I and II and base and apex of segment 
III pale gray. Eyes red. 


Antennae each seven segmented 
(Fig. 34); longer sense cone on seg- 
ment IV, and sense cones on segments 
V and VI moderately long and slightly 
blunted; all of equal length. Segment 
II with a pair of terminally funneled 
setae; segment III with six annulations, 


180 


the distal three with short micro- 
trichia; segment IV with five annula- 
tions, all with longer microtrichia. 


Head (Fig. 52) longer than wide. 
Eyes with four large facets bulging at 
sides of head. Mouth cone moderately 
blunt. Head with four pairs of dorsal 
setae; H1, H3, and H4 subequal and 
terminally funneled. 


Pronotum longer than wide with 
seven pairs of terminally funneled 
setae; P6 and P7 somewhat longer 
than P1—-P5. Mesonotum with seven 
pairs and metanotum with five pairs 
of funneled setae, all of nearly equal 
length. Abdominal tergite I with two 
pairs and tergites II-VIII each with 
three pairs of funneled setae; Al, A2, 
and sometimes A3 on tergite IX (Fig. 
70) funneled, and all subequal to equal 
in length. Abdominal tergite X with 
three pairs of setae, Al funneled. 


Almost all dorsal body setae termi- 
nally funneled and of moderate length. 
Bases of setae with faint brown rings. 
Abdominal and pteronotal cuticle with 
dense stippling and fine microtrichia, 
which are shorter and less obvious than 
those on Scirtothrips and longer than 
those on Sericothrips, stippling forming 
transverse rows on abdominal segments 
IX and X. Segment IX lacking a 
posterior comb. 

Diagnosis—Drepanothrips larvae 
most closely resemble Scirtothrips 
larvae from which they can be dis- 
tinguished by the dorsal body setae, 
all of which are terminally funneled, 
while only a few characteristic ones 
are funneled in Scirtothrips. The cu- 
ticular microsetae are shorter and less 
dense in Drepanothrips than they are 
in Scirtothrips. Larvae of Drepano- 
thrips differ from those of other Serico- 
thripini in having terminally expanded 
setae, the setae of the other genera being 
either totally expanded or long and un- 
expanded. Dendrothrips and Pseudo- 
dendrothrips, which resemble Drepano- 
thrips in the larval stages, can be dif- 
ferentiated by their lack of cuticular 


Inuinois NATURAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


microtrichia and by their having only 
certain dorsal body setae funneled. 


(ttn et A al 


The genus contains only one species, 


D. reuteri, in Illinois. 


Drepanothrips reuteri Uzel (1895) 
(Fig. 34, 52, and 70) 
Larva II.—Body light yellow to yel- 


low. Antennae, tibiae, bases of femora, 
and setal basal rings light brown; an- 


tennal segments II, V—VIII, and apex. 


of IV often darker brown. Apices of 
antennal segments I and II and base 
and apex of segment III very pale gray. 
Eyes dark red. 

Most body setae slender, terminally 
funneled or dilated, and subequal in 
length (14-19 pm). Bases of setae 
with faint unraised brown rings. Dorsal 
sclerotized areas lacking. 
forming transverse rows only on ab- 
dominal segments IX and X. 

Measurements of the D. reuteri larva 
II are shown in Table 6. 

Diagnosis.—D. reuteri occurs on 
grapevines (Vitis sp.), of which it 


Table 6.—Measurements, in microns, of 
three Drepanothrips reuteri larvae I. 


Length Width 

Chatacies Range Mean Range 
Antennal segment 

I 19° 22-23 

II 28-31 22-23 

III 42-46 22 

IV 42-46 17-20 

V 8-11 12-14 

VI 8-9 ote 9 

VIL = 16 ae 6 
Antenna 163-178 171 Box 
Head 70-78 78-85 
Pronotum 93-124 124-140 
Body 660-825 765 
Setae 

Hil 16 

H4 16-19 

P7 16-19 

A(IX)1 14-16 

A(IX)2 17-19 

Ventral setae 

(TX) 15 


Stippling © 


y 


“A single measurement in a range column ‘ 
indicates that all such measurements were — 


identical. 


August, 1974 VANCE: LARVAE OF THE SERICOTHRIPINI 181 


has been reported to be a pest. Bailey ture concerning it. This species has 
(1942) gave an account of the biology _ been recorded only once in Illinois, two 
of this thrips and discussed the litera- adult females having been taken in 


Fig. 39-48.—Head and pronotum (except where indicated) of the second-instar larva. 
39.—Aeolothrips vittipennis. 40.—Merothrips morgani. 4].—Heterothrips arisaemae. 42.— 
Phlaeothripid (Tubulifera). 43.—Heliothrips haemorrhoidalis. 44.—Limothrips cerealium, 
head, pronotum, and left foreleg. 45.—Chirothrips simplex, head, pronotum, and left foreleg. 
46.—Caliothrips indicus. 47.—Oxythrips cannabensis. 48.—Anaphothrips secticornis. 


182 


Urbana from a sparrow nest built in 
a grape arbor. 


Material Examined.—INHS: 3 larvae 
II; 23 August 1965; on grape; col- 
lected by K. Stahlik; Selma, Fresno 
County, California. 


Echinothrips Moulton (1911) 


Larva II.—Cuticular color usually 
yellow to orange. Antennal segments, 
tibiae, bases of femora, and setae gen- 
erally light brown. Eyes red. 


Antennae each seven segmented 
(Fig. 30); longest sense cone on seg- 
ment IV, sense cones on V and VI long 
and pointed; segments II and III each 
with two pairs of fimbriate setae; seg- 
ment III with five annulations, none 
with microtrichia; segment IV with 
five annulations, all with microtrichia. 


Head (Fig. 54) wider than long. 
Eyes with four large facets bulging at 
sides of head. Mouth cone moderately 
blunt. Head with four pairs of dorsal 
setae, all long and fimbriate; H1 lo- 
cated more posteriorly than usual in 
most known thrips larvae and almost 
opposite to H4; H3 and H4 equal and 
shorter than H1. 


Pronotum (Fig. 54) wider than long 
with six pairs of fimbriate setae; P3 
lacking; Pl, 2, 4, and 6 all longer 
than P7. Mesonotum with seven pairs 
and metanotum with four pairs of long 
fimbriate setae of varying lengths. 
Setae Al and 2 of abdominal segment 
IX long; A3 of varying length, some- 
times reduced (Fig. 73). Segment X 
with three pairs of dorsal setae; Al 
and A3 long and fimbriate. 


Most dorsal body setae fimbriate and 
long, the setal lengths on abdominal 
tergites sometimes varying greatly. Cu- 
ticle with minute microtrichia sparsely 
scattered on abdominal tergites, micro- 
trichia becoming pustulelike and form- 
ing transverse rows on the terminal 
abdominal segments. Abdominal seg- 
ment IX lacking a posterior comb. 


Intivois NATuRAL History SURVEY BULLETIN 


Vol. 31, Art. 5 


Larva I.—Cuticle yellow to orange; 
hypodermal pigment lacking. Anten- 
nal segments I, III, and most of IV, 
tibiae, and bases of femora generally 
light brown; segment II, apex of IV, 
and all of V—VII darker brown. Eyes 
red. 


Antennae each seven segmented; su- 
ture between IV and V usually distinct. 
Sense cones on segments IV—VI as in 
larva II, but longer (Fig. 16). Seg- 


ments II and III each with a pair of’ 


long fimbriate setae. Segment III with 
five annulations, with minute micro- 
trichia present ventrally. Segment IV 
with six annulations; microtrichia pres- 
ent randomly on and between annula- 
tions but less dense than in Scirto- 


ae ean aes E- Se Vico alll 


thrips. Apical segment not narrowed ~ 


terminally. 


Chaetotaxy similar to that of larva 
II, except posteroangular setae (P7) 
lacking, the mesonotum with four pairs 
of setae, the metanotum with three 


pairs of setae, and abdominal segment _ 


IX with two pairs of dorsal setae. 


Integument with stippling, as in larva 


II, but fainter. 


Diagnosis.—Echinothrips larvae can — 
be easily recognized by their long fm- 


briate setae and elongate body shape, 
which are unique among the Thripinae 
in Illinois. 
sculpturing are similar to those of 
Sericothrips, but setal length and type, 


body shape, placement of setae Hl ~ 


(more posterior in Echinothrips), and 
loss of-one pronotal setal pair (P3) 
differ from those of Sericothrips. These 


Antennal and cuticular 


" Nee 


features distinguish Echinothrips from _ 


all other genera. 
thripine genus in Illinois possessing 
long setae similar to those in Echino- 
thrips is Scolothrips. 


Interspecific variation in Echinothrips 


is very limited in the two species con- — 


sidered, E. americanus and E. sub- 
flavus: In the one slide of E. subflavus 
studied, body dimensions and setal 


- 
- 


The only other © 


: 
4 
yh, 

nid 

oye 

7 


lengths were larger than those in ~ 


&, | 


August, 1974 VANCE: LARVAE OF THE SERICOTHRIPINI 183 


E. americanus. However, considerable Echinothrips americanus 
variation in setal lengths was found in Morgan (1913) 
americanus; so the extent of variation . 

in both species will have to be investi- (Fig. 16, 30, 54, and 73) 

gated before setal measurements can Larva II.—Cuticle yellow to orange. 
be used as a diagnostic feature. Antennae, tibiae, bases of femora, and 


Fig. 49-56.—Head and pronotum of the second-instar larva. 49.—Scirtothrips niveus. 

50.—Sceirtothrips taxodii. 5].—Dendrothrips ornatus. 52.—Drepanothrips reuteri, 53.— 

Sericothrips annulipes. 54.—Echinothrips americanus. 55.—Taeniothrips simplex. 56.— 
Scolothrips pallidus. 


184 


setae brown; apices of antennal seg- 
ments I and II and base and apex of 
III light brown. Eyes red. 


Most dorsal setae long (50-70 pm) 
and fimbriate for most of their length; 
others (H2; Msl, 2, 5, and 6; and Mt1) 
shorter (20-30 pm). Light brown spot 
present on head anteriorly. Cuticle with 
large stippling on abdomen forming 
transverse rows and becoming finer 
and randomly distributed on pterono- 
tum and posterior portion of pronotum. 


Measurements of the E. americanus 
larva II are shown in Table 7. 


Table 7.—Measurements, in microns, of 
five Echinothrips americanus larvae II. 


Y Length Range 

EES Range Mean Width 
Antennal segment 

I 19-23 28-31 

II 28-36 25-29 

Ill 50-62 23-28 

IV 43-54 19-23 

V 11-12 16-19 

VI 12-16 eae 12-16 

VII 26-28 tren 8 
Antenna 195-217 209 sie 
Head 85-93 96-116 
Pronotum 105-124 148-178 
Body 1,065-1,281 1,167 
Setae 

H1 54 

H4 ~ 39-46 

P7 39-46 

A(IX)1 54-70 

A(IX)2 67-78 

A(IX)3 51-54 

A(X)1 54-67 


_ *A single measurement in a range column 
indicates that all such measurements were 
identical. 


Larva I—Cuticle yellow to orange. 
Antennal segments I, III, and most of 
IV brown; apices of I and II and base 
and apex of III pale brown; segments 
II, apex of IV, and V-VII darker 
brown. Tibiae, bases of femora, and 
setae light brown. Eyes red. 

Setae long and fimbriate as in larva 
II, but generally shorter (30-45 pm 
in larva I). 

Cuticle with stippling visible only on 
abdominal segment X. 


Ituinois NAturRAL History SURVEY BULLETIN 


_ by their somewhat smaller size. E. 


Vol. 31, Art. 5 


Measurements of the E. americanus 
larva I are shown in Table 8. 


Table 8.—Measurements, in microns, of — 
two Echinothrips americanus larvae |. 


Length Width 

Character Range Range 
Antennal segment 

I 16-19 23-26 

II 26-28 23° 

III 39-40 23 

IV 43-46 23 

Vv 6-8 12 

VI 12-16 9 

VII 26 8 
Antenna 168-183 A 
Head 70 88 
Pronotum 85 116 
Body 807-814 
Setae 

H1 32-40 

A(IX)1 50-51 

A(IX)2 40-43 

A(X)1 132 


» A single measurement in a range column 
indicates that all such measurements were © 
identical. 


Diagnosis ——E. americanus larvae — 
cannot at this time be distinguished 
from those of subflavus except by as- 
sociated adults and host-plant data and 


americanus is found on many forest 
plants, in particular on jewelweed 
(Impatiens). B 
Material Examined.—INHS: 2 larvae 
I, 10 larvae II; June—October; on 
Desmodium, hydrangea, and jewelweed — 
foliage; Gallatin, Clark, Henry, and ~ 
Johnson counties, Illinois. 


-Echinothrips subflavus 
“Hood (1927b) 


Larva I1.—Cuticle yellow. Antennal 
segments, tibiae, bases of femora, and _ 
setae brown; apices of antennal seg- 
ments I and II and base and apex of- 
III light brown. Eyes dark red. 

Most dorsal setae long (60-80 pm) — 
and fimbriate for most of their length; 
others (H2; P7; Msl, 2, 5, and 6; 
and Mtl) shorter (25-40 pm). Cuticle 
with larger stippling on abdomen, and _ 


August, 1974 


forming transverse rows and becoming 
finer and randomly distributed on 
pteronotum and posterior portion of 
pronotum. 

Measurements of the E. subflavus 
larva II are shown in Table 9. 


Table 9.—Measurements, in microns, of one 
Echinothrips subflavus larva |. 


Character Length Width 
Antennal segment 

I 23 31 

II 42 28 

Ill 65 25 

IV 57 23 

Vv 11 16 

VI 16 12 

VII 28 8 
Antenna 242 sigs 
Head 116 124 
Pronotum 124 194 
Body 1,350 ee 
Setae 

H1 62 

H4 46 

P7 28 

A(IX)1 85 

A(IX)2 85 

A(IX)3 60 

A(X)1 85 


Diagnosis.—E. subflavus larvae can- 
not at this time be distinguished from 
those of E. americanus except by as- 
sociated adults and host-plant data. 
E. subflavus is found on hemlock 
(Tsuga canadensis (L.) ) in the eastern 
United States and could possibly be 
brought into Illinois on hemlocks in- 
tended for ornamental plantings. 


Material Examined.—USNM: 1 larva 
II; 23 July 1939; on hemlock; col- 
lected by J. D. Hood; Oswegatchie, 
New York. 


Scirtothrips Shull (1909) 


Larva I1.—Cuticular color yellow to 
orange, sometimes with orange hypo- 
dermal subintegumental pigment. An- 
tennae, bases of femora and entire 
tibiae, and an anterior median cephalic 
spot all light brown to brown. Apices 
of antennal segments I and II and base 
and apex of III pale gray. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


185 


Antennae each seven segmented ( Fig. 
33). Longest sense cone on segment IV 
and sense cones on segments V and VI 
long, pointed, and all subequal. Seg- 
ment II with a pair of funneled setae. 
Segment III with six amnulations, fine 
microtrichia present on all annulations. 


Head (Fig. 49) wider than long. 
Eyes with four large facets bulging 
at sides of head. Mouth cone mod- 
erately blunt. Head with four pairs 
of dorsal setae; H1 usually and H4 
sometimes funneled; H1 and H4 of 
equal length; H3 subequal to H1 and 
H4. 


Pronotum (Fig. 49) wider than long 
with seven pairs of dorsal setae. All 
except the posteroangular setae (P7) 
short and pointed; P7 longer and often 
funneled. Mesonotum with seven pairs 
and metanotum with four pairs of short, 
pointed dorsal setae. Abdominal seg- 
ment I with two pairs and segments 
II-VIII each with three pairs of dorsal 
setae. Segment IX (Fig. 72) with four 
pairs of dorsal and lateral setae; A2 
longest and sometimes funneled; ven- 
tral setae about as long as Al. Segment 
X with two pairs of dorsal setae; Al 
sometimes funneled. 


Most setae pointed and fairly short; 
H1 and P7 always, and H4, A(IX)2, 
and A(X)1 sometimes longer and ter- 
minally funneled or knobbed. Integu- 
ment with dense microtrichia, resulting 
in a dense stippling effect over the 
abdomen and pteronotum and forming 
transverse rows on abdominal segments 
IX and X. Abdominal segment IX lack- 
ing a posterior comb. 


Larva I.—Cuticular color light yel- 
low to orange. Antennal segments, 
tibiae, and bases of femora light brown 
to brown. Apices of antennal segments 
I and II and base and apex of III 
pale gray. Terminal antennal segments 
darker brown. 


Antennae each seven segmented 
(Fig. 15), the suture between IV and 
V sometimes indistinct. Sense cones 
as in larva II; segment II with pair 


186 ILtivoris NaTurRAL History Survey BuLLerin Vol. 31, Art.5 


of terminally funneled setae; segment some microtrichia scattered between 
III with six annulations with fine micro- the annulations; segment iv with five 
trichia present on the distal four and annulations, with longer microtrichia 


a ance? 
a 
ye iprreer fs 


= 


‘ a 7, 
ony ay 
1 My 

arene IDyyralsy 


OO 
hye Uy 5 
rary) 000) sal 
uw 39) yy 
oh at oui 
OA Sty Abe DD) I 
as oe ze 


9000 S0g0035 ~ 
CECE N "G0 0.00 2295 
code °0000))0, 
@ip 2.000 ole. ee 


Fig. 57-67.—Abdominal segments VIII-X of the second-instar larva. 57.—Aeolothrips 
vittipennis, 58.—Merothrips morgani. 59.—Heterothrips arisaemae. 60.—Caliothrips indicus. 
61.—Phlaeothripid (Tubulifera) . 62.—Chirothrips simplex. 63.—Heliothrips haemorrhoidalis. 
64.—Limothrips cerealium. 65.—Hercinothrips femoralis, 66.—Oxythrips cannabensis. 67. 
—Anaphothrips secticornis. 


August, 1974 


present on and randomly between the 
annulations; segment VII tapering 
apically. 


Chaetotaxy similar to that of larva 
II, except the posteroangular setae lack- 
ing, mesonotum with four pairs and 
metanotum with three pairs of setae, 
and abdominal segment IX with two 
dorsal pairs and one lateral pair of 
setae. Integument with microtrichia, 
and resultant stippled pattern fainter 
than in larva II. 


Diagnosis.—Scirtothrips larvae can 
be distinguished from the larvae of 
other sericothripines by their long 
dense cuticular microtrichia and by 
their simple reduced dorsal body setae, 
only a few of which are long and ter- 
minally funneled. Drepanothrips and 
Sericothrips have less dense and shorter 
cuticular microtrichia and all setae 
either terminally funneled or fanned. 
Scirtothrips resembles Dendrothrips in 
having small simple setae with only 
certain ones longer and terminally fun- 
neled, but larvae of the latter genus 
lack the dense cuticular microtrichia 
of Scirtothrips. 


Morphological characters used to 
separate the larvae of Scirtothrips are 
uncertain. The extent of intraspecific 
variation in the species is not known 
due to a lack of specimens. Host-plant 
data and associated adults should be 
used where possible to supplement 
larval identifications. 


KEY TO THE MATURE LARVAE II 
OF SCIRTOTHRIPS 


1. Dorsal setae H1, H4, P7, A(TX) 2, and 
A(X)1 all terminally funneled; Hi 
and H4 both longer, about 23 wm 
(Fig. 49); body color yellow; found 
TTS COL ODEs eos ctste sever ta texalaceaie-2. niveus 


At most, only dorsal setae Hi and P7 
funneled; H1 and H4 shorter, about 
15 um (Fig. 50); body color yellow 
or orange; found on red cedar or 
OHLCYMECHS is aot oils nie ie os ae eniadele 2 


2. Body color usually orange; setae 
A(IX)1 and 3 significantly shorter 
than A(IX)2; found on red cedar 
BF Oke hye nd CE tere brevipennis 


VANCE: LARVAE OF THE SERICOTHRIPINI 


187 


Body color yellow, sometimes with 
orange subintegumental pigment; 
setae A(IX)1 and 3 long (19 and 
23-25 um) and subequal to A(IX)2 
(Fig. 72); found on cypress...taxodii 


Scirtothrips brevipennis 
Hood (1914) 


Larva II.—Body pale orange to yel- 
low-orange, sometimes with darker 
orange pigmentation. Antennal seg- 
ments I-IV brown; apices of segments 
I, II, and III pale gray; segments 
V-VII darker brown. Tibiae, bases of 
femora, and anterior median cephalic 
spot brown. Eyes red. 


Most setae simple, pointed, and short; 
Hl and P7 longer and funneled; 
A(IX)2 pointed and decidedly longer 
than A(IX)1 and A(IX)3. Cuticle 
with fine dense microtrichia. 


Measurements of the S. brevipennis 
larva II are shown in Table 10. 


Table 10—NMeasurements, in microns, of 
two Scirtothrips brevipennis larvae ||. 


Length Width 

Character Range Range 
Antennal segment 

I if: 23 

II 31 23 

Ill 46 22 

IV 46 22 

Vv 8 14 

VI 8 12 

VII 15 6 
Antenna 170 ore 
Head 93-108 78 
Pronotum 90-101 140-155 
Body 631 
Setae 

H1 16 

H4 16 

P7 16-17 

A(IX)1 14-16 

A(IX)2 25-26 

A(IX)3 16 


aA single measurement in a range column 
indicates that all such measurements were 
identical, 

Diagnosis.—S. brevipennis larvae are 
distinguished from those of niveus in 
having shorter cephalic setae (H1 and 
H4) and having H4, A(IX)2, and 


188 


A(X)1 pointed instead of terminally 
funneled. S. brevipennis larvae are 
separated from those of taxodii by hav- 
ing orange body color (usually yellow 
in taxodii) and shorter setae (Al and 
A3) on abdominal segment IX than 
taxodii larvae have. S. brevipennis is 
found on eastern red cedar (Juniperus 
virginiana). 

Material Examined.—INHS: 2 larvae 
II; June; on red cedar; Johnson and 
Pope counties, Illinois. 


Scirtothrips niveus 
Hood (1913) 
(Fig. 49) 


Larva II.—Body yellow with darker 
yellow pigmentation. Antennae, tibiae 
and bases of femora light brown. Apices 
of antennal segments I and II and base 
and apex of III pale gray. Eyes red. 

Most setae simple, fairly short (12 
pm), and pointed; H1, H4, P7, A(1X)2, 
and A(X)1 all longer and funneled. 
Cuticle with fine dense microtrichia. 

Measurements of the S. niveus larva 
II are shown in Table 11. 

Diagnosis.—S. niveus larvae can be 
distinguished from other Scirtothrips 


Table 11.—Measurements, in microns, of 
one Scirtothrips niveus larva II. 


Character Length Width 
Antennal segment 

I 16 25 

II 29 22 

Til 46 22 

IV 43 22 

Vv 11 16 

VI 11 11 

VII 16 8 
Antenna 172 stots 
Head 93 93 
Pronotum 108 163 
Body 840 ; 
Setae 

H1 23 

H4 23 

Pi 23 

A(IX)1 20 

A(IX)2 28 

A(TX)3 23 

A(X)1 14 


Inuinois NaturAL History SuRvEY BULLETIN 


Vol. 31, Art. 5 


larvae considered here by the longer 
H1 and Hé4 setae and by knobbed 
rather than pointed H4, A(IX)2, and 
A(X)1 setae. S. niveus occurs on 
leaves of dogwood (Cornus sp.). 

Material Examined.—INHS: 1 larva 
II; June; on dogwood; Cook County, 
Illinois. 


Scirtothrips taxodii Hood (1954) 
(Fig. 15, 33, 50, and 72) 


Larva II.—Body color yellow, often 
with red-orange body pigment. An- 
tennae, tibiae, bases of femora, and an- 
terior median cephalic spot light brown. 
Antennal segments V-—VII usually 
darker brown. Apices of antennal seg- 
ments I and II and base and apex of 
III pale gray. Eyes red. 

Most setae pointed and short (11-12 
ym); H1, H4, P7, A(IX)2, and A(X)1 
longer and knobbed. A(IX)1 and 
A(IX)3 long and subequal to A(IX)2. 
Cuticle with dense microtrichia. 

Measurements of the S. taxodii larva 
II are shown in Table 12. 


Table 12.—Measurements, in microns, of 
six Scirtothrips taxodii larvae |1. 


Character Lene Width 
Range Mean Range 

Antennal segment 

I 16-17 22-24 

II 18-30 20-23 

Ill 43-46 20-23 

IV 43-46 20-23 

Vv 6-8 12-16 

VI 9-11 9-11 

VII 19-23 Hd 84 
Antenna 164-185 174 est 
Head 78-85 93-100 
Pronotum 85-125 140-155 
Body 670-780 720 Actes 
Setae 

H1 14-16 

H4 14-17 

P7 19-23 

A(IX)1 19 

A(IX)2 25-28 

A(IX)3 23-25 

A(X)1 20-23 


_ "A single measurement in a range column 
indicates that all such measurements were 
identical, 


August, 1974 VANCE: LARVAE OF THE SERICOTHRIPINI 189 


Larva I—Body pale yellow, often ments V—VII all darker brown; tibiae 
with red-orange body pigment. Anten- and bases of femora brown. Eyes red. 
nal segments I and II light brown; seg- Most setae pointed and short (8 »m); 
ments HI and proximal portion of IV only H1 knobbed. Posteroangular setae 
brownish orange; apex of IV and seg- lacking. Anterior median cephalic spot 


MMO NOD, 


uyuuorans aoe? 
waning gunn nb POP EPEOP OOO LHLA Leap ay 
6 
be 
Ao 


A Sa ‘ 


s494,) 
Oy 


prey gt! 


eile 


Fig. 68-77.—Abdominal segments VIII-X (except where indicated) of the second-instar 
larva. 68.—-Sericothrips campestris. 69.—-Sericothrips annulipes, abdominal segments IX and 
X. 70.—Drepanothrips reuteri. 71.—-Dendrothrips ornatus. /2.—Scirtothrips taxodii. 73.— 
Echinothrips americanus. 74.—Pseudodendrothrips mori. 75.—Taeniothrips simplex. 76.— 
Ctenothrips bridwelli. 77.—Scolothrips pallidus. 


190 


lacking. Cuticle with very fine micro- 
trichia. 

Measurements of the S. taxodii larva 
I are shown in Table 13. 


Table 13.—Measurements, in microns, of 
two Scirtothrips taxodii larvae |. 


Length Width 

Character Tine Hanes 
Antennal segment 

I 122 18-20 

II 18-20 19-20 

Ill 27-30 24 

IV 38-47 19-27 

Vv 6-7 12 

VI 6-7 8-9 

VII 15 5-6 
Antenna 120-138 ae 
Head 74 85 
Pronotum 85 119-124 
Body 527-542 aie 
Setae 

Hi 9 

A(IX)1 13 

A(IX)2 24 

A(X)1 15 


_ *A single measurement in a range column 
indicates that all such measurements were 
identical. 


Diagnosis.—S. taxodii larvae are dis- 
tinguished from S. niveus larvae by 
shorter Hl and H4 setae and by 


pointed instead of knobbed H4, © 


A(IX)2, and A(X)1 setae. S. taxodii 
is differentiated from S. brevipennis by 
body color, which tends to be yellow 
in taxodii and orange in brevipennis, 
and by setae Al and A3 on abdominal 
segment IX being longer and closer to 
the length of A2. S. taxodii is found 
on leaves of bald cypress (Taxodium 
distichum). 

Material Examined.—INHS: 2 larvae 
I, 11 larvae II; June-August; on bald 
cypress; Alexander and Massac coun- 
ties, Illinois. 


Sericothrips Haliday (1836) 


Larva II.—Body color pale yellow 
to yellow to yellow-orange, several 
species showing light to heavy orange 
or red hypodermal pigmentation. An- 
tennal segments, tibiae, bases of femora, 


Inurnois NaturaL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


setae, and setal basal rings (and ab- 
dominal segments IX and X in S. an- © 
nulipes) light brown to brown. Apices — 
of antennal segments I and II and base 
and apex of III pale gray. Brown ~ 
sclerotized areas present on anterior — 
median head area and, in certain spe- 
cies, on pteronotum. Eyes red. 
Antennae each seven segmented 
(Fig. 31); longest sense cone on seg- 
ment IV and sense cones on segments — 
V and VI moderately long and slightly. — 
blunted, all subequal. Sense cones on 


segment V slightly shorter. Segments — 


II and III each with a pair of fanned 


setae. Segment III with six annula- — 


tions; very fine microtrichia present on 
the distal annulations. Segment IV 
with five annulations, all with longer, 
more prominent microtrichia. 

Head (Fig. 53) wider than long. 
Eyes with four large facets bulging at — 
sides of head. Mouth cone moderately _ 
blunt. Head with four pairs of dorsal 
setae; Hl and H4 always, and H2 — 
and H3 sometimes, fanned; H1 directly _ 
opposite H2; H3 reduced and much — 
smaller than H4; H4 varying from 
shorter than to subequal to H1. 

Pronotum wider than long and with ~ 
seven pairs of setae. Setae Pl, 3, and ~ 
5 usually shorter, and P7 longer than 
P2, 4, and 6. Mesonotum with seven 
pairs of fanned setae, two pairs lo- 
cated medially and five pairs laterally. 
Metanotum with four pairs of fanned — 
setae, two pairs located medially and 


two pairs laterally. Meso- and meta- — 


notum each with two pairs of brown — 
sclerotized areas in annulipes, pulchel- 
lus, and cingulatus. 

Abdominal segment I with two pairs 
and abdominal segments II-VIII each — 
with three pairs of fanned setae; Al 
usually shortest and A3 usually longest, 
their lengths varying among Serico- — 
thrips species. Segment IX with setae 
Al and A2 long and narrowly fanned, — 
A3 reduced and sometimes fanned. Seg- — 
ment X with Al narrowly fanned. 


Most dorsal body setae fanned to — | 


August, 1974 


varying degrees and of varying lengths. 
Setal bases usually with faint brown 
rings (much larger and more promi- 
pulchellus, 


nent in annulipes, and 


VANCE: LARVAE OF THE SERICOTHRIPINI 191 


cingulatus and to a lesser degree in 
variabilis). Integument with dense 
stippling resulting from very fine micro- 
trichia; stippling forming transverse 


Fig. 78-89.—Abdominal segments II1 and IV (except where indicated) of Sericothrips 


species. 78.—S. cingulatus. 79.—S. 


annulipes. 


80.—S. pulchellus. 81.—-S. variabilis. 


82.—S. baptisiae. 83.—S. campestris. 84.—S. beachae. 85.—S. tiliae. 86.—S. nubilipennis. 
87.—S. sambuci. 88.—S. langei. 89.—S. annulipes, meso- and metanotum. 


192 


rows on abdominal segments, particu- 
larly on segments IX and X (minute 
pustules present in cingulatus). Ab- 
dominal segment IX lacking a posterior 
comb. 


Larva I.—Body color pale yellow to 
orange-yellow, with hypodermal pig- 
ment in some species. Antennal seg- 
ments, tibiae, bases of femora, and 
setae light brown. Apices of antennal 
segments I and II and base and apex of 
III pale gray. Terminal antennal seg- 
ments darker brown. 


Antennae each seven segmented, the 
suture between IV and V sometimes 
indistinct. Sense cones on segments 
IV and V as in larva II, but longer; 
sense cone on segment VI as in larva 
II (Fig. 14). Segments II and III each 
with a pair of fanned setae. Segment 
III with six annulations with small 
microtrichia present on most. Segment 
IV with five annulations with longer 
microtrichia present on and randomly 
between the annulations. Segment VII 
tapering apically. 

Chaetotaxy similar to that of larva 
II, but setae much reduced and fanned 
only terminally; posteroangular setae 
(P7) lacking; mesonotum with four 
pairs and metanotum with three pairs 
of setae; abdominal segment IX with 
two dorsal pairs of fanned setae. An- 
terior median cephalic spot, pteronotal 
sclerotized areas, and raised setal basal 
rings all lacking. Integument with 
stippling as in larva II, but fainter. 


Diagnosis.—Sericothrips larvae can 
be distinguished from those of all other 
Thripinae except Zonothrips by the 
presence of fimbriate fan-shaped setae. 
Larvae of Zonothrips were unavailable 
to me, but they can be separated from 
those of Sericothrips through the as- 
sociated adults and host-plant data. A 
description and illustration of Z. karnyi 
(larva IL) were given by Priesner 
(19262), but this description and illus- 
tration are lacking in diagnostic char- 
acters sufficient to separate Zonothrips 
from Sericothrips. 


Ittinois NATuRAL History Survey BULLETIN 


Vol. 31, Art.5 


Characters used to differentiate be- 
tween larvae of Sericothrips species 
vary in their value. Presence or ab- 
sence of large setal basal rings and 
brown sclerotized areas on the pterono- 
tum are always consistent. Presence or 
absence of an anterior median cephalic 
spot and the proportions of the lengths 
of the abdominal dorsal setae are fairly 
reliable. Body color, hypodermal pig- 
mentation, proportionate lengths of cer- 
tain head and pronotal setae, and the 
general length and width of the setae 
are useful only when used in conjunc- 
tion with other characters. 


KEY TO THE MATURE LARVAE II 
OF SERICOTHRIPS 


1. Pteronotum with brown sclerotized 
areas (Fig. 89); abdominal and 
pteronotal setae with basal rings 
moderately or greatly enlarged (7- 
15 um in diameter) ............... 2 


Pteronotum lacking brown sclerotized 
areas; abdominal and pteronotal 
setae with basal rings reduced and 
faint (6-7 wm in diameter) (the 
lateral abdominal setae with larger 
rings in variabilis) ............05% 4 


2. Cuticle with microtrichia on small 
pustules; setal basal rings only 
moderately enlarged (Fig. 78) .... 

a info w Ble, Shane fe ave, 6 ab aeete ca) everett een cingulatus 


Cuticle with microtrichia alone result- 
ing in a stippled pattern; setal basal 
rings greatly enlarged ............ 5 


3. Abdominal segments IX and X usu- 
ally brown; abdominal segment IV 
with setal pair A2 significantly 
shorter than A3, and Al nearly sub- 
equal to A2 (Fig. 79); found on 
black locust 


Abdominal segments IX and X not 
brown; ~abdominal segment IV with 
setal pair A2 subequal to A3, and Al 
significantly shorter than A2 (Fig. 
80); found on wafer ash...pulchellus 


4. Dorsal body setae generally short, 
the longest setae rarely exceeding 
20-25 wm and either widely or nar- 
rowly fanned; abdominal segment 
IV with setae Al and A2 subequal F 
and significantly shorter than A3... 5 — 


Dorsal body setae generally long, the 
longer setae measuring up to 30-35 
“um; proportions of abdominal setae 
variable 


August, 1974 


5. Body setae widely fanned (Fig. 82); 
found on false indigo ....... baptisiae 

Body setae narrowly fanned (Fig. 
88); found on water lily....... langei 

6. Setal pair A2 on anterior abdominal 
segments subequal to A3, the length 
BALAI VAR MINE Tee eS hie «onal dlaca ove 3. 5 elseoce ni 


Setal pair A2 on anterior abdominal 
segments significantly shorter than 
A3, and Al usually significantly 
SV QOT HITS gS 0 SNAP ee a ene EY 8 


7. Setal pair Al subequal to A2 (Fig. 
83); A3 not over 25 um long on 
abdominal segment IV; found on 
wild four-o’clock .......... campestris 

Setal pair Al shorter than A2 (Fig. 
87); A3 up to 30 um long on abdom- 
inal segment IV; found on elder- 
TET ects ie aie Gans ia > pia cts ete wee sambuci 


8. Setae generally widely fanned (Fig. 

81 and 84) 
Setae generally narrowly fanned (Fig. 

SOCAN OO )e an chia ala achietss-ctceus aaesle 10 


9. Lateral abdominal setae with large 
basal rings (Fig. 81); body color 
yellow to orange, sometimes with 
red hypodermal pigmentation ..... 

variabilis 


Lateral abdominal setae with reduced 
basal rings (Fig. 84); body color 
pale white to yellow without red 
hypodermal pigmentation ....beachae 


10. Body color usually white to whitish 
yellow; found on various forest 
trees and in forest debris. .nubilipennis 


Body color usually more yellow; found 
on basswood 


Sericothrips annulipes Hood (1927b) 
(Fig. 31, 53, 69, and 79) 


Larva If.—Body yellow with promi- 
nent red-orange hypodermal pigmenta- 
tion dorsally in mature larvae. Anten- 
nae, tibiae, bases of femora, abdominal 
segments IX and X, setae, setal rings, 
and cephalic spot brown. Apices of 
antennal segments I and II and base 
and apex of segment III pale gray. 
Eyes red. 


Most dorsal body setae fairly long 
and fairly narrowly fanned. Longer 
body setae generally twice the length 
of the shorter setae; longer setae, 22— 
30 pm, shorter setae, 9-16 pm. Bases 
of setae with prominent raised brown 


VANCE: LARVAE OF THE SERICOTHRIPINI 


193 


rings (7-13 »m in diameter) on ab- 
domen and pteronotum. Head some- 
times with a truncate apical point; 
anterior median brown cephalic spot 
usually present. Mesonotum and meta- 
notum each with two pairs of brown 
sclerotized areas (Fig. 89). Abdominal 
segment IV with setal pair Al subequal 
to A2, and both significantly shorter 
than A3 (Fig. 79); Al usually shorter 
than A2 on segment IX. 


Measurements of the S. annulipes 
larva II are shown in Table 14. 


Table 14.—Meesurements, in microns, of 
10 Sericothrips annulipes larvae ||. 


Length Width 
Character Range Mean Range 
Antennal segment 
Ill 48-54 21-22 
IV 45-52 18-21 
V 6-7 10-13 
VI 1) 36 7-10 
Vil 18-22 hee 4-6 
Antenna 170-189 176 Aas 
Head 132-150 87-97 
Pronotum 90-127 135-160 
Body 882-1,061 995 
Setae 
Hl 25-30 
12 Y/ 25-32 
A(IV)1 9-13 
A(IV)2 13-18 
A(IV)3 24-30 
A(IX)1 22-28 
A(IX)2 28-31 
Ventral setae 
(1X) 9-15 


Larva I.—Body pale yellow to yel- 
low. Antennal segments I-IV, legs, and 
setae light brown; antennal segments 
V-VII darker brown. Eyes red. 


Most body setae fanned or expanded 
only terminally, fimbriate for most of 
their length, and shorter than in larva 
II (10-20 »m in larva I). Setal basal 
rings and brown sclerotized areas of 
pteronotum lacking. Abdominal cuticle 
with faint stippling, but more promi- 
nent and forming transverse rows on 
segments IX and X. 

Measurements of the S. annulipes 
larva I are shown in Table 15. 


194 


Table 15.—Measurements, in microns, of 
seven Sericothrips annulipes larvae |. 


Length Width 

Character Range Range 
Antennal segment 

I 14-19 19-25 

II 22-28 20-25 

III 28-36 22-25 

IV 46-50 22-25 

Vv 58 11-12 

VI 6-8 8 

VII 19-23 5 
Antenna 140-169 Ha 
Head 165-178 105-120 
Pronotum 120-135 180-186 
Body 571-681 
Setae 

H1 11-12 

A(IX)1 12-19 

A(IX)2 12-20 


_ * A single measurement in a range column 
indicates that all such measurements were 
identical, 


Diagnosis.—S. annulipes larvae are 
similar to those of pulchellus and cingu- 
latus. All possess brown sclerotized 
areas on the pteronotum and enlarged 
rings at the setal bases. S. annulipes 
can be distinguished from cingulatus 
by the lack of cuticular pustules and 
by the presence of some shorter body 
setae in annulipes. It differs from 
pulchellus by the brown color of ab- 
dominal segments IX and X and in 
the proportions of the anterior abdomi- 
nal setae. In annulipes setal pair 
A(IV)1 is shorter than A(IV) 2 and 
both are shorter than A(IV)3; in 
pulchellus (Fig. 80) A(IV)1 is shorter 
than A(IV)2, which is subequal to 
A(IV)3. S. annulipes is found on 
locust trees, particularly black locust, 
throughout the state. 

Material Examined.—INHS: 14 
larvae I, 10 larvae II; on black locust; 
Johnson, Piatt, Putnam, Stephenson, 
and Union counties, Illinois. 


Sericothrips baptisiae Hood (1916) 
(Fig. 82) 
Larva Il.—Body yellow to yellow- 


orange, without hypodermal coloring. 
Antennae, tibiae, bases of femora, 


ILntinois NATURAL History SurvEY BULLETIN 


Vol. 31, Art. 5 


setae, setal rings, anterior cephalic spot, 
and sometimes abdominal segment X 
light brown to brown. Apices of an- 
tennal segments I and II and base and 
apex of III pale gray. Eyes red. 
Most setae, except Hl, P7, and 
A(IX)1 and 2, short (22-33 pm) and 
widely fanned; shorter setae, 9-13 pm; 
longer setae, 15-18 »m. Bases of setae 
with small unraised brown rings (6 
ym in diameter). Head with anterior 
median brown spot. No dorsal brown , — 
sclerotized areas on pteronotum. Ab- 
dominal segment IV with setal pair Al 
subequal to A2 and both significantly 
shorter than A3 (Fig. 82). Stippling — 
on abdominal segments forming irregu- 
lar transverse rows. j 


Measurements of the S. baptisiae 
larva II are shown in Table 16. 


Table 16.—Measurements, in microns, of 
three Sericothrips baptisiae larvae II. 


: Length Width 
COT Range Mean Range 
Antennal segment 
Ill 52-55 Bes 22-24 
IV 51-55 Hae 21-22 
Vv 1 ae 10-12 
VI 7 ok, 10 
VII 21-24 iv 6-7 
Antenna 185-202 198 se 
Head 165-178 105-120 ae 
Pronotum 120-135 hi 180-186 
Body 791-889 936 
Setae 
H1 22-25 
P7 21-24 
A(IV)1 10 
A(IV)2 10-13 
A(IV)3 18-19 
A(IX)1 21-30 
A(IX)2 29-33 
Ventral setae 
(1X) 15-18 


4A single measurement in a range column ~ 
indicates that all such measurements were — 
identical. 


Diagnosis.—S. baptisiae larvae are 
distinguished from other Sericothrips 
larvae by having short and widely _ 
fanned dorsal body setae and by setae 
Al and A2 on abdominal segment TV _ 
being equal and significantly shorter 


August, 1974 


than A3. S. langei larvae are similar 
to baptisiae larvae in setal length and 
proportions, but have narrower setae. 
S. baptisiae is found exclusively on 
false indigo (Baptisia) throughout the 
state. 


Material Examined.—INHS: 3 larvae 
II; September; .on Baptisia; Adams 
and Vermilion counties, Illinois. 


Sericothrips beachae Hood (1927a) 
(Fig. 84) 


Larva Il.—Body very pale yellow or 
white without hypodermal pigmenta- 
tion. Antennae, tibiae, bases of femora, 
setae, and anterior cephalic spot light 
brown. Apices of antennal segments I 
and II and base and apex of III pale 
gray. Eyes red. 

Most body setae fairly long and mod- 
erately fanned. Longer body setae 
(24-36 »m) about twice the length of 
shorter body setae (10-16 »m). Bases 
of setae with very faint small rings. 
Apex of head sometimes pointed and 
with an anterior median brown spot. 


Table 17.—Measurements, in microns, of 
four Sericothrips beachae larvae II. 


Character penee wie 
Range Mean Range 
Antennal segment 
Ill 45-60 ahs 228 
IV 45-52 % 17-19 
Vv 6-7 ec te 
VI 7-9 nets 7-9 
VII 24 Aa 6 
Antenna 157-202 189 an 
Head 150-180 82-97 
Pronotum 115-135 150-180 
Body 725-995 890 
Setae 
H1 30-36 
P7 29-31 
A(IV)1 9-13 
A(IV)2 12-18 
A(IV)3 18-27 
A(IX)1 31-33 
A(IX)2 31-34 
Ventral setae 
(TX) 18-23 


ee single measurement in a range column 
indicates that all such measurements were 
identical, 


VANCE: LARVAE OF THE SERICOTHRIPINI 


195 


No brown sclerotized areas on pterono- 
tum. On abdominal segment IV setal 
pair Al subequal to A2 (occasionally 
decidedly shorter) and both signifi- 
cantly longer than A3 (Fig. 84). Stip- 
pling on abdominal segments faint and 
forming irregular transverse rows. 

Measurements of the S. beachae 
larva II are shown in Table 17. 

Diagnosis.—S. beachae larvae can be 
distinguished from other light-colored 
Sericothrips larvae with longer body 
setae by their wide body setae and by 
setal pair A(IV)2 being significantly 
shorter than A(IV)3. S. beachae can 
be distinguished from variabilis, a 
closely related species, by the total 
lack of any hypodermal and cuticular 
pigmentation and by the absence of 
small basal rings on lateral abdominal 
setae. S. beachae is found on hops in 
many areas of the state. 

Material Examined.—INHS: 4 larvae 
Il; 2 June 1970; on hops; Iroquois 
County, IHlinois. 


Sericothrips campestris Hood (1939) 
(Fig. 68 and 83) 


Larva II.—Body usually yellowish 
orange without hypodermal pigmenta- 
tion. Antennae, tibiae, bases of femora, 
setae, and setal rings brown. Antennal 
segment I, apex of II, and base and 
apex of III pale gray. Eyes red. 

Setae fairly long and moderately 
fanned. Shorter body setae (12-22 
um) usually about two-thirds the length 
of longer setae (22-31 »m). Setae with 
faint, unraised, brown basal rings (8 
pm in diameter). Apex of head ob- 
tusely pointed; anterior median cephalic 
spot lacking. Dorsal sclerotized areas 
lacking. Lengths of setae on abdominal 
segment IV unequal to subequal (Fig. 
83). Setal pair Al usually subequal to 
A2 on segment IX (Fig. 68). Stippling 
on abdominal segments forming definite 
transverse rows on terminal segments 
and to a lesser extent on others. 

Measurements of the S. campestris 
larva II are shown in Table 1S. 


196 


Table 18.—Measurements, in microns, of 
10 Sericothrips campestris larvae |I. 


Length Width 
Character Range Mean Range 
Antennal segment 
Ill 52-55 22-27 
IV 48-60 19-23 
Vv 6-9 12-14 
VI 9-12 9-10 
Vil 21-24 cg 6-8 
Antenna 195-220 202 an 
Head 112-150 175-210 
Pronotum 147-180 97-120 
Body 990-1,179 1,001 
Setae 
H1 25-31 
P7 28-34 
A(IV)1 15-22 
A(IV)2 16-30 
A(IV)3 22-30 
A(IX)1 24-34 
A(IX)2 29-35 
Ventral setae 
(1X) 22-30 


Larva I—Body yellow to yellow- 
orange. Antennae, tibiae, and setae 
very light brown. Antennal segment 
I, apex of II, and base and apex of 
III pale gray. Eyes red. 

Most dorsal body setae fanned or 
expanded only terminally, fimbriate for 
most of their length, and shorter than 


Table 19.—Measurements, in microns, of 
three Sericothrips campestris larvae |. 


Length Width 

Character epiee Range 
Antennal segment 

I 16-19 23-26 

II 25-28 23-25 

Ill 23-39 25-28 

IV 46-54 23-25 

Vv 5-8 11-12 

VI 8-11 8" 

Vil 20-22 5-6 
Antenna 150-170 sat 
Head 62-85 78-85 
Pronotum 85-93 116-140 
Body 573-636 
Setae 

H1 12-14 

A(IX)1 14 

A(IX)2 19-22 


_ “A single measurement in a range column 
indicates that all such measurements were 
identical, 


Intinors NatuRAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


in Jarva IL (10-22 pm in Jarva 1). Setal 
rings lacking. Abdominal cuticle with 
faint stippling, becoming more promi- 
nent on terminal segments. 
Measurements of the S. campestris — 
larva I are shown in Table 19. 
Diagnosis.—Larvae of S. campestris 
resemble those of sambuci in having 
long setae and having setal pair A2 
subequal to A3 on segment IV; Al is 
subequal to A2 on segment IV in, 
campestris, but is shorter in sambuci, 
and sambuci has slightly longer setae. 


S. campestris occurs on wild four- 
oclock (Mirabilis nyctaginea) along 
gravelly railroad embankments in IIli- 
nois. 

Material Examined.—INHS: 3 larvae 
I, 12 larvae II; June-August; on wild 
four-o’ clock; Champaign and Vermilion 
counties, Illinois; Lathrop, Missouri; 
and Ogallala, Nebraska. 


Sericothrips cingulatus 
Hinds (1902) 
(Fig. 78) 


Larva II—Cuticle yellow to orange 
without hypodermal pigmentation. An- 
tennal segments, tibiae, bases of femora, 
setae, setal basal rings, sclerotized areas 
on pteronotum, and abdominal segment 
X brown to dark brown. Apices of an- 
tennal segments I and II and base and 
apex of III pale gray. 

Most dorsal body setae long and 
widely fanned. Bases of setae on ab- 
domen_and pterothorax with prominent 
brown rings (8-12 »m in diameter). 
Mesonotum and metanotum each with 
a pair of brown sclerotized areas as 
in S. annulipes (Fig. 89). Anterior ab- 
dominal segments with dorsal setae 
equal or subequal (Fig. 78); Al on 
segment IX subequal to A2. Abdomi- 
nal stippling large, almost forming 
small pustules (Fig. 78). 

Measurements of the S. cingulatus 
larva II are shown in Table 20. 

Diagnosis.—S. cingulatus larvae re- 
semble the larvae of the annulipes 


August, 1974 


Table 20.—Measurements, in microns, of 
one Sericothrips cingulatus farva ||. 


Character Length Width 
Antennal segment 

I 23 39 

II 39 29 

III 59 28 

IV 37 23 

Vv 6 17 

VI 12 12 

VII 28 6 
Antenna 204 epee 
Head 101 116 
Pronotum 140 202 
Body 1,050 
Setae 

H1 31 

P7 28 

A(IV)1 26 

A(IV)2 29 

A(IV)3 29 

A(IX)1 31 

A(IX)2 39 


Ventral setae (IX) 12 


group of Sericothrips in possessing 
pteronotal sclerotized areas and promi- 
nent (although smaller) rings at the 
bases of the setae, and in having ab- 
dominal segment IX brown. S. cingu- 
latus can be recognized by its dark 
brown body color, its having almost 
all dorsal body setae long and equal or 
subequal, and the presence of small 
pustules. 


Larval and adult morphology of 
cingulatus indicate that this species is 
atypical of most Sericothrips. Adults 
of cingulatus possess dense abdominal 
microtrichia completely covering the 
tergites, whereas the tergites of most 
species in this genus possess micro- 
trichia only laterally. In the larvae the 
cuticular stippling is modified into 
small pustules, and the brown sclero- 
tized areas are similar to those found 
in the more primitive Chirothripini. 


S. cingulatus is found scattered state- 
wide in grassland areas and particularly 
in grass-sedge marshes. 


Material Examined.—INHS: 1 larva 
ll; 18 March 1971; on clover and 
vetch; Amite, Louisiana. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


197 


Sericothrips langei Moulton (1929) 
(Fig. 88) 


Larva II.—Body white to light yel- 
low, sometimes light orange, without 
subintegumental pigmentation. Anten- 
nae, tibiae, bases of femora, setae, and 
setal rings light brown. Most of an- 
tennal segment I, the apex of II, and 
the base and apex of III pale gray. 
Eyes red. 

Most body setae short and all nar- 
rowly expanded. Shorter body setae 
(7-13 wm) usually about one-half the 
length of longer setae (13-24 ym). Setae 
with small faintly brown unraised rings 
(6 »m in diameter). Head often with 
an apical point. Anterior cephalic spot 
lacking. Abdominal segment IV with 
setal pair Al subequal to A2 and both 
significantly shorter than A3 (Fig. 88). 
Setal pair Al on segment IX signifi- 
cantly shorter than A2. Stippling on 
abdomen forming prominent transverse 
rows on the terminal segments, becom- 
ing less prominent on the others. 


Measurements of the S. langei larva 
II are shown in Table 21. 


Table 21.—Measurements, in microns, of 
nine Sericothrips langei larvae |. 


Length 


Character Width 
Range Mean Range 
Antennal segment 
Ill 48-55 22-27 
IV 21-23 21-23 
V 7-12 13-15 
VI 10° 10-12 
VII 22-30 6-7 
Antenna 202-227 214 anor 
Head 180-190 ine 123-130 
Pronotum 123-140 so 180-202 
Body 901-1,159 1,040 
Setae 
Hi 18-22 
P7 25-31 
A(IV)1 7-11 
A(IV)2 8-15 
A(IV)3 15-24 
A(IX)1 13-17 
A(TX)2 18-25 
Ventral setae 
(TX) 12-15 


2A single measurement in a range column 
indicates that all such measurements were 
identical. 


198 


Larva I—Body light yellow to yel- 
low-orange. Antennal segments I-IV, 
tibiae, and setae very light brown; an- 
tennal segments V—VII darker brown. 
Eyes red. 

Most body setae expanded only ter- 
minally and shorter than those of larva 
II (6-15 »m in larva I). Setal rings 
lacking. Cuticle with faint stippling 
becoming more prominent on the ter- 
minal segments. 

Measurements of the S. langei larva 
I are shown in Table 22. 

Diagnosis.—S. langei larvae can be 
distinguished from all other Serico- 
thrips larvae considered here by the 
very short and very narrowly fanned 
setae. The species is found on water 
lilies of the genus Nymphaea through- 
out the state. 

Material Examined.—INHS: 5 larvae 
I, 20 larvae II; June-August; on water 
lily; Lake and Monroe counties, IIli- 
nois, and Au Train, Michigan. 


Table 22.—Measurements, in microns, of 
four Sericothrips langei larvae |. 


i Length Width 

Character fries anes 
Antennal segment 

I om weal) 26-28 

II 28-31 23-26 

Ill 31-37 25-28 

Vv 5-6 11-12 

VI 9-12 8 

Vil 20-25 5 
Antenna 162-1.74 fate 
Head 85-100 93-100 
Pronotum 85-124 140 
Body 642-734 os 
Setae 

Hi 9-12 

A(IX)1 8-9 

A(TX)2 14-16 


“A single measurement in a range column 
indicates that all such measurements were 
identical, 

Sericothrips nubilipennis 
Hood (1924) 
(Fig. 86) 

Larva II.—Body whitish yellow with- 

out hypodermal pigmentation. Anten- 


Ittivois NATuRAL History SuRVEY BULLETIN 


Vol. 31, Art. 5 


nae, tibiae, and setae light brown. An- 
tennal segment I, base of II, and base 
and apex of III pale gray. Eyes red. 
Setae fairly long and moderately — 
fanned. Shorter body setae (12-18 »m) 
generally two-thirds the length of the 
longer setae (19-31 pm). Setae with 
very faint basal rings (6 »m in di- 
ameter). Apex of head usually rounded; ~ 
median anterior spot lacking. Dorsal — 
sclerotized areas lacking. Abdominal — 
segment IV with all setae varying from 
decidedly unequal to subequal in cer- 
tain cases (Fig. 86). Setal pair Al of 
segment IX usually shorter than A2. 
Abdominal stippling generally faint, 
transverse rows being prominent mainly — 
on the terminal segments. h 
Measurements of the S. nubilipennis 
larva II are shown in Table 23. o 


Table 23.—Measurements, in microns, of — 
five Sericothrips nubilipennis larvae ||. 


Length Width — 
Sa Range Mean Range ~ 
Antennal segment y 
Ill 45-55 45 20-22 — 
IV 48-60 Sra 18-21 
Vv 7-13 og 12-15 
VI 10-12 ash 9-10 
VII 26-30 ass 5-7 
Antenna 189-214 202 eae 
Head 135-165 =e 90-112, 
Pronotum 112-135 157-202 
Body 850-1,033 932 nn 
Setae 
H1 ov-el 
P7 22-56 
A(IV)1 10-18 
A(IV)2 12=22 
A(IV)3 1S=25 
A(TX)1 _ 22-30 
A(IX)2 22-31 
Ventral setae 
(IX) 15-22 


Diagnosis.—Larvae of S. nubilipen- 
nis are distinguished by their narrowly — 
fanned setae and by A(IV)2 being — 
significantly shorter than A3. This — 
species is similar to and often indis-— 
tinguishable from S. tiliae. The body 
color of nubilipennis tends to be white, 
whereas that of tiliae tends to be yel- 


August, 1974 


low. Host-plant data are unreliable 
criteria, too, since both species can 
occur on adjacent forest plants with 
accidental transfers being made from 
one host to the other. 


S. nubilipennis generally occurs on 
various forest trees, such as_hack- 
berry (Celtis) or dogwood (Cornus) 
throughout the state. 


Material Examined.—INHS: 5 larvae 
II; June—October; on hackberry leaves 
and forest leaf litter; Champaign, Hen- 
derson, Macon, and Piatt counties, IIli- 
nois. 


Sericothrips pulchellus Hood (1908) 
(Fig. 80) 


Larva I.—Cuticle orange with prom- 
inent red-orange hypodermal pigmenta- 
tion, often faint. Antennae, tibiae, bases 
of femora, setae, setal rings, anterior 
cephalic spot, and dorsal sclerotized 
pteronotal areas light brown to brown. 
Apices of antennal segments I and II 
and base and apex of III gray. 


Table 24.—Measurements, in microns, of 
10 Sericothrips pulchellus larvae |I. 


VANCE: LARVAE OF THE SERICOTHRIPINI 


199 


Most dorsal body setae fairly long 
and moderately fanned. Shorter body 
setae (12-20 pm) usually two-thirds 
the length of the longer setae (22-31 
pm). Bases of setae with prominent, 
raised brown rings (7-15 »m in di- 
ameter) on abdomen and pterothorax. 
Head usually without an apical point; 
anterior median cephalic spot present. 
Mesonotum and metanotum each with 
a pair of brown sclerotized areas. Ab- 
dominal segment IV with setal pair 
A2 subequal to A3 and both signifi- 
cantly longer than Al (Fig. 80). Setal 
pair Al on segment IX usually shorter 
than A2. Abdominal stippling forming 
prominent transverse rows on most 
abdominal segments (Fig. 80). 


Measurements of the S. pulchellus 
larva II are shown in Table 24, 


Larva I—Body orange. Antennae 
and setae generally light brown. Apex 
of antennal segment IV and all of 
segments V—VIII brown. Eyes red. 


Body setae expanded terminally and 
shorter than in larva II (9-25 pm in 
larva I). Setal basal rings, cephalic 
spot, and sclerotized pteronotal areas 
lacking. Abdominal stippling faint. 


Length Width 
Character 4 ‘ 
Range Mean Range Table 25.—Measurements, in microns, of 
five Sericothrips pulchellus larvae |. 
Antennal segment 
III 44-52 “Ne 22° Length Width 
IV 44-52 20-22 Character Range Range 
Vv 7 12-15 
VI 10 ee 10-13 Antennal segment 
VII 24-30 cers 7 I 11-16 23-25 
Antenna 160-189 176 Seen II 23-25 22-23 
Head 150-165 97-112 III 26-31 23-26 
Pronotum 97-142 on 150-210 IV 42-48 23-26 
Body 867-1,128 945 Vv 5-6 9-12 
Setae VI 8-9 8" 
H1 27-32 VII 22-2 5 
P7 30-32 Antenna 150-158 sere 
A(IV)1 12-18 Head 70-78 78-85 
A(IV)2 19-27 Pronotum 70-85 101-132 
A(IV)3 22-30 Body 496-611 
A(IX)1 21-30 Setae 
A(IX)2 27-33 H1 11-19 
Ventral setae A(IX)1 19-25 
(1X) 15-18 A(IX)2 12-16 
* A single measurement in a range column * A single measurement in a range column 
indicates that all such measurements were indicates that all such measurements were 


identical. 


identical. 


200 


Measurements of the S. pulchellus 
larva I are shown in Table 25. 

Diagnosis.—S. pulchellus larvae can 
be distinguished from all other Serico- 
thrips larvae considered here except 
annulipes and cingulatus by the large 
raised rings at the bases of the setae 
and by the brown sclerotized areas on 
the pteronotum. Red hypodermal pig- 
mentation is also characteristic of this 
species although it is sometimes faint 
or absent. 

S. pulchellus can be distinguished 
from cingulatus by the absence of 
cuticular pustules and by some body 
setae being shorter than others. S. 
pulchellus differs from annulipes in 
the absence of brown coloration on 
abdominal segments IX and X and in 
the length proportions of setae on ab- 
dominal segment V; in annulipes setal 
pair Al is subequal to A2 and both 
are shorter than A3; in pulchellus A2 
is subequal to A3 and both are longer 
than Al. 

S. pulchellus is found on wafer ash 
(Ptelea), sometimes in great numbers. 
Adults and larvae feed together, often 
causing the foliage to turn white be- 
cause of the feeding scars. At times of 
great abundance, adults may be scat- 
tered and found resting on a variety of 
trees and shrubs (Stannard 1968). 

Material Examined.—INHS: 6 larvae 
I, 23 larvae II; June-August; on wafer 
ash; Carroll, Kankakee, Mason, and 
Winnebago counties, Illinois. 


Sericothrips sambuci Hood (1924) 
(Fig. 87) 


Larva II.—Body yellow. Antennal 
segments I and II yellow; segments 
III-VII and setae light brown. Base 
and apex of antennal segment III pale 
gray. Eyes red. 

Most body setae long and moderately 
fanned. Shorter body setae (10-16 »m) 
usually less than half the length of 
longer setae (24-29 pm). Bases of 
setae without rings. Apical cephalic 
point, median cephalic spot, and dorsal 


Intutinois NAturAL History SurvEY BULLETIN 


Vol. 31, Art. 5 


sclerotized areas lacking. Abdominal 
segment IV with setal pair A2 subequal 
to A3 and much longer than Al (Fig. 
87); Al on segment IX subequal to 
A2. Abdominal stippling forming fine 
transverse rows. 

Measurements of the S. sambuci 
larva II are shown in Table 26. 


Table 26.—Measurements, in microns, of 
10 Sericothrips sambuci larvae II. 


Length Width . 

OREO Range Mean Range 
Antennal segment 

III 48-60 22-26 

IV 51-60 21-33 

Vv 7-10 13-15 

VI 10-13 ie 10-13 

VII 27-32 mate 6-7 
Antenna 202-233 220 =e 
Head 150-180 98-120 
Pronotum 120-142 165-202 
Body 900-1,216 1,046 
Setae 

H1 30-39 

P7 30-42 

A(IV)1 10-15 

A(IV)2 24-31 

A(IV)3 25-33 

A(IX)1 27-36 

A(IX)2 30-42 

Ventral setae 

(IX) 22-30 


Larva I.—Body yellow. Antennal 
segments I-IV pale brown; apex of 
IV, all of V—VII, and setae light brown. 
Eyes pale red. 


Body setae fairly long and expanded 


terminally; setae generally shorter than 


in larva II (10-25 »m in larva I). 
Stippling on abdominal cuticle faint. 

Measurements of the S. sambuci 
larva I are shown in Table 27. 

Diagnosis.—Larvae of S. sambuci re- 
semble those of campestris in having 
long setae and A(IV)2 subequal to 
A(IV)3. However, A(IV)1 is subequal 
to A(IV)2 in campestris and shorter 
than A2 in sambuci. 

S. sambuci is found statewide on 
elderberry (Sambucus). 

Material Examined.—INHS: 1 larva 
I, 17 larvae II; August—-October; on 


August, 1974 


Table 27.—Measurements, in microns, of 
one Sericothrips sambuci larva |. 


Character Length Width 
Antennal segment 
I 16 26 
II 28 26 
Ill 34 26 
IV 50 26 
av 5 12 
VI 8 9 
VII 23 4 
Antenna 160 
Head 78 78 
Pronotum 93 124 
Body 621 
Setae 
H1 21 
A(IX)1 16 
A(IX)2 19 


Sambucus; Calhoun, Iroquois, Marion, 
and Union counties, Illinois. 


Sericothrips tiliae Hood (1931) 
(Fig. 85) 
Larva II.—Body yellow (dark orange 


in one specimen). Antennae, legs, and 
setae uniformly light brown. Eyes red. 


Table 28.—Measurements, in microns, of 
seven Sericothrips tiliae larvae ||. 


Length Width 
ees Range Mean Range 
Antennal segment 
Ill 52-56 22-24 
IV 49-57 16-22 
Vv 7-9 13-15 
VI 9-12 9-12 
VII 23-30 Aen fe 
Antenna 189-214 202 wee 
Head 135-180 90-107 
Pronotum 105-135 157-172 
Body 838-1,014 882 
Setae 
H1 30-36 
PT 27-34 
A(IV)1 9-13 
A(IV)2 15-18 
A(IV)3 24-29 
A(IX)1 24-30 
A(IX)2 27-35 
Ventral setae 
(IX) 9-12 


_ “A single measurement in a range column 
indicates that all such measurements were 
identical. 


VANCE: LARVAE OF THE SERICOTHRIPINI 201 


Most dorsal body setae fairly long 
and moderately fanned; longer setae 
(22-33 »m) usually twice the length 
of shorter setae (9-18 »m). Anterior 
median cephalic spot, brown pteronotal 
areas, and setal basal rings all lacking. 
Abdominal segment IV with setal pair 
A(IV)1 shorter than A(IV)2, and 
A(IV)2 shorter than A(IV)3 (Fig. 85); 
Al on segment IX subequal to A2. 

Measurements of the S. tiliae larva 
II are shown in Table 28. 

Larva I.—Body yellow. Antennal 
segments I-IV and setae light brown; 
segments V—VII darker brown. Eyes 
red. Setae only slightly expanded and 
5-17 »m long. 

Measurements of the S. tiliae larva 
I are shown in Table 29. 


Table 29.—Measurements, in microns, of 
one Sericothrips tiliae larva |. 


Character Length Width 
Antennal segment 

III 36 20 

IV 50 20 

Vv 6 11 

VI 8 8 

VII 14 5 
Antenna 165 a5 
Head 78 70 
Pronotum 85 110 
Body 622 
Setae 

H1 9 

A(IX)1 9 

A(IX)2 iff 


Diagnosis.—Larvae of S. tiliae are 
distinguished by their narrowly fanned 
setae and by A(IV)2 being significantly 
shorter than A(IV)3. This species is 
similar to and often indistinguishable 
from S. nubilipennis. The body color 
of tiliae is usually yellow, whereas that 
of nubilipennis is usually white. 

S. tiliae is found statewide on linden 
(Tilia), being most common in the 
northern part of the state. 

Material Examined.—INHS: 1 larva 
I, 8 larvae II; July-September; on 
linden; Effingham and Kankakee coun- 
ties, Illinois. 


202 


Sericothrips variabilis (Beach 1896) 
(Fig. 14, 32, and 81) 


Larva I.—Body color white, chang- 
ing to yellow and orange with increas- 
ing maturity of larva; red hypodermal 
pigment occasionally present in mature 
larva. Antennae, tibiae, setae, setal 
rings, and anterior median cephalic 
spot brown. Antennal segment I, apex 
of II, and base and apex of III pale 
gray. Eyes red. 


Most dorsal body setae moderate in 
length and moderately fanned. Shorter 
body setae (13-19 »m) between one- 
half and two-thirds the length of the 
longer setae (25-33 pm). Bases of 
pteronotal and abdominal setae with 
small, faint brown rings (6 »m in di- 
ameter); the lateral abdominal setae 
with larger (8 »m in diameter) and 
more prominent rings. Anterior me- 
dian cephalic spot present but often 
not visible in balsam mounts. Brown 
pteronotal areas lacking. Abdominal 
segment IV with setae Al, A2, and 
A3 progressively longer (Fig. 81); seg- 
ment IX with Al usually subequal to 
A2. 


Table 30.—Measurements, in microns, of 
10 Sericothrips variabilis larvae II. 


Inuivors NAturRAL History SURVEY BULLETIN 


oe ENE Width 
ree Range Mean Range 

Antennal segment 

Ill 53-55 22-24 

IV 48-55 18-21 

Vv 7-9 10-13 

VI 10-12 : 9-10 

VII 25-27 shee 6-7 
Antenna 189-202 195 Bees 
Head 142-165 105-112 
Pronotum 120-150 Bran 165-195 
Body 882-1,089 1,021 
Setae 

H1 30-31 

ely 29-39 

A(IV)1 12-19 

A(IV)2 15-24 

A(IV)3 22-31 

A(IX)1 25-31 

A(IX)2 30-34 

Ventral setae 

(TX) 8-11 


Vol. 31, Art. 5 


Measurements of the S. variabilis 
larva II are shown in Table 30. 

Larva I—Body color white to yel- 
low. Antennal segments I-IV light 
brown; apex of IV and all of V—-VII 
darker brown; apices of segments I 
and II and base and apex of III pale 
gray. Setae expanded terminally and ~ 
measuring 6-16 »m. Anterior median 
cephalic spot and setal rings lacking. 

Measurements of the S. variabilis , 
larva I are shown in Table 31. 


Table 31.—Measurements, in microns, of 
five Sericothrips variabilis larvae |. 


Giamacten Length Width 
Range Range 

Antennal segment 

III 30-34 25-26 

IV 46-50 22-26 

Vv 5-6 11-12 

VI 8 6-9 

VII 20-22 5 
Antenna 158-171 wale 
Head 54-85 78-85 
Pronotum 78-101 115-132 
Body 490-621 nets 
Setae 

H1 9-12 

A(IX)1 11-12 

A(IX)2 16-20 


aA single measurement in a range column 
indicates that all such measurements were 
identical. 

Diagnosis.—Larvae of S. variabilis 
resemble those of beachae in having 
wider setae and A(IV)2 shorter than 
A(IV)3. From beachae, variabilis can 
be distinguished by the larger lateral 
setal basal rings and by often having 
cuticular and hypodermal coloration. 

S. variabilis occurs statewide on 
many legumes, particularly soybeans. 
Its life history is discussed earlier in 
this report. 

Material Examined.—INHS: 24 
larvae I, 74 larvae II; August; on and — 
reared from soybeans; Champaign — 
County, Illinois. 


Zonothrips Priesner (1926a) 


Larva II.—Cuticle yellow, generally — 
with red hypodermal pigment. Eyes 


August, 1974 


red. Antennae, setae, and probably 
tibiae and bases of femora light brown. 


Antennae six (possibly seven) seg- 
mented; segments II and III each with 
a pair of fanned setae; segment III 
with six annulations, all with micro- 
trichia. 


Head constricted below eyes and 
longer than wide. Eyes with four large 
round facets bulging at sides of head. 
Head with four pairs of dorsal setae, 
H2 fanned. Pronotum with seven pairs 
of expanded setae. Mesonotum with 
seven pairs and metanotum with four 
pairs of fanned setae. Abdomen with 
two pairs of expanded setae on tergite 
I and three pairs on tergites II-VIII; 
segment IX with three pairs of dorsal 
setae, Al and A2 both being fanned 
and A3 pointed and reduced. Segment 
X with three pairs of dorsal setae, Al 
fanned. 


Major dorsal body setae all mod- 
erately to widely fanned and mod- 
erately long. Cuticle with fine stippling 
probably resulting from fine micro- 
trichia. Abdominal segment IX lacking 
a posterior comb. 

Larva I.—Cuticle yellow without 
hypodermal pigmentation. Eyes red. 
Antennae six segmented; segments II 
and III with weakly expanded setae. 


Chaetotaxy similar to that of larva 


VANCE: LARVAE OF THE SERICOTHRIPINI 


203 


II, but posteroangular setae lacking, 
and setae shorter and less expanded. 
Cuticular sculpture similar to that of 
larva IH, but fainter. Abdominal seg- 
ment IX lacking a posterior comb. 


Diagnosis.—This description of the 
genus Zonothrips is based on a de- 
scription and illustration by Priesner 
(19262) of Z. karnyi. Zonothrips can 
be distinguished easily from all other 
thripine genera except Sericothrips by 
the widely fanned, dorsal body setae. 
Differentiation of Zonothrips and Seri- 
cothrips is more difficult. Stannard 
(1968 ) reported the adults of these two 
genera as being similar, only separated 
by the number of antennal segments 
and the placement of abdominal sternal 
setae. The only way of separating the 
two genera at the present time is by 
considering host-plant data and as- 
sociated adults. 


Zonothrips osmundae 
Crawford, J.C. (1941) 


No larvae of this genus and species 
were available for study. Adults were 
collected in Illinois at Volo Bog, Lake 
County, from September to October 
on and around cinnamon fern (Os- 
munda cinnamomea) by L. J. Stannard, 
Jr., and are deposited at the Illinois 
Natural History Survey. 


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INDEX 


A 
Abdominal segments, 149, 167 
Abdominal spiracles, 167 
Adult characters of Sericothrips variabilis, 
153 
Aeolothripidae, 148, 157, 158, 160, 161, 
169-171 
Aeolothrips, 152, 159, 160, 168, 170, 171 
bicolor, 171 
fasciatus, 155, 171 
kuwanai, 155 
vittipennis, 171, 172, 174, 181, 186 
African violet, 173 
AGA (preserving solution), 146 


168, 


Anaphothripini, 148, 149, 157, 160, 161, 162, 
168, 169, 173, 175-176, 178 
Anaphothrips, 149, 159, 160, 169, 175, 176, 


178 
secticornis, 172, 175, 176, 177, 178, 
186 
Ankothrips, 170 
Antenna, 147, 148, 156, 158, 161, 162, 
172, 174, 177 
annulations, 148, 156, 158, 161, 167 
microtrichia, 148, 156, 158, 161, 162, 167 
sense cones, 148, 167 
Aptinothrips, 159, 160, 169, 176 
rufus, 176 
Arisaema (see jack-in-the-pulpit) 
dracontium, 178 


B 
Bald cypress, 187, 190 
Baliothrips, 169, 176 
Banana, 178 
Baptisia (see false indigo) 
Barley, 175 
Basswood (see linden) 
Berlese collecting method, 178 
Black locust, 171, 192, 194 
Bouteloua eriopoda, 175 
Bregmatothrips, 176 


181, 


167, 


Cc 
Caliothrips, 159, 160, 168, 173, 178 
fasciatus, 150, 154, 155 
indicus, 173, 174, 181, 186 
Canada balsam, 146 
Celtis (see hackberry) 
Cercyothrips, 179 
Chaetanaphothrips, 159, 160, 169, 175, 176, 
178 
Character states, 
156, 158, 162 
Charassothrips, 179 
Chilothrips, 159, 160, 162, 169, 176 
pini, 176 
Chirothripini, 148, 149, 157, 160, 161, 162, 
166, 168, 173, 175, 178 
Chirothrips, 148, 159, 160, 162, 169, 175 
simplex, 174, 175, 181, 186 
Chrysopa californica, 155 
Cinnamon fern, 203 
Clover, 178, 197 
Collecting methods, 146 
Color of immatures, 147, 148, 165 
Comb on abdominal tergite IX, 149, 156, 
158, 161 
Cordyline feminalis, 173 
Cornus (see dogwood) 
Corrodentia, 157, 161 
Cotton, 178 
Cottonwood, 176 
Ctenothrips, 159, 160, 169, 176 
bridwelli, 176, 177, 189 
Culver’s root, 178 
Cuticle, 147, 148, 149, 156, 158, 162 
hypodermal pigmentation, 148, 156, 158, 
162, 165 
microtrichia, 149, 156, 161, 162, 165 
pustules, 149, 156, 158, 161, 162, 165 


primitive and derived, 


sclerotized areas, 148, 156, 158, 161, 162, 


165 
sculpturing, 147, 149, 156 


August, 1974 VANCE: LARVAE OF 


D 
Dendrothripini, 148, 157, 159, 160, 161, 162, 
163, 168, 173, 175, 176, 178, 179 
Dendrothrips, 159, 160, 168, 176, 180, 187 
ornatus, 172, 176, 177, 183, 186, 189 
Desmodium, 184 
Dogwood, 187, 188, 199 
Dorcadothrips, 169, 176 
Drepanothrips, 159, 160, 163, 164, 165, 168, 
178, 179-180, 187 
reuteri, 156, 163, 164, 165, 177, 180-182, 
183, 186, 189 
E 
Echinothripina, 163, 165, 166, 168, 178, 179 
Echinothrips, 159, 160, 163, 164, 165, 166, 
168, 179, 182-183 
americanus, 156, 165, 172, 177, 182, 183- 
184, 189 
subflavus, 165, 182, 184-185 
Echinothrips complex, 179 
Economic assessment of Sericothrips 
variabilis, 155-156 
Elderberry, 193, 200, 201 
Endopterygota, 150 
Enneothrips, 179 
Ensiferothrips, 179 
Exopterygota, 150 
Eye facets, 149, 161, 167 
F 
False indigo, 193, 195 
Femur, 167 
Frankliniella, 159, 160, 169, 176, 178 
fusca, 150, 178 
parvula, 178 
tritici, 154, 172, 178 
Frankliniella-Thrips-Taeniothrips complex, 
147, 163 
Franklinothrips, 159, 160, 168, 169, 170, 174 
G 
Gladiolus, 178 
Grape, 180, 182 
H 
Hackberry, 199 
Haplothrips verbasci, 150 
Head, 149, 167 
Heliothripinae, 148, 149, 157, 160, 161, 162, 
168, 171, 178, 175, 179 
Heliothripini, 157 
Heliothrips, 159, 160, 168, 173 
haemorrhoidalis, 154, 172, 173, 174, 181, 
186 
Hemlock, 185 
Hercinothrips, 159, 160, 168, 173 
femoralis, 154, 173, 174, 186 
Heterothripidae, 148, 157, 158, 160, 161, 168, 
171 
Heterothrips, 152, 159, 160, 168 
arisaemae, 171, 172, 174, 181, 186 
Hippodamia convergens, 155 
Holometabola, 150 
Hops, 195 
Hoyer’s mounting medium, 146 
Hydrangea, 184 


THE SERICOTHRIPINI 207 


I 
Immature stages, 149, 151-153, 170 
Impatiens (see jewelweed) 
Tridothrips, 169, 176 


J 
Jack-in-the-pulpit, 171 
Japanese mulberry, 176 
Jewelweed, 178, 184 
Juniperus virginiana (see red cedar) 


L 

Larva, 166, 167 
first instar, 149, 151-152, 167, 170 
second instar, 149, 152, 167, 170 

Leucothrips, 159, 160, 168, 176 
piercei, 176 

Limothrips, 159, 160, 161, 162, 169, 173, 175 
cerealium, 150, 174, 175, 181, 186 
denticornis, 172, 175 

Linden, 193, 201 

Liothrips vaneecki, 154 


M 
Marijuana, 176 
Mating, 154-155 
Melanthrips, 170, 171 
Merothripidae, 148, 157, 158, 160, 161, 168, 

169, 171 
Merothrips, 152, 159, 160, 168, 169, 171 
morgani, 171, 174, 181, 186 

Mesonotum, 167 
Mesothoracic spiracle, 167 
Metamorphosis, 149-150 
Metanotum, 167 
Microcephalothrips, 159, 160, 169, 176, 178 
Mirabilis nyctaginea (see wild four-o’clock ) 
Morphology, external of larva, 167 
Mounting methods, 146-147 


N 
Nymphaea (see water lily) 


oO 
Oats, 178 
Octothrips, 179 
Odontothrips, 169, 176 
Orius 
indicus, 155 
insidiosus, 155 
tristicolor, 155 
Osmunda cinnamomea (see cinnamon fern) 
Oxythrips, 159, 160, 162, 169, 176 
cannabensis, 176, 177, 181, 186 


P 
Parthenothrips, 159, 160, 168, 173 
dracaendae, 173 
Phlaeothripidae (see Tubulifera) 
Photoperiod, effect on development, 
152, 153-154 
Phylogeny 
of the Sericothripini, 163-166 
of the Thysanoptera, 157-163 
Phytoseiid mites, 155 
Pigmy cypress, 176 


151, 


208 


Pliesiopsothrips, 179 

Plesiothrips, 169, 176, 179 

Predators of Sericothrips variabilis, 155 

Prepupa, 149, 152-153, 166, 170 

Privet, 176 

Pronotum, 149, 167 

Prosopothrips, 176 

Pseudodendrothrips, 159, 160, 169, 176, 180 
mori, 176, 177, 189 

Ptelea (see water ash) 

Pteridothrips, 179 

Pupa, 149, 153, 166, 167, 170 
site of development, 151, 154 


R 
Rearing methods, 150-151 
Redbud, 176 
Red cedar, 187, 188 
“Remetabola,” 150 
Repositories, 146 
Rhaphidothrips, 169, 170, 176 


Ss 
Sambucus (see elderberry) 
Scirtodothrips, 179 
Scirtothripina, 162, 163, 165, 166, 168, 178, 
179 
Scirlothrips, 159, 160, 163, 164, 165, 168, 178, 
179, 180, 182, 185-187 
brevipennis, 163, 164, 165, 187-188, 190 
citri, 150, 154, 156 
longipennis, 156 
niveus, 163, 164, 165, 183, 187, 188 
taxodii, 163, 164, 165, 172, 177, 183, 187, 
188-190 
Scolothrips, 159, 160, 169, 176, 182 
pallidus, 178, 183, 189 
Sericopsothrips, 179 


Sericothripina, 162, 163, 164, 165, 168, 176, 


178-179 = 
Sericothripini, 148, 156, 157, 159, 160, 161, 
162, 163, 164, 165, 166, 168, 173, 175, 176, 
178, 179 
Sericothrips, 150, 159, 160, 162, 163, 164, 165, 
168, 170, 175, 178, 180, 182, 187, 190-193, 
203 
annulipes, 163, 164, 165, 166, 178, 183, 
189, 190, 191, 192, 193-194, 196, 200 
annulipes group, 162, 164, 165, 166, 175, 
196-197 
baptisiae, 163, 164, 165, 166, 191, 193, 
194-195 
beachae, 163, 164, 165, 166, 191, 193, 195, 
202 
campestris, 163, 164, 165, 189, 191, 193, 
195-196, 200 
cingulatus, 162, 163, 164, 165, 166, 178, 
190, 191, 192, 194, 196-197, 200 
langei, 163, 164, 165, 166, 167, 168, 191, 
193, 195, 197-198 
nubilipennis, 163, 164, 165, 166, 191, 193, 
198-199, 201 
pulchellus, iv, 163, 164, 165, 166, 190, 
191, 192, 194, 199-200 


ILtino1is NATURAL History Survey BULLETIN 


Vol. 31, Art. 5 


sambuci, 168, 164, 165, 166, 191, 193, 196, 
200-201 
liliae, 168, 164, 165, 166, 191, 193, 198, 
201 
tiliae group, 163, 164, 165, 166 
variabilis, 150-156, 163, 164, 165, 166, 
170, 172, 177, 191, 192, 193, 195, 202 
Setae, 149, 156, 158, 162, 165, 167 
anal, 167 
basal rings, 149, 156, 162, 165 
length, 149, 156, 162, 165 
ornateness, 149, 156, 158, 162, 165 
Sexing of immatures, 151, 152, 153 
Soybeans, 150-151, 154, 155, 171, 178, 202 
Spanish moss, 178 4 
Spines on abdominal tergite IX, 156, 158, 
161 
Sticky traps, 151, 154 


T 
Taeniothrips, 148, 159, 160, 169, 176 
inconsequens, 154 
nigricornis, 155 
simplex, 154, 177, 178, 183, 189 
Tanglefoot sticky traps, 151, 154 
Tarsus, 167 
Taxodium distichum (see bald cypress) 
Temperature, effect on development, 151, 
152, 153-154 
Tentorium, 157, 161 
Terebrantia, 149, 156, 157, 166-167 
Thripidae, 157, 158, 159, 160, 161, 162, 168, 
169, 171-173 
Thripinae, 161, 162, 163, 168, 173, 175 
Thripini, 147, 148, 157, 159, 160, 161, 163, 
168, 169, 173, 175, 176, 178, 179 
Thrips, 159, 160, 169, 176 
impar, 178 
physapus, 178 
tabaci, 150, 154, 178 
Tibia, 167 
Tilia (see linden) 
Timothy, 176 
Tsuga canadensis (see hemlock) 
Tubulifera, 149, 157, 158, 159, 160, 161, 166- 
167, 171, 174, 181, 186 


Vv 
Vetch, 197 
Vitis (see grape) 

WwW 
Wafer ash, iv, 192, 200 


Water lily, 193, 198 
Wild four-o’clock, 193, 196 


Y 
Yarrow, 178 


Zz 
Zonothrips, 159, 160, 165, 168, 179, 192, 202- 
203 
karnyi, 192, 203 
osmundae, 165, 203 
Zut Slide Ringing Compound, 147 


Some Publications of the ILLINOIS NATURAL HISTORY SURVEY — 


BULLETIN 


Volume 30, Article 7—A Comparative Study 
of Two Components of the Poinsettia Root 
Rot Complex. By Robert S. Perry. Au- 
gust, 1971. 35 p., index, 


Volume 30, Article 8—Dynamics of Condi- 
tion Parameters and Organ Measurements 
in Pheasants. By William L. Anderson. 
July, 1972. 44 p., index. \ 


Volume 31, Article 1—The Effects of Sup- 
plemental Feeding and Fall Drawdowns 
on the Largemouth Bass and Bluegills at 
Ridge Lake, Illinois. By George W. Ben- 
nett, H. Wickliffe Adkins, and William 
F. Childers. January, 1973. 28 p., index. 


Volume 31, Article 2.—The Reproductive 
Cycle of the Raccoon in Illinois. By Glen 
C. Sanderson and A. V. Nalbandov. July, 
1973. 57 p., index. 


Volume 31, Article 3—Nutritional Respon- 
ses of Pheasants to Corn, with Special 
Reference to High-Lysine Corn. By Ron- 
ald F. Labisky and William L. Anderson. 
July, 1973. 26 p., index. 

Volume 31, Article 4—An Urban Epiphy- 
totic of Phloem Necrosis and Dutch Elm 
Disease, 1944-1972. By J. Cedric Carter 
and Lucile Rogers Carter. May, 1974. 31 
p., index. 


BIOLOGICAL NOTES 

80.—Illinois Birds: Hirundinidae. By Rich- 
ard R. Graber, Jean W. Graber, and 
Ethelyn L. Kirk. August, 1972. 36 p. 


81.—Annotated Checklist of the Butterflies 
of Illinois, By Roderick R. Irwin and 
John C, Downey. May, 1973. 60 p. 

82.—Lactate Dehydrogenase Isozymes of 
Darters and the Inclusiveness of the 
Genus Percina. By Lawrence M. Page 
and Gregory S. Whitt. May, 1973. 7 p. 

83.—Illinois Birds: Laniidae. By Richard 
R. Graber, Jean W. Graber, and Ethelyn 
L. Kirk. June, 1973. 18 p. 


84.—Interactions of Intensive Cultures of 
Channel Catfish with Largemouth Bass in 
1-Acre Ponds. By D. Homer Buck, Rich- 
ard J. Baur, and C. Russell Rose. Febru- 
ary, 1974. 8 p. 


List of available publications mailed on request 


No charge is made for publications of the Intrno1s NaTurAL History Survey. A 
copy of most publications will be sent free to anyone requesting it until the supply bi 
low. Costly publications, more than one copy of a publication, and publications in 
supply are subjects for special correspondence. Such correspondence should ident 
writer and explain the use to be made of the publication or publications. 


Address orders and correspondence to the Chief, 
Illinois Natural History Survey 
Natural Resources Building, Urbana, Illinois 61801 


85.—The Literature of Arthropods 
ated with Soybeans. III. A Bibli 
of the Bean Leaf Beetle, Ceroton 
cata (Forster) and ©. rujicornis (0 
(Coleoptera: Chrysomelidae). By 
Nichols, M. Kogan, and G. P, Wal 
February, 1974. 16 p. 
86.—Illinois Birds: Tyrannidae. By 
ard R. Graber, Jean W. Grabe 
Ethelyn L. Kirk. February, 1974, 
87.—The Literature of Arthropods 
ated with Alfalfa. I. A Bibliog 
the Spotted Alfalfa Aphid, The 
maculata (Buckton) (Homoptera: 
dae). By D. W. Davis, M. P. Nichol: 
HE. J. Armbrust. February, 1974. 
88.—The Literature of Arthropods 
ated with Alfalfa, II, A Bibliogr 
the Sitona Species (Coleoptera 
lionidae). By W. P. Morrison, B 
M. P. Nichols, and E. J. Armbr 
ruary, 1974. 24 p. 
89.—The Life History of the Spottail 
er, Etheostoma squamiceps, in 
Illinois, and Ferguson Creek, 
By Lawrence M. Page. May, 197 
90.—A Bibliography of the North 
Rootworm, Diabrotica longicornis 
and the Western Corn Rootwo: 
brotica virgifera LeConte (Co! 
Chrysomelidae). By W. H. Luckn 
H. C. Chiang, E. E. Ortman, and M 
P, Nichols. April, 1974. 15 p. 


CIRCULAR 

46.—Illinois Trees: Their Diseases, 
Cedric Carter, June, 1964. (Thir 
ing, with alterations.) 96 p. 

47.— Illinois Trees and Shrubs: Theii 
Enemies. By L. L. English. Jul 
(Fifth printing, with revisions.) 91 

51.—Illinois Trees: Selection, Planti 
Care. By J. Cedric Carter, August, 
123 p. 

52.—Fertilizing and Watering Tre 
Dan—Neely and E. B. Himelick. — 
ber, 1971. (Third printing.) 20 p. 

53.—Dutch Elm Disease in Illinois. 
Cedric Carter. October, 1967. 19 p. 


4 


L |) = ILLINOIS 


ittural History Survey 
1 BULLETIN 


Root Infection 
of Woody Hosts with 
Verticillium albo-atrum 


NATURAL HISTORY SURVEY 
OCT Bgl 
LIBRARY 


TN ENT OF REGISTRATION AND EDUCATION 


TURAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 


| 33a] ) VOLUME 31, ARTICLE 6 


ILLINOIS 


atural History Survey 
BULLETIN 


Root Infection 
of Woody Hosts with 
Verticillium albo-atrum 


d L. Born 


OF ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


URAL HISTORY SURVEY DIVISION 
ANA, ILLINOIS 
VOLUME 31, ARTICLE 6 
AUGUST, 1974 


STATE OF ILLINOIS 
BOARD OF 


DEAN Barrincer, Ph.D., Chairman; THoMAS Park, 


DEPARTMENT OF REGISTRATION AND EDUCATION — 
NATURAL RESOURCES AND CONSERVATION 


Ph.D., Biology; L, L. Suoss, Ph.D., Geology; Herpert §, 
GuTowsky, Ph.D., Chemistry ; Ropert H. ANDERSON, B.S.C.E 


., Engineering ; CHARLES E, OuMsrED, Ph.D., Forestry ; 


W. L. Everirr, E.E., Ph.D., Representing the President of the University of Illinois; Euserr H, Hapuny, Ph.D., 
Representing the President of Southern Illinois University. 
NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 
SCIENTIFIC AND TECHNICAL STAFF Hi 
GEORGE SPRUGEL, JR., Ph.D., Chief 
Avice K, Apams, Secretary to the Chief 3 
Section of Economic Entomology 


Wittiam H. Luckmann, Ph.D., Entomologist and Head 

Wixuis N. Bruce, Ph.D., Entomologist 

Wayne L. Hows, Ph.D., Entomologist 

STEVENSON Moors, III, Ph.D., Entomologist, Extension 

Howarp B. Perry, Ph.D., Entomologist, Extension 

James E, APPLEBY, Ph.D., Associate Entomologist 

Epwarp J, ArmBRustT, Ph.D., Associate Entomologisl 

Marcos Kogan, Ph.D., Associate Entomologist 

JoserpH V. Mappox, Ph.D., Associate Entomologist 

Ronatp H. Meyer, Ph.D., Associate Entomologist 

Rogert D. PauscH, Ph.D., Associate Entomologist 

Raupew E. Securiest, Ph.D., Associate Entomologist 

Joun K. BouspmMan, M.S., Assistant Entomologist 

Georce L, Goprrey, Ph.D., Assistant Entomologist 

Witiiam G. Ruesink, Ph.D., Assistant Entomologist 

James R. SanBorn, Ph.D., Assistant Entomologist 

Douauas K. Sevu, Ph.D., Assistant Entomologist 

Joun L, WepserG, Ph.D., Assistant Entomologist 

CLARENCE E, WuitsE, B.S., Assistant Entomologist 

Keun S. Park, M.S., Assistant Chemist 

Sup E. Watkins, Supervisory Assistant 

DonaLp E, KuHLMAN, Ph.D., Assistant Professor, Exten- 
sion 

Roscoe RANDELL, Ph.D., Assistant Professor, Extension 

Tim Cooney, M.A., Assistant Specialist, Extension 

Kurr E. Reppora, M.S., Assistant Specialist 

Joun F. Watt, M.S., Assistant Specialist, Extension 

Jpan G. Wiuson, B.A., Supervisory Assistant 

DANIEL P. BarteL, Ph.D., Research Associate 

Martua P. Nicuous, M.S., Research Associate 

Ropert J. Barney, B.S., Research Assistunt 

Tzu-Suan Cuu, M.S., Research Assistant 

Srepuen D. Cowan, B.S., Research Assistant 

SrerpHeN K. Evranrp, B.S., Research Assistant 

BarBarRa J. Forp, M.A., Research Assistant 

Raymonp A, Korex, M.Mus., Research Assistant 

Rose ANN Meccout, B.S., Research Assistant 

BarBarRA E. Perprson, B.S., Research Assistant 

KerurAH ReINBOLD, M.S., Research Assistant 

STEPHEN Roserts, B.S., Junior Professional Scientist - 

Joun T. SHaw, B.S., Junior Professional Scientist 

LowE.u Davis, Lechnical Assistant 

CHARLES G. Heum,-M.S., Technical Assistant 

LinpA IsENHOWER, Technical Assistant 

Lu-pina Len, M.S., Technical Assistant 


Section of Botany and Plant Pathology 

Ropert A. Evers, Ph.D., Botanist 

Evucene B. Himenick, Ph.D., Plant Pathologist 

R. Dan Newuy, Ph.D., Plant Pathologist 

D. F, ScHorneweiss, Ph.D., Plant Pathologist 

J. LeuanD Oran, Ph.D., Associate Mycologist 

Water Hartstirn, Ph.D., Assistant Plant Pathologist 
Berry S. Neuson, Junior Professional Scientist 

Gene E. Rew, Technical Assistant 


Section of Aquatic Biology 

D. Homer Buck, Ph.D., Aquatic Biologist 

WituiAM F. Cuinpers, Ph.D., Aquatic Biologist 

R. Weipon Larimore, Ph.D., Aquatic Biologist 

Roser C. Hintrpran, Ph.D., Biochemist 

ALLISON BRIGHAM, Ph.D., Assistant Aquatic Biologist 
Warren U. Bricuam, Ph.D., Assistant Aquatic Biologist 
RicHarp E, Sparks, Ph.D., Assistant Aquatic Biologist 
JoHN TRANQUILLI, M.S., Assistant Aquatic Biologist 
DonaLp W. Durrorp, M.S., Junior Professional Scientist 
Mary Frances Martin, Junior Professional Scientist 
Joun M. McNurney, M.S., Junior Professional Scientist 
Tep W. Storck, Ph.D., Junior Professional Scientist 


CONSULTANTS AND RESEARCH AFFILIATES: 


Systematic Enromouocy, Roperick R. Irwin, Chicago, Illi- 


‘Yom Huu, M.S., Research Assistant 

RicHarp Kocuer, B.S., Research Assistant 

RoBer? Moran, M.S., Research Assistant : 
C. Russet Ross, Field Assistant A 


RicHarD J. Baur, M.S., Research Assistant : 
{ 


Section of Faunistic Surveys and 
Insect Identification 


Puitip W, SmitH, Ph.D., Taxonomist and Head ‘= 
Waxvace E, LaBerce, Ph.D., Taxonomist 

Minton W. Sanperson, Ph.D., Taxonomist 

Lewis J. STANNARD, JR., Ph.D., Taxonomist 2 
Larry M. Pas, Ph.D., Assistant Tazonomist ‘ 
Joun D. Unzicker, Ph.D., Assistant Taxonomist 
Donatp W. Wess, M.S., Assistant Taxonomist 

BERNICE P. SWEENEY, Junior Professional Scientist 


Section of Wildlife Research 


GLEN C, Sanperson, Ph.D., Wildlife Specialist and Head 

Prank C, BELLROSE, B.S., ” Wildlife Specialist 

Jean W. Graper, Ph.D., Wildlife Specialist { 

Ricwarp R. Grazer, Ph.D., Wildlife Specialist 

Harotp C, Hanson, Ph.D., Wildlife Specialist 

Ronaup F, Lapisky, Ph.D., Wildlife Specialist 

WiuiiamM L. ANDERSON, M,A., Associate Wildlife Special- 
ist 

W. W. Cocuran, JR., B.S., Associate Wildlife Specialist 

Winuiam R. EDWARDS, M. g., Associate Wildlife Special-— 4 
ist 

G. Buairk JosELYN, M.S., Associate Wildlife Specialist 

CHARLES M. Nixon, M.S., Associate Wildlife Specialist 

KENNETH E. SmirH, Ph.D., Associate Chemist 

Bea L. WESTEMEIER, M. S., Associate Wildlife Spe- 
cialist 5 

STEPHEN P. Havers, M.S., Assistant Wildlife Specialist — 

Davin R. Vance, M.S., Assistant Wildlife Specialist 

Ronautp E. Duzan, Junior Professional Scientist 

HeL.en C. Scuuutz, M.A., Junior Professional Scientist — 

ELEANORE WixLson, Junior Professional Scientist { 

SHARON FRADENBURGH, B.A., Laboratory Technician 

Rosert D, Crompton, Field Assistant : 

James W. Seers, Laboratory Assistant 


Section of Administrative Services 
Rosert O, Watson, B.S., Administrator and Head 


Supporting Services 

Jerry McNear, Maintenance Supervisor ] 
Witma G. Dinuman, Property Control and Trust Ac- 

counts 
Parry L, Duzan, Technical Assistant 
Rorert O. Evuis, Assistant for Operations q 
Larry D. Gross, Maintenance Supervisor 
Luoyp E. HurrmMan, Stockroom Manager 
J. Witu1am Lusk, Mailing and Distribution Services 
Metvin-E. ScHwartz, Financial Records 
James E, SERGENT, Greenhouse Superintendent 


/ 
Publications and Public Relations 
Owen I’, GuissenDoRF, M.S., Technical Editor 
Rospert M. ZewaDskI, M. S., ‘Associate Technical Editor — 
SuiRLEY McCLELLAN, Assistant Technical Editor | 
LAWRENCE S. FARLow, Technical Photographer ; 
Luioyp LeMereg, Technical Illustrator ’ 
Technical Library 

Doris F. Dopps, M.S.L.S., Technical Librarian 

Doris L, SUBLETTE, MS.LS., Assistant Technical L 
brarian 


nois; WILDLIFE RESEARCH, WILLARD D. Kuimstra, Ph.D., Professor of Zoology and Director of Cooperative Wild- 
life "Research, Southern Illinois University ; PARASITOLOGY, Norman D. Levine, Ph.D., Professor of Veterinary 
Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, University of 
Illinois ; ENtomoLocy, Ropert L. Mutcaur, Ph.D., Professor of Zoology and of Entomology and Head of the De- 
partment of Zoology, University of Illinois ; and GILBERT P. WALpBAUER, Ph.D., Professor of Entomology, Uni-— 
oer of Illinois ; Statistics, Horace W. Norton, Ph.D., Professor of Statistical Design and Analysis, University 
0. MNO, 


CONTENTS 


SR IARVATETUGEINEPIN'ES) oe sea iat eRe a eN hose tena peed e oes han aeectee, eon eateeneee ce NA 209 
(Ets a Tere dtr 4 BUONO ity eee a a, ee me a Ore een a Ceeeeerre 210 
CopE OF VERTICILLIUM ALBO-ATRUM ISOLATES .............-22-2---2-0c2---eeeeeeeeeeeeeee cece 212 
iiatonia istandeniethodsmsr. sess ens LE Ne Ne) et ee ee 912 
RELATIONSHIP OF Root WounpDs & AGE OF WOUNDS ON INFECTION ............... 213 
Materials and Methods 
Type of wound ............ 
Eee SEA eT eh Me 2 a Bis aoe Cee ey eee aT ae PS 
"ECTEULES, co dcmecans chose tee Bie ne ser eee ts Se See IR Se Pe Ste As eee eS ANE A Se 
Sey PEO HRV GUC yRemen mth ae eats 8 RES Se he tet eS ee ee 
ENC CRO le VOULICeememe. ete cometeere.. NT Ae RT ie BU ee 
Discussion and Conclusions ................. ids thee NE Se ES PC Ree Eas Ry 214 


PENETRATION AND DEVELOPMENT OF Y. ALBO-ATRUM IN Roots oF Woopy 
Hosts ..... oh ae Boe sea EP eA ee Re ROR A a Sag ee Tee ee ey 


RIPReTTAl SHAN OMNLCCHOUS ie ceo ccc Soee en ne eee Ee ee 

IEE STIUL Spee ees ee eee ae a sre eee ae aero ES eles 
Fungus growth on root surface 
TECOLOXE (ih ae) YS) AVS) Wipe 10) 0 Nee toe fe tee Sr en ne eran an ee er Pee 
Penetration in root hair region 
Penetration in area of lateral root formation —...........-.-..- 217 
Cortical invasion—Susceptible hosts ..........<..-.-.-.------------2--0-0---00000e--* 218 
Cortical invasion—Resistant hosts -................222..22.-202--2-222-220 22 eee 218 
Penetration of vascular region of susceptible hosts -...................--...... 218 
Penetration of vascular region of resistant hosts -......................--.-- 221 

Diseesstom eiel (Cone rei e a ee 224 


Errecr oF Root INFECTION ON GROWTH RESPONSE OF REDBUD & GREEN 
PRES RDIATN CS ee ee, eee ees Sal a wiht. 


Materials and Methods 
[RveS3bULES Soa cae rae) A es SE Se, A a ee 
RS VATA | COLES Pee eee ee rece cee Mee Se ARS ee Bee 
TDUAy REEL, Sea Meet oc et es cele ca ean eee 
TLE de TOUT G ETE a a-tee dost tees Bi cia eee epee pote aay aaa ean te aR ere OE 
(SiS: 1 aYEN Gedo esa ea ea eR a ee eee eee 
Nitrogen content -................ 
Water content of leaves 
TL Coen see Sr a i A cee 
Discussion and Conclusions 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It is 
@ contribution from the Section of Botany and Plant Pathology of the Illinois Natural 
History Survey. 


(58200—4,000—S8-74) 


LE 


Errect OF TEMPERATURE & HEAT TREATING ON DEVELOPMENT OF V. ALBO- 
ATRUM IN ROOTS c.0200:00-000 So Ee Or 2 
Materials and Methods .......2::..:---2 002.2: Ss. ee 
BReSta lt: cc. secccccoecnesceccee-venccdcnnseense ee oeeee see ee er 
Effect of temperature: :...:--...:.-52 cc: sesepeseettenee oor 

Effect of heat treating of soil -.---.2.2-.--ccecceecesecsene--ooo se eee 
Discussion and Conclusions: 2.22. - 12-2 eee -<csts-oorcee 


EVALUATION OF SYSTEMIC FUNGICIDES AGAINST V, ALBO-ATRUM ........---------------- 
Materials and Methods) o...25..0o.ce--cntecee coerce ce ene see 
Laboratory Studies a2. .2--<c-:co:--:sccsttecaceen scene! onsen a 
Greenhouse: studies” 22.20.0000 ee 
Soil drenches «.....2:..-.#220.5- eA eee eee 

Foliar treatments 

Root treatments .... 

Restallts: ic 25 eae Se er Bs nner ire ee 
Symptoms) 2202 2S Ne ee eee eee 
Laboratory studies 
Greenhouse studies .2.....22......0000 ee 
Soil drenches) 2..:20...08.24.00) ste 

Foliar treatments: .....0.00.0..- haces te 

Root treatments .....2.-.0 20-0. - thee le 
Discussion and CGomelusiOns io. o2 ecco ssscc ce ees 


| 
| 


Root Infection of Woody Hosts 


: Verticillium albo-atrum 


VERTICILLIUM WILT is a plant 
disease caused by the fungus Verticil- 
‘lium albo-atrum Reinke and Berthold. 
| This pathogen is peculiar in that it does 
\not confine its attacks to one host, or 
a few closely related hosts, as is so 
‘frequently the case with most other 
/pathogenic fungi; it attacks a large 
/number of widely unrelated plants, 
jmany of which are of economic im- 
portance. The disease does not often 
Joccur in forest stands, but it is be- 
|coming increasingly prevalent in plant- 
jings of ornamental trees and shrubs, 
particularly in temperate regions of the 
world. 

Symptoms of Verticillium wilt on 
woody hosts are variable and often 
difficult to recognize. Usually the first 
visual symptom is sudden wilting of 
foliage on one or several twigs on a 
branch. A yellowing of foliage some- 
times precedes wilting. Most plants 
exhibit leaf symptoms in early July, 
but some trees may first show symp- 
toms in early spring or late fall. Leaves 
on affected ash species may drop while 
still green and before noticeable yel- 
lowing or wilting has occurred. 


Other symptoms suggesting Verticil- 
lium wilt are decline in current twig 
growth, stunting, and dieback of indi- 
vidual twigs and branches. Occasion- 
ally trees such as maple and tulip tree 
develop elongated dead areas of bark 
on the diseased branches or trunk. 
Water-soaked areas sometimes develop 
under the killed bark. 


Trees that develop a limited amount 
of branch wilt during the summer may 
show additional wilt and dieback the 
following year, and others may recover 
and not wilt in succeeding years. Trees 
that have extensive wilt throughout the 
crown usually die before the end of the 
summer. 


Gerald L. Born 


The present study initiated in 1970 
and completed in 1972 deals with (1) 
the influence of root wounds and age 
of wounds on infection, (2) penetra- 
tion and development of the fungus in 
susceptible and resistant woody hosts, 
(3) analysis of the growth response of 
young tree seedlings after root infec- 
tion, (4) the influence of temperature 
and heat treating of soil on develop- 
ment of V. albo-atrum in excised roots, 
and (5) laboratory and greenhouse 
evaluation of fungicides against V. 
albo-atrum. 


This report is adapted from a thesis 
submitted to the University of Illinois 
in partial fulfillment of requirements 
for the degree of Doctor of Philosophy 
in Plant Pathology. 


ACKNOWLEDGMENTS 


Much of the work on this project was 
conducted with the cooperation of the 
Illinois Natural History Survey staff 
and through the use of Survey facilities. 
The study was carried out under the 
supervision of Dr. J. C. Carter, Head 
of the Section of Botany and Plant 
Pathology at the Survey, to whom I 
express my greatest appreciation. 

Others on the Natural History Survey 
staff who deserve special recognition 
for helping to make this a successful 
project are Dr. E. B. Himelick, Dr. 
Dan Neely, Dr. J. L. Forsberg, and 
Dr. J. L. Crane. Dr. Crane was es- 
pecially helpful in his assistance in 
microscopic photography. 

I gratefully acknowledge the assis- 
tance given by Dr. Richard E. Ford, 
Professor of Plant Pathology and Head 
of Department, University of Illinois, 
and Mrs. Betty Nelson and Mrs. Bar- 
bara Little of the Survey staff in prep- 
aration of the original thesis report, 


209 


210 


and by O. F. Glissendorf, Survey Tech- 
nical Editor, who edited the manuscript 
for this Bulletin article. 


LITERATURE REVIEW 


The first reference to a wilt disease 
was made by Reinke & Berthold (1879). 
They isolated a fungus from potato 
plants with the Krauselkrankheit dis- 
ease which they named Verticillium 
albo-atrum. Their investigations were 
not appreciated until 30 years later 
when Blattrollkrankheit and Krausel- 
krankheit were causing severe losses in 
the potato fields of Europe. 


Van Hook (1904) described a typi- 
cal case of wilt in ginseng (Panax 
quinquifolium L.) which he attributed 
to Acrostalagmus albus Preuss. How- 
ever, this name is synonymous with the 
earlier name Verticillium which was 
established by Nees von Esenbeck 
(1816). Corda (1838) did not describe 
the genus Acrostalagmus until 1838. 


Klebahn (1913) isolated a Verticil- 
lium from dahlia plants which he con- 
sidered distinct from V. albo-atrum, 
and he named this fungus V. dahliae 
Kleb. Since 1913 the relationship be- 
tween V. albo-atrum and V. dahliae has 
been the subject of much controversy. 
Many investigators have disagreed in 
their interpretations of the drawings 
and descriptions found in previous re- 
ports. Wollenweber (1929), Rudolph 
(1931), Presley (1941), Wilhelm & 
Taylor (1965), and Van den Ende 
(1958) maintained that the fungi that 
produce sclerotia and resting mycelium 
are members of a large variable species. 
Others, e.g., Klebahn (1913), Van der 
Meer (1925), Isaac (1949), and Smith 
& Walker (1930), have preferred to 
treat them as separate species. 


In 1957 Verticillium wilt was re- 
ported as affecting plants in at least 
18 orders, 38 families, 98 genera, and 
137 species in the temperate climates 
of the world (Caroselli 1957). 


Numerous papers in the past have 


Inurnois NAaturAL History SuRVEY BULLETIN 


Vol. 31, Art. 63 


dealt with factors that influence the - 
incidence of Verticillium wilt. Nutri- 
tion, soil type, soil moisture, soil and | 
air temperature, and light have all been) 
shown to have an effect on the inci 


Wilhelm TEU 


Although many workers have men- 
tioned wounds as a source of entry by 


the host. Selman & Pegg (1057) failed d 
to show any appreciable increase in” 


damage but, in these cases, the “un- 
damaged” control plants had been 
transplanted. Under normal conditions » 
of root growth in soil, where the inocu-~ 
lum potential of the fungus may be 
expected to be relatively low, it seems 
probable that entry into the xylem ves-_ 
sels occurs largely through wounds. _ 


Armstrong & Armstrong (1958) 


hosts were cut prior to inoculation with 
Fusarium spp. Also, the average num- 
ber of days for wilt to occur signifi- - 
cantly decreased when roots were cut 
immediately prior to inoculation. Fu | 
ton (1952) showed that more infection 
occurred when the canes or roots O& 
raspberry were injured prior to inocu- 


lation. 4 


: 
: 


Aug., 1974 


dolph (1931) reported that the fungus 
was found only in the xylem in the 
early wilt stage, and later invaded the 
pith, cambium, and cortex in the ad- 
vanced wilt stages. McWhorter (1962), 
working with Pelargonium infected 
with V. albo-atrum, found only traces 
of mycelium in tissues that had con- 
siderable discoloration. He rarely found 
large amounts of mycelium in diseased 
tissue. Therefore, the amount of mycelia 
in the vessels is not always indicative 
of the severity of wilt. 


Talboys (1958) observed that acute 
symptoms of hop wilt were associ- 
ated with extensive development of 
mycelium in the xylem vessels but 
sparse production of tyloses; con- 
versely, mild symptoms were associated 
with the development of limited my- 
celium but abundant tyloses in the ves- 
sels. Talboys (1964) suggested a simple 
explanation of the inverse correlation 
he had found between density of 
mycelium and frequency of tyloses in 
infected xylem vessels of the hop plant 
by postulating that a low concentra- 
tion of fungal metabolites in the xylem 
stimulates the formation of tyloses but 
that a high concentration inhibits for- 
mation. 


Until recently, spread of the fungus 
throughout the plant has received little 
attention. Sewell & Wilson (1964) con- 
cluded that V. albo-atrum conidia are 
transported in xylem sap of hops and 
occasionally they become lodged in 
vessels where they germinate and pro- 
duce more conidia. In cotton and 
tomatoes, conidia may spread through- 
out the plant in 12 hours to 6 days 
following inoculation (Garber 1957; 
Green 1954). 


Many earlier workers noted that the 
hyphae are very slender and reduced 
in diameter at the point where they 
pass through the cell walls, but once 
through they swell to a much greater 
size (Garber 1957; Garber & Houston 
1966; Klebahn 1913; Reinke & Berthold 
1879). In the vascular system, the 


Born: Root Inrecrion wit V. albo-atrum 


211 


fungus moves from one vessel element 
to another through pits (Garber 1957; 
Garber & Houston 1966; Green 1954). 
There is an apparent inability of the 
mycelium to penetrate new cellular 
growth lateral to the invaded cells as 
rapidly as the new cells are formed 
(Green 1954). 


Klebahn (1913) and Rankin (1914) 
reported microsclerotia in the vessels 
of infected plants. Talboys (1958) ob- 
served that penetration of vascular 
tissue of hop by V. albo-atrum de- 
pended on the amount of suberin in 
the endodermal cell walls. Garber & 
Houston (1966) observed gum-like de- 
posits in tolerant cotton plants which 
impeded the fungus from penetrating 
the vascular element. They reported 
that the splitting apart of cells was a 
mechanical process and not enyzmatic 
although they observed enzymatic ac- 
tion on the middle lamella of cell walls 
when the inoculum potential was high. 


Symptom appearance is variable, re- 
quiring days to many weeks after in- 
fection for expression. Yellowing of 
foliage and sudden wilting are usually 
the first visual symptoms. General 
stunting accompanied by shortening of 
the internodes may accompany wilt. 
Young tomato plants infected with V. 
albo-atrum may show neither leaf yel- 
lowing nor wilting in the initial stages, 
but only a stunting of the whole plant 
(Selman & Pegg 1957). 


Selman & Pegg (1957) found that 8 
weeks after inoculation the dry weights 
of tomato leaves, stems, and roots were 
decreased by 72, 70, and 65 percent 
respectively. Of the growth charac- 
teristics studied, leaf area was most 
reduced by infection and this was due 
to a failure of the leaves to expand 
rather than to a reduction in leaf pro- 
duction. 

After infection, symptom deyelop- 
ment, and necrosis, the fungus may 
overwinter within the plant as micro- 
sclerotia. Benken & Khakimoy (1964) 
observed abundant microsclerotia of V. 


212 
albo-atrum in veins and petioles of 
overwintering cotton leaves. The fungus 
spread unchecked in the field within 
the necrotic tissues of infected cotton 
seedlings and sporulated freely over 
the surface of the stems for a short 
distance above ground level} eventu- 
ally forming numerous microsclerotia 
in stems and roots. Nadakavukaren 
(1965) observed that V. albo-atrum 
microsclerotia survived best at low 
temperatures and high moisture levels. 
Heale & Isaac (1963) reported that rest- 
ing mycelium remained viable for 9 
months in pieces of necrotic lucerne 
buried 12 inches (30 cm) in soil. 
Brinkerhoff (1969) observed that micro- 
sclerotia were elongated in leaves incu- 
bated at 28 to 30 C and round in leaves 
incubated at 18 C. V. albo-atrum sur- 
vived for relatively long periods in cot- 
ton tissue, and infested debris con- 
stituted a ready source of inoculum 
when incorporated into either sterile 
or nonsterile soil. Evans et al. (1966) 
suggested that further colonization by 
V. albo-atrum was arrested when cotton 
plants were plowed under prior to 
microsclerotial formation in the tissues. 


Many recent papers have shown the 
value of systemic fungicides for the 
control of vascular wilts. Most of the 
work has been done with Benlate 
(benomyl) and thiabendazole (TBZ). 
Schreiber et al. (1971) found that 
benomyl was taken up equally well 
when either applied as a drench or in- 
corporated directly into the potting 
media. The planting medium affected 
the concentration as well as the rate 
of accumulation of benomyl. Highest 
levels of accumulation of the fungitoxi- 
cant were in seedlings grown in media 
that had the lowest content of organic 
matter and the highest pH. Heat 
sterilization of soil prior to benomyl 
treatment resulted in greater accumula- 
tion of benomyl] in elm seedlings than 
when the plants were grown in non- 
sterile soil. 


Erwin et al. (1971) found that the 


Intrnois NAtruRAL History SURVEY BULLETIN 


Vol. 31, Art. 6 


addition of thiabendazole to soil re- 
duced the incidence and severity of 
cotton wilt in plants subsequently inoc- 
ulated with V. albo-atrum. Rawlins & 
Booth (1968) reported that the addi- 
tion of surfactant Tween 20 increased 
the effectiveness of benomyl and thi- 
abendazole against V. albo-atrum, prob- — 

ably by increased absorption of the 
fungicide by the roots. Erwin et al. — 
(1968) found that thiabendazole not — 
only translocates from the roots to the 
stems of cotton plants but also can be 
detected in the bark. They concluded — 
that thiabendazole diffused laterally — 
from the xylem to the bark. 3 


Soil treatment, or seedling root dips — 
with difolatan (Bankuti 1964) gave © 
good control of Fusarium oxysporum — 
f. sp. lycopersici and V. albo-atrum on 
tomatoes in greenhouse and field tests. 
Complete protection against V. albo- — 
atrum was provided for seedlings — 
planted up to 140 days in soil treated ~ 
with difolatan. 


Applying systemic fungicides to the 
foliage and allowing the chemical to 
be translocated downward may be the 
method used in the future. However, 
this method presents many problems. — 
Many fungicides, such as benomyl, are — 
extremely insoluble in water. Hock — 
(personal communication) has been — 
able to solubilize benomy! using inor- 
ganic acids, heat, and constant stirring. 
Buchenauer & Erwin (1971) found that 
benomyl and thiabendazole induced ~ 
curative effects when sprayed on inocu- 
lated cotton plants that showed initial — 
symptoms of Verticillium wilt. Both 
fungicides were detected by bioassay — 
and chemical analysis in xylem tissue — 
and in nontreated stems and leaves 
above the place of application. 


CODE OF VERTICILLIUM 
ALBO-ATRUM ISOLATES 


MATERIALS AND METHODS 


All isolates used throughout this 
study were obtained from actively wilt- 


Aug., 1974 


ing hosts in Illinois. They were main- 
tained on freshly prepared potato dex- 
trose agar (PDA) tube slants and 
transferred periodically. An isolate used 
to inoculate a particular species was 
obtained earlier from another of the 
same species. Resistant species were 
inoculated with a mixture of all isolates. 
Below are the code numbers used in 
this study to identify each isolate. Also, 
the host, date of isolation, and location 
of host plant in Illinois are given for 
each isolate. 


Code Host Date Place 

1 Sugar maple 1969 Urbana 
2 Russian olive 1956 Wheaton 
3 Redbud 1960 Urbana 
4 Green ash 1961 Decatur 


RELATIONSHIP OF 
ROOT WOUNDS & AGE OF 
WOUNDS ON INFECTION 


MATERIALS AND METHODS 


Two hundred twenty each of 2-year- 
old bare-rooted sugar maples (Acer 
saccharum Marsh.) and redbud (Cercis 
canadensis L.) seedlings were selected 
as test plants. The plants were break- 

ing dormancy when received from a 
commercial nursery. The average height 
was 45 to 60 cm. The roots of each 
plant were washed with tap water and 
rinsed with distilled water prior to 
planting. The plants were potted in a 
medium-grade perlite and fertilized bi- 
weekly with a balanced liquid fertilizer. 

Isolate 1 was used to inoculate sugar 
maple and Isolate 2 was used to inocu- 
late redbud. 


Type of Wound 

Two weeks after potting, 100 plants 
were removed from the perlite and 
treated. Treatments immediately pre- 
ceding inoculation included: (1) no 
wound, (2) abrasion, (3) puncture, 
(4) vascular incision. Wounds 
were made on the primary root ap- 
|proximately 5 cm below the ground 
|line. With the abrasion-type wound, 


Born: Root Inrecrion wiru V. albo-atrum 


213 


the root surface was injured by rubbing 
moist 400 grade carborundum against 
the root surface. Puncture wounds were 
made by forcing a balsam wood block, 
in which five pins were embedded, 
against the root. This produced pin 
prick wounds 3 mm deep into the root. 
The vascular incisions were made by 
cutting a V-shaped wedge approxi- 
mately 0.5 cm deep into the root. 
When no wound was made, a mycelial 
disc was placed against the root sur- 
face. 


All treated plants were inoculated 
with a mycelial disc and the wound 
area covered with vinyl grafting tape 
to prevent moisture loss. The control 
plants were treated identically except 
that a sterile agar disc was placed on 
the wounded area and covered with 
grafting tape. Twenty plants of each 
species were used for each treatment. 


After 30 days, all plants were re- 
moved from the pots and _ isolations 
were attempted from the plant roots 
and stems. 


Age of Wound 

In an additional experiment 120 
plants of each species were tested to 
determine the importance of wound 
age on infection. Two weeks after 
potting, the plants were gently removed 
from the potting medium and V-shaped 
wounds were made on each plant ap- 
proximately 5.0 cm below the soil line 
on all plants. Fifteen plants were 
inoculated with a mycelial disc im- 
mediately after wounding and the 
wounds were covered with vinyl graft- 
ing tape. All other wounds were 
wrapped with vinyl grafting tape and 
the wounded plants replaced in perlite. 
At intervals of 1, 2, 4, 8, 16, and 32 
days, 15 plants were removed from the 
potting mixture, inoculated at the 
wound site, rewrapped with grafting 
tape, and planted back in perlite. Thirty 
days after each inoculation date, the 
plants were removed from the pots and 
isolations were made from the roots 
and stems of each plant. 


214 


RESULTS 


Type of Wound 

No infection occurred on unwounded 
roots. Root wounds were a prerequisite 
for fungus entry into the plant (Table 
1). Any disruption of the periderm on 
the older roots which allowed the fun- 
gus to by-pass these tissues was suitable 
to fungal entry. The percentages of 
infection for abrasive, puncture, and 
vascular wounds were 75, 80, and 85 
respectively on redbud, and 50, 55, 
and 80 on sugar maple. The most effi- 
cient wound on both hosts was a vascu- 
lar wound which placed the pathogen 
in direct contact with the vessel mem- 
bers. 


Age of Wound 

Root wounds remained as infection 
courts up to 32 days on redbud and 16 
days on sugar maple seedlings (Table 
2). As the age of the wound increased 
the number of plants infected through 
wounds decreased. Only 13 percent 
of the redbud plants became infected 
when inoculated at wound sites that 
were 32 days old and no infection oc- 


curred through wound sites 32 days old 


on sugar maple. 


Thirty two-day-old wounds had sev- 
eral layers of dead cells which were oc- 


Table 1.—The effect of root wounds on number of redbud and sugar maple plant 


infected with Verticillium albo-atrum. 


Inurvors NaturAL History SuRVEY BULLETIN 


Vol. 31, Art. 6 


cluded with heavily pigmented ma-— 
terials. This condition was a barrier 
against penetration by the fungus. 
Many vessel members adjacent to- 
wounds were occluded with tyloses and 
wound reaction materials. Callus was 
beginning to form at the margins of 
the wound after 32 days. 

Wounds were not made on other 
areas of the root. Therefore, location” 
of the wounds may have some signifi- 
cance because wounds made on older 
secondary tissue may require a longer 
time for initiation of repair tissue. 
Younger tissue, ie., at the root tip or 
lateral roots, may heal faster and reduce 
the time a wound remains as an infec- 
tion court. 


DISCUSSION AND CONCLUSIONS 


The periderm consists of the phello- 
derm, phellem, and phellogen which 
completely surrounds the vascular cyl- 
inder of woody plant roots. The cells 
of the phelloderm are parenchyma and 
remain alive and active. The cells of 
the phellem become suberized, which 
renders them virtually waterproof, 
and at maturity they die, forming a 
rather impervious, protective layer 
around the outside of the root. 


The fungus gains entrance through 


Type of Wound" 


Redbud — 2 years old 
No wounds 
Abrasion 
Puncture 
Vascular incision 
Noninoculated (controls) 


Sugar maple — 2 years old 
No wounds 
Abrasion 
Puncture 
Vascular incision 
Noninoculated (controls) 


Number of Plants Infected 


Roots 
and Stems 


Roots 
Only 


“Twenty plants were used per treatment. 


Aug., 1974 


Table 2.—The effect of age of root wounds 
prior to inoculation with Verticillium albo- 
atrum. 


Age of Wound 
at Inoculation* 


Number of 
Plants Infected 


Redbud — 2 years old 
i Immediate 
1 day 
2 days 
4 days 
8 days 
16 days 
32 days 
Noninoculated (controls) 


= 


CNNWRAD SO 


Sugar maple — 2 years old 


Immediate 11 
1 day 10 
2 days 6 
4 days 3 
8 days 3 

16 days al 

32 days 0 

Noninoculated (controls) 0 


a Fifteen plants were used for each treatment. 


root wounds into the vascular system 
while by-passing the periderm. Any 
injury acts as an infection court but 
a wound deep into the stele places the 
fungus in direct contact with the vessel, 
thus the infection court is more con- 
ducive for penetration by the fungus. 
Moisture and temperature optima may 
interact with age of wounds for maxi- 
mum infection. 


The older the wound the less chance 
for infection by the fungus. This may 
be correlated with growth responses 
by the plant at the wound site. Follow- 
ing wounding, a layer of dried cells 
forms on the pruned surface. These 
cells die as the result of injury by the 
knife. Adjacent to the dead cells is a 
zone which becomes infiltrated with 
wound substances. The trachieds re- 
main intact and ultimately become oc- 
cluded with wound substances. Tyloses 
develop in vessel members adjacent 
to the wound site. This growth re- 
sponse results in a barrier that prevents 
the fungus from invading the func- 
tional vessel members. The sequence 
of wound healing may take place much 


Born: Root Inrection wituH V. albo-atrum 


215 


faster on young root tissue. Vigor of 
the host plant will affect the time in 
which root wounds heal over. 


Good cultural practices should be 
followed when planting susceptible 
hosts in soil that may be infested with 
V. albo-atrum. When digging plant ma- 
terial, care should be taken to keep 
wounds to a minimum. Digging a ball 
larger than normal may decrease the 
chances of severing large roots. Root 
pruning should be avoided. After plant- 
ing, the application of fertilizer and 
water will decrease transplanting shock 
and increase plant vigor. If the vigor of 
the plant can be increased, root wounds 
will heal more quickly and this will 
decrease the chances of infection. 


PENETRATION AND 
DEVELOPMENT OF 
V. ALBO-ATRUM IN 


ROOTS OF WOODY HOSTS 


MATERIALS AND METHODS 


Redbud and green ash [Fraxinus 
pennsylvanica Marsh. var. subinteger- 
rima ( Vahl.) Fern.] are hereafter desig- 
nated as susceptible, and honey locust 
(Gleditsia triacanthos L.) and syca- 
more (Platanus occidentalis L.) are 
hereafter designated as resistant. The 
susceptible species were selected from 
a list of susceptible hosts of Verticillium 
albo-atrum as reported by Himelick 
(1969). The resistant hosts were so 
designated from unpublished work of 
E. B. Himelick (personal communica- 
tion). Isolates 3 and 4 were used to 
inoculate redbud and green ash re- 
spectively. A mixture of Isolates 1, 
2, 3, and 4 was used to inoculate honey 
locust and sycamore. The soil used in 
the greenhouse was a 1:1:1 ratio by 
volume of loam soil, peat, and river 
sand, steamed for 4 hours at 100 C. 


To obtain seedlings, seeds were col- 
lected in early fall and cold-stratified 
in sand for 90 days at 5 C. The strati- 
fied seeds were immersed for 2 minutes 
in a 10 percent sodium hypochlorite 


216 


solution and germinated in perlite un- 
der glass. Twenty seedlings of each 
species in the 2-leaf stage were inocu- 
lated by dipping the roots into an ap- 
proximate 1 x 10‘/ml conidial density 
of V. albo-atrum, by placing 3-mm 
blocks of PDA containing the fungus 
on selected areas, and by placing a 
Verticillium-infested oat seed adjacent 
to a selected area. After inoculation, 
the seedlings were placed horizontally 
in 150-mm petri dishes containing 
sterile peat moss or planted in sterile 
soil in pots in the greenhouse. 


Selected seedlings were sectioned for 
microscopic examination at intervals 
after inoculation. The seedlings were 
removed from the petri dishes or soil 
and the portions to be sectioned were 
killed and fixed in FAA, dehydrated 
in tertiary butyl alcohol, embedded in 


Table 3.—Root colonization of susceptible redbud and green ash, and of resistant hone 
locust and sycamore seedlings, with Verticillium albo-atrum. 


Intensity of Colonization* in Susceptible and Resistant Plants 
in Specified Regions of Penetration 


Days 
Exposure 
toInoculum Host 


1 Redbud 
Green ash 
Honey locust 
Sycamore 


2 Redbud 
Green ash 
Honey locust 
Sycamore 


4 Redbud 
Green ash 
Honey locust 
Sycamore 


6 Redbud 
Green ash 
Honey locust 
Sycamore 


8 Redbud 
Green ash 
Honey locust 
Sycamore 


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Tuutinois Naturat History SuRvEY BULLETIN 


Root Tip Epidermis Cortex 


Vol. 31, Art. 6 


paraffin, and sectioned at 12 to 15y 
using the technique described by Jo- 
hansen (1940). The sections were 
stained with thionin in phenol and 
counterstained with orange G. in ab: 
solute alcohol (Stoughton 1930), then 
examined under the microscope. 


RESULTS 


Fungus Growth on Root Surface 

The four genera of hosts used were 
essentially alike morphologically and 
no differences were detected in the way 
the fungus penetrated them (Table 3) 


The fungus colonized the exterioi 
surface of the epidermis (Fig. 1). The 
fungal growth was appressed over thi 
entire epidermal surface with conidio- 
phores arising at right angles from th 
surface. Tissue around the area of 
penetration became necrotic. Brown 


Inner 
Cortex 


Outer 
Xylem Phloem’ 


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a Symbols shown represent infection intensity as follows: 0=no colonization; 1=slight 
and 3=severe colonization. 


> Passage of the fungus through the phloem into the vessel members occurred but no phloem 


colonization; 2—=moderate colonization ; 


colonization occurred. 


Aug., 1974 Born: 


Roor Inrecrion wirn V. albo-atrum 


oS | 


Fig. 1.—Verticillium microsclerotia completely colonizing the exterior surface of a green 


ash root (X 250). 


necrotic flecks could be seen extending 
above but not below the point of infec- 
tion. 


Root Tip Penetration 

The fungus penetrated the root cap 
within 48 hours. The hyphae pene- 
trated both intercellularly and intra- 
cellularly, but intracellular penetration 
Was most common. There was no ten- 
dency for the cells to separate, which 
might have occurred if a weakening of 
the middle lamella took place, unless an 
extremely high inoculum potential oc- 
curred on the root surface. 


In the region of root elongation 
and maturation, the fungus penetrated 
‘through the epidermis. Penetration was 
either direct through the cell wall or 
between the epidermal cells. The 
hyphae or germ tubes produced appres- 
sorium-like swellings over the epidermis 
within 48 hours. A penetration peg de- 
veloped from the appressorium and was 
smaller in diameter than the parent 
hypha. 


Penetration in Root Hair Region 


In the epidermal area between the 
root hairs, the fungus penetrated at 
random, both inter- and intracellularly. 
Germ tubes developed over the root 
hairs but none was seen penetrating the 
root hairs. The base of the root hair 
frequently was penetrated but no fur- 
ther growth occurred. 


Penetration in Area 
of Lateral Root Formation 


Another avenue for fungus penetra- 
tion into roots is the area of lateral 
root formation. Rupture of the primary 
root tissue did not occur until the 
lateral root primordia were well de- 
veloped. The fungus penetrated the 
torn areas where the lateral root 
emerged. Mycelia could be seen in the 
cortical layers of the lateral root but 
none was observed in the xylem. At 
this point in the process of invasion no 
differences were detected between the 
susceptible and resistant hosts. 


218 


Cortical Invasion — 

Susceptible Hosts 

Most mycelial growth in the cortex 
was intracellular. Mass penetration re- 
sulted from a high inoculum potential at 
the invading point, and the mycelial de- 
velopment was centripetal (Fig. 2). 
Many hyphae at the point of penetra- 
tion formed appressorium-like swellings 
against the cortical cell wall and pene- 
trated to the next cell layer (Fig. 3a). 
Other hyphae that penetrated the corti- 
cal cells were constricted in diameter 
at the point of penetration (Fig. 3b). 


When the invasion of the inner corti- 
cal layers was limited to a few hyphae, 
no marked centripetal alignment of hy- 
phal strands occurred. Hyphal strands 
sometimes deviated from the centripetal 
development and developed tangen- 
tially and intercellularly for several cell 
layers and then penetrated directly 
through the wall. 


Intrinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 6 


Cortical Invasion — 

Resistant Hosts 

Most mycelial growth in the cortex 
was intracellular. The mycelium was 
hyaline but became heavily pigmented 
after 3 days. After 8 days, most hyphae 
were dark brown, regularly septate, and 
swollen between the septa so as to 
appear torulose. These hyphal strands 
gave rise to microsclerotia by re- 
peated budding (Fig. 4a). Micro- 
sclerotia varied in shape, from elongate 
to irregularly spherical, and varied in 
size, from 15 to 75, in diameter. These 
microsclerotia continued to enlarge, 
which caused cortical cells to be sepa- 
rated or expanded many times their nor- 
mal size (Fig. 4b). 


Penetration of Vascular Region 
of Susceptible Hosts 

If the fungus penetrated the cortical 
cells of the susceptible hosts, it in- 


Fig. 2.—Longitudinal section of redbud cortex showing mass penetration of cortical cells 
resulting from a high inoculum potential at the invasion point (X 400). 


Born: Roor Inrectrion wire V. albo-atrum 


Fig. 3.—Cortical cells in longitudinal section. A) Appressorium-like swellings against 
the cortical cell wall (X 2500). B) Hyphal constriction in diameter at the point of penetration 
through a cortical cell wall (X 2000). 


220 Intiwo1s NaturaL History SURVEY BULLETIN Vol. 31, Art. 6 — 


Fig. 4.—Cortical cells of honey locust in longitudinal section. A) Dark brown, septate, 
budding hypha (X 2200). B) Microsclerotia causing cortical cells to be separated or expanded 
many times their normal size (X 250). 


Aug., 1974 


variably penetrated the endodermis and 
vessel members. The fungus grew to 
the endodermal layer within 4 days. 
The quantity of vessel members in- 
vaded appeared related to the number 
of points of entry and to the mass of 
mycelia that developed from the points 
of entry. 

The hyphae that penetrated the en- 
dodermis usually penetrated the vessel 
members through pits. The hypha nar- 
rowed to a thin, peg-like projection as 
it grew through the pit. Hyphae did 
not necessarily stop at the first vessel 
member contacted, but in many in- 
stances they grew out through a pit 
on the wall of the vessel into an ad- 
jacent vessel on the side opposite the 


entry point (Fig. 5). The mycelium 


was generally unbranched, hyaline, 
3.5 in width. No typical conidiophores 
were observed. 


Born: Roor Inrecrion wire V. albo-atrum 


221 


Verticillium conidia were observed 
in the xylem 8 days after inoculation. 
In most cases, the conidia appeared to 
be free-floating in the xylem stream 
and in no way connected with the 
mycelium present (Fig. 6 and 7). The 
conidia often were found lodged at 
the end walls of the vessel members 
(Fig. 8). No defense mechanism such 
as tyloses or gum deposits was observed 
in the xylem members. The lack of a 
defense mechanism on susceptible hosts 
is in complete disagreement with other 
workers’ data on hops and cotton 


(Table 4). 


Penetration of Vascular Region 
of Resistant Hosts 


Although the fungus penetrated the 
cortical cells, few hyphae penetrated 
the endodermis and vessel members. 
The quantity of vessel members in- 


Fig. 5.—Longitudinal section through the vascular cylinder of redbud showing a hypha 
within a vessel member (X 850). 


Intivois NaturAL History SuRVEY BULLETIN 


Vol. 31, Art. 6 


eee 


Aug., 1974 Born: Roor Inrecrion wirn V. albo-atrum 223 


Fig. 8.—Longitudinal section showing lodged conidia at the end walls of the vessel 
member of a redbud root (X 2300). 


Table 4.—A comparison of penetration and development of Verticillium albo-atrum in 
roots of herbaceous and woody hosts. 


Intensity of Colonization in Susceptible (S) and Resistant (R) Plants*® 


Hops» Cotton” Woody Ornamentals* 
S R S R Ss 8S R R 

Region of Infection Daltapine Acata Green Honey 
and Signs of Disease Fruggle OR/55 15 4-42 Redbud Ash Locust Sycamore 
Epidermis 3 3 3 3 3 3 3 3 
Root hairs 2 2 2 2 0 0 0 0 
Lateral roots 0 0 - 0 0 1 1 1 a 
Cortical colonization 3 3 3 3 3) 3 3 3 
Endodermis 3 3 3 3 3 3 at 1 
Phloem colonization 0 0 0 0 0 0 0 0 
Xylem colonization 3 2 3 2 3 3 1 1 
Conidia (xylem) 3 1 3 2 3 3 0 0 
Microsclerotia (cortex) 0 0 0 0 0 0 3 3 
Mechanical plugging 

(xylem) 1 2 2 2 0 0 1 1 


8 Symbols shown represent infection intensity as follows: 0—=no colonization; 1=slight 
colonization; 2 — moderate colonization; and 3 = massive colonization, 


b Data on hops from Talboys (1958); data on cotton from Garber & Houston (1966). 
© Redbud and green ash are susceptible; honey locust and sycamore are resistant. 


vaded did not appear related to the The hyphae that penetrated the 
number of points of entry or to the mass endodermis and vessel members did 
of mycelia that developed from the so through pits in the same manner as 
points of entry. in the susceptible hosts. Few hyphae 


224 


were observed in the vessel mem- 
bers. The mycelium was hyaline, un- 
branched, and 2.71 in diameter. The 
mycelia did not ramify throughout the 
vessel members as they did in the sus- 
ceptible hosts. No conidia could be 
seen in the xylem members although 
mycelium was present. 

Frequently, microsclerotia developed 
in the vessel members, completely plug- 
ging the vessel members (Fig. 9a). 
They arose from single hyphae by re- 
peated budding of heavily pigmented, 
thick-walled cells. The microsclerotial 
cells often grew through pit pairs and 
moved into adjacent vessel members 
where repeated budding took place 
(Fig. 9b and 9c). Germinated micro- 
sclerotial cells were also observed that 
grew through pit pairs into adjacent 
vessel members. 

The ray parenchyma was heavily 
colonized with microsclerotia. Germ 
tubes from microsclerotia grew from 
one parenchyma cell to another through 
pit pairs or plasmodesmata (Fig. 9d). 
This may be an avenue for lateral 
growth of the fungus outward from the 
central vascular cylinder. 


DISCUSSION AND CONCLUSIONS 

Conidia of V. albo-atrum germinated 
on the surface of both the susceptible 
and resistant roots and grew in random 
directions. Some germ tubes grew 
away from the host; others penetrated 
the epidermis. Although germ tube 
penetration occurred, most epidermal 
penetration was by either hyphae or 
germinated microsclerotia. Intercellular 
and intracellular penetration occurred 
within 48 hours after inoculation. Nel- 
son (1950) found that V. albo-atrum 
penetrated peppermint roots 6 hours 
after inoculation. Reid (1958) reported 
intercellular penetration but observed 
no intracellular penetration of melon 
roots by F. bulbigenum Cook and 
Massee. 

According to Anderson & Walker 
(1935), F. conglutinans Wollenw. pen- 


Ituinois NATURAL Hisrory SuRVEY BULLETIN 


Vol. 31, Art. 6 


etrated the cell walls of cabbage plants — 
by mechanical pressure. Talboys (1958) 
found that the splitting apart of hop 
cells by V. albo-atrum was a mechani- 
cal rather than an enzymatic process, 
My evidence through visual observa: 
tion did not suggest that an enzyme 
was involved in either epidermal pene- 
tration or cortical invasion unless the 
cells were invaded by a mass of hyphae. » 
This is in agreement with Garber & 
Houston (1966) on Verticillium inva-— 
sion of cotton. Direct penetration was 
either by constriction of a hypha as it 
passed through the wall or by a peg-- 
like projection of an appressorium-like — 
swelling. Garber & Houston (1966) 
noted similar structures in cotton cells 
invaded by Verticillium. q 


I did not observe penetration of root 
hairs although it has been reported by 
Smith & Walker (1930) for Fusarium 
invasion of cabbage roots and by Gar- 
ber & Houston (1966) for Verticillium 
invasion of cotton roots. 


The areas of lateral root emergence 
were not important as infection court 
The fungus penetrated the lateral root 
and ramified throughout the cortical 
tissue, but no mycelia were found in 
vading the vascular tissues. Many un- 
injured roots were invaded to the same 
cortical layers. Smith & Walker (1930) 
reported similar observations; however, 
Reid (1958) suggested that penetration 
of emerging lateral roots might provide 
a mechanism for a vascular pathogen to 
avoid the penetration barrier of the 
endodermis. 


The progress of infection in the sus- 
ceptible green ash and redbud and the 
resistant honey locust and sycamore 
was identical after the point of cortical 
colonization. The species were alike 
in morphology and were penetrated by 
the fungus in a comparable fashion. 
Differences in fungus growth were 
noted immediately as the fungus pro- 
gressed beyond the initial cortical col- 
onization. 


In the susceptible species, mycelia 


Aug., 1974 Born: Roor Inrection wiru V. albo-atrum 225 


Fig. 9.—Longitudinal section through the vascular cylinder of a sycamore root. A) Ger- 
minating microsclerotium of V. albo-atrum which has completely plugged a vessel member 
(X 550). B, C) Budding cells growing through pit pairs into adjacent vessel members 
(X 1000). D) Ray parenchyma heavily colonized with microsclerotia and microsclerotium 
germinating (X 1000). 


226 


ramified throughout the tissues and 
reached the endodermis and xylem ele- 
ments within 4 days. Conidia were 
found in the vessels of roots 8 days 
after inoculation. Lack of mycelial con- 
nections between fungus parts present 
in the xylem and conidia at secondary 
sites higher in the root system can be 
explained by conidial movement. It 
is reasonable to assume that free-float- 
ing conidia moved to secondary infec- 
tion sites and provided for rapid fungus 
dispersal throughout the plant. In some 
susceptible cotton plants Schnathorst 
et al. (1967) found that 30,000 conidia/ 
ml of tracheal fluid were present 96 
hours after inoculation. 


In the resistant species, microsclerotia 
were produced in abundance in the 
cortex. These structures enlarged by 
repeated budding and ruptured the 
walls of the cortical cells. Few hyphae 
penetrated the endodermis and reached 
the xylem members. Few hyphae were 
found in the xylem members and 
conidia were not observed. Schnathorst 
et al. (1967) found that tolerant va- 
rieties of cotton depressed conidial 
numbers more than 20 fold. 


Talboys (1964) postulated that the 


xylem defense-response is much the 
same in different species and cultivars 
of plants. Since it is a generalized re- 
sponse to physical damage and infec- 
tion, the difference in host resistance to 
vascular infection is constituted by a 
difference in response of the extra- 
vascular tissue at the early stage of 
infection. This I found to be only 
partially true. The endodermis pro- 
hibited mycelial penetration to some 
extent in the resistant hosts. However, 
a xylem-defense response took place 
after penetration of the vessel members. 
Few hyphae were found in the vessel 
members after penetration and no 
conidial production occurred. Beckman 
et al. (1962) inoculated bananas with 
Fusarium by means of a standard dose 
of microconidia introduced into the 
xylem elements and found a highly sig- 


Iuxtivo1s NAturRAL History SURVEY BULLETIN 


Vol. 31, Art. 6 


nificant difference between the xylem- 
defense response of the resistant Laca- 
tan and susceptible Gros Michel ba 
nanas. Therefore, Talboys’ postulate 
should be expanded to include the in 
fection sequence in the vascular sys- 
tem in trees. 


EFFECT OF ROOP 
INFECTION ON GROWTH ~ 
RESPONSE OF REDBUD | 
& GREEN ASH SEEDLINGS 


MATERIALS AND METHODS 

Redbud and green ash seeds were 
collected and germinated as previously 
described. After 3 weeks, 80 seedlings 
of each species were removed from the 
germination beds and the roots dipped 
in inoculum for 5 minutes. After root- 
dipping, 5 plants were planted in each 
of 32 No. 10 potting cans. 


Isolates 3 and 4 were used to ino 
late redbud and green ash respectively. 
Each isolate was grown on PDA for 
14 days at 24 C. The fungus and agar 
were macerated with water in a Waring 
blendor to produce a thick suspension 
of inoculum. An equal number of con: 
trol plants were root-dipped in a PDA 
blended suspension which did not con- 
tain the fungus and potted as described 
above. 


The plants were inoculated on March 
29. The first samples of healthy and 
infected plants were taken on April 
12 and at 2-week intervals thereafter 
until July 19. Ten plants per treatment 
were sampled on eight occasions mak- 
ing a total of 160 redbud and 160 
green ash plants. The following data 
were obtained from each treatment: 
stem height, leaf area, total number 
of leaves produced, fresh and d 
weights, water content of leaves, and 
nitrogen content of stems, leaves, and 
roots, 

Dry weights were obtained by drying 
the plant parts in an electric oven at 
80 C for 72 hours. Leaf areas were 
determined by weighing a_ specific 


Aug., 1974 


known leaf area as compared to the 
weight of the whole leaf. 

Micro-Kjeldahl determinations for 
toal nitrogen were made on bulk sam- 
ples of leaves, stems, and roots from 
healthy and infected plants. 

All data for stem height, leaf area, 
and dry weight were analyzed statistic- 
ally using a one-way analysis of vari- 
ance and student “T” tests. 


RESULTS 


Symptoms 

Fourteen days after inoculation, 
young inoculated plants were retarded 
in growth but no wilt symptoms were 
apparent. Two weeks later the plants 
were stunted and the leaves had failed 
to expand. 

Sectioned roots and stems showed 
extensive invasion of the vessel mem- 
bers by the fungus. The hyphae were 
confined to the primary xylem vessel 
members 16 weeks after inoculation. 


Dry Weight 

Infection markedly reduced dry-mat- 
ter production on both redbud and 
green ash seedlings. The mean values 
for the dry weight of whole plants 
for controls and infected plants are 
shown in Table 5 and Fig. 10. All 
weight data for leaves, stems, and roots 
were analyzed statistically and the 
mean values for the dry weights on all 


Born: Roor Inrecrion wiru V. albo-atrum 297 


sampling periods after inoculation are 
given in Tables 6 and 7 and Fig. 11 
and 12. When comparing healthy and 
infected plants, a significant difference 
in dry weight was evident for leaves 
and stems of redbud and leaves of 
green ash 14 days after inoculation. A 
significant difference in dry weight of 
roots of both hosts occurred 28 days 
after inoculation. On July 19, 112 days 
after inoculation, the percentage dif- 
ferences for healthy and infected plants 
were 45, 53, and 47 for leaves, stems, 
and roots of redbud, and 36, 17, and 
24 for leaves, stems, and roots of green 
ash, respectively. 


Leaf Number 

The mean values for the number of 
leaves for healthy and infected redbud 
and green ash plants are given in Table 
8 and Fig. 13. The infected plants 
showed limited leaf production 28 days 
after inoculation, and thereafter the 
rate of leaf production differed little 
in the two groups. 


Stem Height 

The mean values for stem height of 
healthy and infected redbud and green 
ash plants are given in Table 9 and 
Fig. 14. A significant difference in stem 
height of redbud and green ash was not 
evident until 42 days and 28 days after 
inoculation, respectively. The initial 
reduction in growth due to infection 


Table 5.—The dry weight of redbud and green ash seedlings infected with Verticillium 


albo-atrum. 
Mean Dry Weight (g per plant)* 
(10 Plants) 

Days After Redbud Green Ash 

Inoculation Noninoculated Inoculated Noninoculated Inoculated 
14 23 .16* 22 .16* 
28 aie 22** 1.20 A4t* 
42 17 .30** 1.49 -50** 
56 2.29 bi** 2.87 YALs* 
70 3.36 2.20** 3.89 2.13** 
84 4.62 2.96** 5.25 3.04** 
98 7.12 3.56** 7.23 4,09** 
112 8.21 4.29** 10.54 7.20** 


* An asterisk denotes a significant difference (0.05) between noninoculated and inoculated 
means, and two asterisks denotes a highly significant difference (0.01). 


228 Inurnois Natura History Survey BULLETIN Vol. 31, Art. 6 


GREEN ASH 


7.0 , 
6. Check = / 
5.0 Inoculated = ——-— / 
4.0 ay 


Fig. 10.—The dry 
weights of green ash 
and redbud seedlings 
after inoculation with 


REDBUD V. albo-atrum, 


MEAN DRY WEIGHT (g per plant) 


9.0 
8.0 Check= 
7.0 Inoculated= ———-—-— 


14 28 42 56 70 84 =—98 n2 
DAYS 


Table 6.—Dry weights of leaves, stems, and roots of redbud seedlings infected with 
Verticillium albo-atrum. 


Mean Dry Weight (g per plant)* 
(10 Plants) 


Leaves Stems Roots 

Days After Noninoc- Inoc- Noninoc- Tnoc- Noninoc- Inoc- 

Inoculation ulated ulated ulated ulated ulated ulated 
14 silal .06* .04 AVES? 07 .05 
28 36 .10** 14 .OT** Al .O7* 
42 82 .13** 22 .05** 16 .08** 
56 1.02 .25** 67 -15** 58 .11** 
70 1.61 1.07** 97 .64** 82 AQ** 
84 2.17 1.53** 1.23 13** 1.16 .69** 
98 3.45 1.70** 1.99 195** 1.68 -90** 

112 3.97 2.18** 2.36 1.10** 1.91 1.01** 


®An asterisk denotes a significant difference (0.05) between noninoculated and inoculated 
means, and two asterisks denotes a highly significant difference (0.01). 


Aug., 1974 Born: Roor Inrecrion witu V. albo-atrum 229 
LEAVES STEMS 
45 45 
Check = Check = 
4.0 Inoculated = — —— A Inoculated =— — — 


MEAN DRY WEIGHT (g per plont ) 
MEAN DRY WEIGHT (g per plant ) 


ROOTS 


4,5 
Check = 


Inoculated«—— — 


MEAN DRY WEIGHT (9g per plant ) 


DAYS 


Fig. 11.—The dry weights of leaves, stems, and roots of redbud seedlings after inocula- 
tion with V. albo-atrum. 


230 Inuinois NaturAL History SurvEY BULLETIN Vol. 31, Art. 6 


MEAN DRY WEIGHT (g per plant) 


lation with V. albo-atrum. 


LEAVES 


Check= 
Inoculated > ——— 


STEMS 
Check = 


Inoculated= ——— 


MEAN DRY WEIGHT (g per plant) 


ROOTS 
Check =————— 
Inoculated-——— 


MEAN DRY WEIGHT (g per plant) 


DAYS 


Fig. 12.—The dry weights of leaves, stems, and roots of green ash seedlings after inocu- 


Aug., 1974 


Table 7.—Dry weights of leaves, stems, 
Verticillium albo-atrum. 


Born: Root INFecrioN wiry V. albo-atrum 


231 


and roots of green ash seedlings infected with 


Mean Dry Weight (g per plant)* 


(10 Plants) 


Leaves Stems Roots 
Days After Noninoc- Inoc- Noninoc- Inoc- Noninoc- Inoc- 
Inoculation ulated ulated ulated ulated ulated ulated 
14 allal .06* 02 02 04 03 
28 1.04 .16** 21 1** 15 .07* 
42 74 .25** 35 L2** 25 .07** 
56 2.08 T1** 49 .20** 35 .14** 
70 2.13 £30** 90 .56** -80 AT#* 
84 3.08 1.43** 1.18 .92** 1.00 .69** 
98 4.02 2.05** 1.67 1.08** 1.45 1.04* 
112 6.26 4.01** 2.11 L6t* 1.91 1.45** 


8 An asterisk denotes a significant difference (0.05) between noninoculated and inoculated 
means, and two asterisks denotes a highly significant difference (0.01). 


Table 8.—Influence of root infection of 
of leaves produced per plant. 


redbud and green ash seedlings on total number 


Mean Number of Leaves Per Plant 


(10 Plants) 
Days After Redbud Green Ash 
Inoculation Noninoculated Inoculated Noninoculated Inoculated 
0 3.20 3.50 7.56 7.68 
14 4.80 4.02 10.00 8.50 
28 7.60 5.37 14.35 10.20 
42 8.60 6.20 16.27 12.73 
56 10.08 7.48 18.75 14.88 
70 10.90 8.65 20.95 16.90 
84 12.20 10.06 22.67 19.06 
98 13.10 11.30 23.80 20.80 
112 14.00 12.20 24.00 22.80 


was slight, but further growth of the 
inoculated plants was reduced. The 
difference in stem height between 
healthy and infected plants 112 days 
after inoculation was 37.5 percent for 
redbud and 30 percent for green ash. 


Nitrogen Content 


The nitrogen content percentages of 
redbud and green ash leaves, stems, 
and roots of healthy and infected plants 
are given in Table 10. There was 26 
percent less N in infected redbud stems 
and 31 percent less in infected green 
ash stems when compared with the 
controls 112 days after inoculation. The 
N content in the leaves and roots was 


higher in the infected plants than in 
the healthy controls. 


Water Content of Leaves 

From the fresh-weight and dry- 
weight data, the percentage water con- 
tent of leaves was determined. The 
leaf data for healthy and infected plants 
are given in Table 11. 


There was no definite pattern of 
water content between infected and 
healthy redbud or green ash seedlings. 
Frequently (but not consistently) the 
water content of the infected seedlings 
was above that of the healthy controls. 
Wilt symptoms did not occur at any 
time during the experiment. No cor- 


232 
7 REDBUD 
Check = 
Inoculated = ——— 


MEAN NUMBER OF LEAVES PER PLANT 


42 


56 
DAYS 


70 84 98 N2 


Fig. 13.—Influence of root infection of red- 
bud and green ash seedlings on total number 


ItLinois NaturAL History SuRVEY BULLETIN 


Vol. 31, Art. 6 


27 


GREEN ASH 


24 Check = 


Inoculated = ——— 


MEAN NUMBER OF LEAVES PER PLANT 
a 


14 


28 «#442 «(56 


DAYS 


70 84 98 2 


of leaves produced per plant. 


Table 9.—Influence of root infection on stem height of redbud and green ash seedlings. 


Mean Stem Height (cm per plant )* 


(10 Plants) 
Days After Redbud Green Ash 

Inoculation Noninoculated Inoculated Noninoculated Inoculated 

0 6.20 6.01 6.31 5.96 

14 *. 6.90 6.35 7.58 6.80 
28 8.37 7.30 13.45 9.04** 
42 9.25 Ti5Lee 15.16 11.58** 
56 9.75 7.64** 19.86 13.46** 
70 12.00 8.02** 22.53 15.20** 
84 13.50 8.70** 25.67 16.83** 
98 15.00 9.15** 28.25 18.75** 
112 16.00 10.00** 32.00 22.40** 


“An asterisk denotes a significant difference (0.05) between noninoculated and inoculated 
means, and two asterisks denotes a highly significant differenee (0.01). 


relation could be made on water con- 
tent between healthy and infected seed- 
lings due to sampling time or green- 
house watering maintenance 


Leaf Area 


The mean values for leaf area of 
redbud and green ash are given in 
Table 12. A significant difference in 
leaf area of healthy and infected red- 
bud and green ash was found 14 days 


after inoculation. A highly significant 
difference occurred on both hosts after 
28 days. Although leaf area was less in 
infected plants, deformity of the leaves 
was not observed. 


DISCUSSION AND CONCLUSIONS 


The presence of V. albo-atrum might 
be expected to affect the metabolism 
of the host in any or all of the following 
ways: a) obstruction to water absorp- 


Aug., 1974 Born: Roor Inrecrion wirx V. albo-atrum 233 
REDBUD 35 GREEN ASH 
18 
eee = Check= 
Inoculated = —— —— — < a 
a] Inoculated-——— 
a 
é 
i= 
g 
z = 
i : 
z x 
E Fa 
v = 
~— ” 
= 
ee = 
= 
w 
a 
Zz 
< 
w 
= 


14 28 42 56 70 84 98 = 112 
DAYS 


Fig. 


14.—Influence of root infection on 
stem height of redbud and green ash seedlings 
after inoculation with V. albo-atrum. 


Table 10.—Influence of root infection on total nitrogen content of leaves, stems, and 


roots of redbud and green ash seedlings. 


Total Nitrogen (percent dry weight) 


Redbud Green Ash 
Days After 
Inoculation Noninoculated Inoculated Noninoculated Inoculated 
Leaves 
14 2.61 3.26 4.00 3.50 
56 2.00 3.63 5.45 4.98 
112 2.87 3.41 3.03 4.11 
Stems 
14 1.71 1.23 4.50 4.10 
56 2.17 1.36 4.34 3.56 
112 1.47 1.09 4.04 2.75 
Roots 
14 2.13 1.86 3.02 3.31 
56 1.64 2.00 2.36 2.50 
112 1.50 1.69 2.10 3.95 


tion and movement; b) obstruction to 
the uptake and translocation of mineral 
nutrients; and c) production of toxic 
substances. The data have been ex- 
amined in the light of these hypotheses. 


Infection leads to a drastic reduction 
in dry-matter production of all parts 
of the plant. The greatest effect of 
infection was a reduction in stem height 
and leaf area. Total leaf area decreased 
significantly in the inoculated plant 
when compared with the control. The 


number of leaves per plant exhibited 
only a slight initial reduction and there- 
after was little affected. 


Nitrogen is one of the most important 
major nutrients affecting leaf expansion, 
but there was no evidence of a reduc- 
tion in the uptake of nitrogen. Fre- 
quently, the nitrogen content was 
higher in the infected plants than in 
the control plants. The results are sur- 
prising since the root system is the first 
part of the plant to be affected by the 


234 Inurnois NaturaL History SuRVEY BULLETIN Vol. 31, Art. 6 


Table 11.—The water content of redbud and green ash leaves in response to root infec- 
tion with Verticillium albo-atrum. 


Percentage Water Content of Leaves* 


Redbud Green Ash 
Days After 
Inoculation Noninoculated Inoculated Noninoculated Inoculated 4 
14 70 71 78 75 "| 
28 60 73 74 84 
42 59 63 78 75 
56 52 63 55 62 4 
70 53 50 67 58 > 
84 52 48 60 71 
98 52 50 62 70 
112 51 54 55 52 ‘ 


gate 


4 Percentage water content was calculated from the difference between the dry weight and 
the fresh weight. 


Table 12.—Influence of root infection on leaf area of redbud and green ash seedlings. 


Mean Value of Leaf Area Per Plant (cm2)* : 

Dane Ajier Redbud Green Ash ‘ 
Inoculation Noninoculated Inoculated Noninoculated Inoculated 

14 32.62 22.65* 29.84 21.32* 

28 72.70 36.21** 63.81 31.37** 

42 99.04 40.41** 108.43 65.72** ; 

56 107.54 54.75** 141.01 83.20** 4 

70 138.18 109.64* 186.39 103.21** : 

84 145.23 129.88* 207.46 136.81** ? 

98 190.31 147.14* 265.78 183.21** af 

112 203.08 157.86* 298.76 201.21** rd 


“An asterisk denotes a significant difference (0.05) between noninoculated and inoculated 
means, and two asterisks denotes a highly significant difference (0.01). 


fungus, and thus mineral absorption 
might be impaired. No other mineral 
nutrients were determined but it would 
seem unlikely that infection would in- 
terfere with their uptake or transloca- 
tion. 


A low water supply might be ex- 
pected to account for the general stunt- 
ing which occurred. However, the 
water content in this experiment was 
approximately the same for the infected 
plants and controls, and there were no 
symptoms of general wilt. 


The results of the growth analysis 
may be interpreted in terms of a toxin 
theory. The reduction in growth may 
be initiated by toxins entering the stems 
and leaves at concentrations below the 
level that would cause wilt or death. 
This could affect cell extension or re- 


duce photosynthesis. Therefore, at the 
meristems the toxins may interfere with 
stem elongation and thus reduce inter- 
node growth. The fact that general 
wilting was never observed would indi- 
cate a greater tolerance of toxin by 
young plants. 


EFFECT OF TEMPERATURE 
& HEAT TREATING 
ON DEVELOPMENT OF 
V. ALBO-ATRUM IN ROOTS 


MATERIALS AND METHODS 

Redbud and green ash seeds were 
collected and germinated as previously 
described. At the 2-leaf stage, plants 
were removed from the germination 
bed and root-dipped in inoculum for 
5 minutes. 


Aug., 1974 


Isolates 3 and 4 were used to inocu- 
late redbud and green ash respectively. 
Each isolate was grown on PDA for 14 
days at 24 C. The fungus mycelia and 
agar were macerated with water in a 
Waring blendor to produce a _ thick 
suspension of inoculum. The control 
plants were root-dipped in a PDA solu- 
tion without the fungus. 


After the roots were dipped, the 
plants were potted in perlite and al- 
lowed to grow for 14 days. The plants 
were then removed from the perlite and 
the roots were excised at the ground 
line. 

To study the effects of temperature 
on microsclerotial development, the ex- 
cised roots were incubated at continu- 
ous temperatures ranging from 5 to 
35 C at 5-degree intervals. The roots 
were wrapped in moist paper towels 
and then placed in capped bottles to 
maintain a moist atmosphere. 


Cultures of the fungus on PDA were 
grown at the same range of tempera- 
tures. Observations were made on the 
production of microsclerotia. 


The influence of the soil microflora 
on microsclerotial formation was de- 
termined by incubating whole roots in 
sterile and nonsterile soil in capped 
bottles. Two soil-moisture levels were 
used. One level approximated field 
capacity; the other approximated one- 
half field capacity. The temperature 
was maintained at 25 C for the 28-day 
test. 

Both tests had four root systems per 
treatment replicated three times. Ob- 
servations were made at 7-day intervals 
for 35 days. For microscopic observa- 
tion, roots were cut into small pieces, 
sectioned on a freezing microtome, and 
stained in cotton blue. 


RESULTS 


Effect of Temperature 


Abundant microsclerotia were ob- 
served in roots after 14 days incubation 
at 15, 20, 25, and 30 C. Microsclerotia 


Born: Root InFrection witu V. albo-atrum 


235 


did not develop at 35 C and were not 
observed in roots incubated at 5 and 
10 C until after 35 days. The micro- 
sclerotia tend to develop as compact 
balls of dark-walled cells (Fig. 15). 
At the lower temperatures, individual 
microsclerotia tended to be elongated, 
and some were reduced to single 
strands of rounded, dark-walled cells. 


Although the fungus failed to form 
microsclerotia on PDA at 35 C, a 
limited amount of mycelial growth 
occurred. After 14 days’ growth, abun- 
dant microsclerotia were produced 
(Fig. 16) at 15, 20, 25, and 30 C. Fewer 
microsclerotia developed at 30 and 
10 C. Little growth occurred on PDA 
after 14 days at 5 C, but measurable 
hyphal growth occurred after 35 days. 
Thus, microsclerotial development on 
PDA closely paralleled development in 
moistened roots at similar tempera- 
tures. 


Effect of Heat Treating of Soil 


Microsclerotia developed in dead 
roots incubated in both steamed and 
nonsteamed soil. Moisture levels near 
the field capacity of the soil were more 
favorable for microsclerotial develop- 
ment. Relatively few microsclerotia de- 
veloped in nonsterile soil at the low 
moisture level. Although microsclerotia 
developed uniformly and more abun- 
dantly in steamed soil, appreciable 
numbers of microsclerotia were found 
in nonsteamed soil. 


DISCUSSION AND CONCLUSIONS 


The microsclerotia of V. albo-atrum 
develop rapidly at 15 to 30 C in excised 
green ash and redbud roots after being 
incubated at high moisture levels. 
Microsclerotia were produced at 5 C, 
but a longer incubation period was re- 
quired. Temperature requirements for 
microsclerotial development on PDA 
and in dead host tissue were similar. 


The development of microsclerotia 
at low temperatures is important in 
inoculum increases in overwintering de- 


236 Intrinois NATuRAL History SURVEY BULLETIN Vol. 31, Art. 6 


4 re 


ER, 


— 


| 


Fig. 15.—V. albo-atrum microsclerotia consisting of compact balls of dark-walled cells 
on dead root tissue (X 250). 


Fig. 16.—V. albo-atrum microsclerotial development on PDA (X 250). 


| 
: 
} 
| 
| 
. 


Aug., 1974 


bris. Evans et al. (1966) found large 
numbers of microsclerotia in over- 
wintering cotton stalks where fall and 
winter weather temperatures were rela- 
tively low and there was sufficient 
moisture. The range of temperatures at 
which microsclerotia form permits the 


fungus to compete favorably with or- 


ganisms that decompose root debris. 
Born (1971) found that heat-treating 
the soil increases symptom develop- 
ment because of a decrease in compe- 
tition with other fungi. 

The present study indicates that with 
high temperature and low soil moisture 
prior to microsclerotial development, 
the inoculum level was significantly re- 


duced. 


EVALUATION OF 
SYSTEMIC FUNGICIDES 
AGAINST V. ALBO-ATRUM 


MATERIALS AND METHODS 


V. albo-atrum Isolates 1 and 2 were 
used throughout this study. Inocula 
for laboratory studies were prepared by 
growing the fungus for 14 days at 24 C 
in petri dishes containing PDA. 


Greenhouse experiments were initi- 
ated in March and ran through June. 
The day and nighttime temperatures 
were approximately 25 and 16 C, re- 
spectively. The soil consisted of a mix- 
ture of equal parts by volume of loam, 
peat, and river sand, steamed at 100 C 
for 4 hours. Soil pH varied from 6.5 
to 7.2. 


Inocula for infesting soil were pro- 
duced by growing the fungus for 14 
days at 24 C in petri dishes containing 
PDA. The fungus mats, containing both 
microsclerotia and conidia, were frag- 
mented in tap water in a Waring 
blendor for 2 minutes. The fungus was 
added to the soil at the rate of one 
culture mat in 100 ml of water/20,000 g 
of soil. The soil was stirred after add- 
ing the inoculum to distribute the fun- 
gus uniformly throughout the soil. To 
determine the inoculum potential in the 


Born: Root INFecrion wirH V. albo-atrum 


237 


soil, the soil mixture was air-dried and 
screened to break up large particles. 
One-g samples were diluted with sterile 
water to 10° g/ml, and 1 ml aliquots 
plated out on PDA + streptomycin. 
This measured an inoculum potential 
of 250,000 propagules/g of dry soil. 


When plants were inoculated di- 
rectly, a V-shaped wound was made 
with a scalpel on the primary root ap- 
proximately 5 cm below the soil line. 
A 5-mm mycelial disc was inserted 
under each flap, pressed in place, and 
wrapped with vinyl grafting tape to 
prevent drying of the inoculum and 
wound area. 


The fungicides subjected to labora- 
tory and greenhouse evaluation were: 
Benlate 50 percent WP [Methyl-1- 
(butylearbamoyl) 2-benzimidazolecar- 
bamate]—benomyl; Thiabendazole 60 
percent WP [2-4-(thiazolyl) benzimi- 
dazole|=TBZ; Bravo-6F 54 percent 
(tetrachloroisophthalonitrile); and 
Vitavax 75 percent WP (5, 6-dihydro- 
2-methyl-1, 4-oxathiin-3-carboxanilide ). 


Laboratory Studies 


The four fungicides were tested in 
vitro to determine the antifungal ac- 
tivity of each against V. albo-atrum. 
Concentrations of 1,000, 500, 100, and 10 
ng/ml (active ingredient ) aqueous sus- 
pension of each fungicide were pre- 
pared and 10-mm Whatmen filter dises 
were soaked for 5 minutes in each con- 
centration. Sterile PDA culture plates 
were seeded with a conidial suspension 
and discs from each fungicide were 
placed on the seeded culture plates, 
two per plate. This was replicated 
four times using a factorial arrange- 
ment of treatments (4 trials x 4 fungi- 
cides x 4 levels) in a completely ran- 
dom design with four plates per treat- 
ment combination. Zones of inhibition 
were measured after 4 or 5 days, at 
which time growth on control plates 
had entirely covered the agar surface. 

Laboratory bioassays were conducted 
on plant materials used in fungicide 


238 


tests in the greenhouse. The plants 
were severed at the base of the stem 
and divided into three regions: (1) 
terminal, characterized by fully ex- 
panded terminal leaves; (2) center; 
and (3) bottom, located about 5 cm 
above the severed base of the stem. 
Leaf discs (9 mm diameter ) and wood 
and bark sections (100 mm long) from 
each region were frozen at —10 C for 
24 hours prior to being placed into 
petri dishes which contained 15 ml 
PDA seeded with a conidial suspension 
of V. albo-atrum. After the plates were 
incubated at 24 C for 7 days, the di- 
ameters of the zones of inhibition were 
measured to determine the relative con- 
centration of fungitoxicant present in 
the sample. 


Greenhouse Studies 


Som DrencuEes.—Three hundred 
twenty seedlings each of sugar maple 
and Russian olive were used as plant 
material. The seedlings were 2 years 
old, bare-rooted, and 45-60 cm in 
height. The seedlings had no previous 
treatment and were just breaking dor- 
mancy. Plants wound inoculated or 
placed in infested soil were potted 2 
weeks prior to fungicide treatment. 
Control plants were treated identically, 
but without the fungus. Eight different 
treatment combinations for each of the 
four fungicides were tested; with 36- 
treatment combinations arranged as a 
4 x 3 x 3 factorial [four fungicides 
x three levels (two rates and a con- 
trol)] x three infestations (with and 
without fungus) in a completely ran- 
domized design giving a total of 320 
observations per species. The eight 
treatments were: (1) infested soil, non- 
treated plants; (2) wound inoculated, 
nontreated plants; (3) infested soil, 
plants treated with 1,500 pg/ml; (4) 
infested soil, plants treated with 500 
vg/ml; (5) wound inoculated, plants 
treated with 1,500 pg/ml; (6) wound 
inoculated, plants treated with 500 
vg/ml; (7) noninfested soil, plants 
treated with 1,500 pg/ml; (8) non- 


ILtino1is NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 6 


infested soil, plants treated with 500 — 
vg/ml. Each plant was placed in a 
No. 10 potting can. In each pot, 200 
ml of the fungicide at the designated 
concentration were applied as soil 
drenches three times at weekly inter- 
vals. Water was applied and the soil 
kept moist by watering when required. 
All fungicide treatments had 10 plants 
per treatment except that the benomyl 
treatments had 25 plants per treatment. 
Disease control was calculated by using 
the following formula. 


Percent disease control — 


Disease Disease 
incidence —_— incidence 
in control in treated x 100 


Disease incidence in control 


The noninfested treated pots were 
used for detection of fungicide phyto- 
toxicity on seedlings. 


FouiaR TREATMENTS.—A _benomyl 
derivative was applied to the foliage 
of sugar maple and Russian olive seed- 
lings to evaluate its effectiveness as a 
foliar fungicide. Solutions of the beno- 
myl derivative were prepared as fol- 
lows: benomyl (5.0 g of active chemi- 
cal) was dissolved in 100 ml of 85- 
percent concentrated lactic acid over 
heat and brought up to a liter with 
distilled water (5,000 »g/ml); benomyl 
(5.0 g of a.c.) was dissolved in a liter 
of distilled water over heat in which 
2 ml of concentrated sulfuric acid had 
been added (5,000 pg/ml); benomyl 
(5.0 g of a.c.) was suspended in a liter 
of distilled water (5,000 »g/ml). The 
pH of the benomyl-lactic acid-water 
solution was 1.2-1.5, and of the beno- 
myl-sulfuric acid-water solution was 
2.5-3.0. 


Each formulation of the benomy] de- 
rivative was applied to an equal num- 
ber of plants 2 weeks prior to soil in- 
festation. Another group of plants was 
treated with each formulation 2 weeks 
after soil infestation. Foliage was 
dipped twice to run-off in late after- 
noon to retain moisture on the foliage 
as long as possible. The fungicide was 


Aug., 1974 


prevented from contaminating the soil 
by the placing of a cardboard cover on 
the top of each can before dipping. 

To determine if the benomy] deriva- 
tive could be translocated from the 
place of application to new growth in 
sugar maple seedlings, foliar dips were 
applied to localized areas. A benomly- 
lactic acid-water solution was prepared 
as previously described. Treatments 
with 5,000 »g/ml were applied in three 
different ways — to the top three 
leaves, applied to leaves on the lower 
two branches, and applied to all leaves 
on one side of the plant. 


The agar diffusion bioassay method 
was used to detect fungitoxic chemicals 
in 9-mm leaf discs above and below 
the area of treatment or in 10-mm sec- 
tions of xylem tissue. 


Roor TrEATMENTS.—Benomyl, thi- 
abendazole, Bravo-6F, and Vitavax 
were applied as root dips to evaluate 
each fungicide as a prophylactic against 
root penetration by the pathogen. Four 
liters of each fungicide were formu- 
lated at 1,500 »g/ml in distilled water. 
Ten plants of each species were al- 
lowed to stand in each fungicide for 
5 minutes. Only the roots were covered 
with the fungicide. After 5 minutes 
each plant was removed from the dip, 
shaken to remove excess liquid, and 
planted in infested soil. Each plant 
was potted in a No. 10 potting can. 
Data on phytotoxicity and symptom 
development were recorded. 


Born: Roor INFeEcrION wirH V. albo-atrum 


RESULTS 


Symptoms 

Initial wilt symptoms occurred with- 
in 7-10 days on both sugar maple and 
Russian olive seedlings after being in- 
oculated by the wound method. When 
the plants were placed in infested soil, 
symptoms occurred within 12 to 14 
days. The progression of symptom de- 
velopment was the same regardless of 
the inoculation method. The leaves 
rapidly lost their turgidity within 2-3 
days. Browning of the leaves and pre- 
mature leaf drop occurred soon after 
the leaves had wilted. Unlike larger 
trees where only a branch or several 
branches may wilt, these seedlings 
wilted quickly and completely. 


Laboratory Studies 

With the paper disc bioassay in vitro, 
benomyl and TBZ were highly inhibi- 
tory at a concentration of 10 pg/ml 
(Table 13). Vitavax was somewhat less 
fungitoxic, and Bravo-6F was much 
less active. As the concentration of 
each fungicide decreased the zone of 
inhibition decreased proportionately 
(Fig. 17). The minimum concentration 
of benomyl and TBZ, that inhibited 
growth was 0.01 and 0.1 »g/ml, respec- 
tively. The minimum concentration of 
Vitavax was 0.1 »g/ml and for Bravo-6F 
it was 1 pg/ml. 

In PDA plates containing benomyl 
or TBZ, conidia germinated but failed 
to grow more than a few microns in 


Table 13.—Paper disc bioassay of fungicides against Verticillium albo-atrum in vitro. 


Concentration® Pungicide 
ug/ml Benomyl Thiabendazole Bravo-6F Vitavaxr 
Diameter of zone of inhibition (mm)?° 

1000.000 48 55 12 47 

500.000 41 53 8 41 

100.000 30 43 6 18 
10.000 25 39 5 15 
1.000 11 8 ik 7 
0.100 3 al PIG 2 
0.001 1 


“ug/ml based on weight of active ingredient of fungicide. 
>» Zone of inhibition computed as average of four trials with four replications per trial. 


240 Intinois NaturAL History SURVEY BULLETIN Vol. 31, Art. 6 


length. When single conidia were trans- Greenhouse Studies 
ferred from these plates to PDA slants 


after 10 days, more than 90 percent Som. Drencues.—When benomyl, 
gave rise to established colonies, TBZ, Vitavax, and Bravo-6F were ap- 


Fig. 17.—Filter paper disc bioassay of four fungicides for the control of V. albo-atrum 
illustrating zones of inhibition outward from filter discs. A) Benomyl. B) Thiabendazole. 
id aa D) Vitavax (1 =10 ug/ml; 2=100 ug/ml; 3 =500 ug/ml; 4= 1,000 
ug/ml). 


Aug., 1974 


. 


plied as soil drenches to sugar maple 
and Russian olive seedlings, each gave 
some degree of disease control except 
Bravo-6F at 500 pg/ml (Table 14). 
Benomyl, TBZ, Vitavax, and Bravo-6F, 
in descending order, were effective 
when applied 2 weeks after soil infesta- 


tion. Benomyl at 1,500 »g/ml gave the 


best control of Verticillium wilt of both 
sugar maple and Russian olive seed- 
lings. The fungicide concentration, 
whether at 1,500 »g/ml or 500 pg/ml, 
at the time of application made little 
difference in the percentage of disease 
control. Benomyl] at 1,500 »g/ml and 
500 pg/ml gave 47.5 and 42.5 percent 


Born: Roor Inrecrion wirx V. albo-atrum 241 


disease control, respectively, on sugar 
maple seedlings. Differences were no- 
ticed when comparisons were made be- 
tween fungicides and fungicide con- 
centrations. On Russian olive seedlings, 
TBZ at 500 pg/ml gave the same 
amount of control as Vitavax at 1,500 
pg/ml. Benomyl at 500 pg/ml gave 
less control than TBZ at 1,500 pg/ml. 
Therefore the rate of soil application of 
any one fungicide is important in the 
control of the disease. 


Bioassay of terminal, center, and 
lower leaves of plants treated with a 
soil drench with each fungicide showed 
the highest accumulation of the fungi- 


Table 14:—Effect of soil drenches for the control of Verticillium wilt of sugar maple 


and Russian olive seedlings. 


Intensity of Infection” 


Fungicide and Number Plants With Plants Without Percent 
Concentration® of Plants Wilt Symptoms Wilt Symptoms Disease Control 
Sugar maple 
Benomyl 
1,500 50 21 29 47.5 
500 50 23 27 42.5 
Thiabendazole 
1,500 20 10 10 37.5 
500 20 11 9 31.2 
Bravo-6F 
1,500 20 15 5 6.3 
500 20 16 4 0.0 
Vitavax 
1,500 20 12 8 25.0 
500 20 14 6 12.0 
Control 20 16 1 0.0 
Russian olive 
Benomyl 
1,500 50 19 31 55.0 
500 50 24 26 43.5 
Thiabendazole 
1,500 20 a 11 47.0 
500 20 10 10 41.1 
Bravo-6F 
1,500 20 15 5 12.0 
500 20 17 3 0.0 
Vitavax 
1,500 20 10 10 41.1 
500 20 all 9 35.3 
Control 20 17 3 0.0 


8 Pungicides applied three times as a soil drench at the rate of 200 ml of aqueous suspension 


per pot. 
dient-aqueous suspension, 


Plants bioassayed 30 days after last treatment. 


Concentration at wz/ml active ingre- 


> Data on symptom development taken 30 days after last treatment. 


249 Ituivors NaturAL History SuRvEY BULLETIN Vol. 31, Art. 6 


toxicant in the lower leaves and stems all other fungicides in both leaves and 
(Tables 15 and 16). Benomyl was de- stems whether it had been applied at 
tected in higher concentrations than 1,500 or 500 pg/ml. Bravo-6F could 


Table 15.—Effect of soil drenches on uptake and translocation of fungitoxic materials 
by sugar maple seedlings. 


Tissues and Portions of Plant Sampled” 
Fungicide and eaves Wood: 
Concentration" 


Terminal Center Lower Top Center Bottom 


Diameter of zone of inhibition (mm) 


Benomyl 
1,500 27 34 41 21 19 23 
500 18 23 24 13 17 17 
Thiabendazole 
1,500 18 23 25 14 16 18 
500 13 16 17 9 11 11 
Bravo-6F 
1,500 0 0 0 0 0 0 
500 0 0 0 0 0 0 
Vitavax 
1,500 19 25 28 16 21 23 
500 16 15 18 14 13 12 
Control 0 0 0 0 0 0 


a Fungicides applied three times as a soil drench at the rate of 200 ml of aqueous suspension 
per pot. Plants bioassayed 30 days after last treatment. Concentration at ug/ml active ingre- 
dient-aqueous suspension. 


> Leaf disc (9 mm diameter) ; wood sections (10 mm long). 


© Top (characterized by fully expanded terminal leaves) ; bottom (5 cm above severed base 
of stem). 


Table 16.—Effect of soil drenches on uptake and translocation of fungitoxic materials 
by Russian olive seedlings. 


= Tissues and Portions of Plant Sampled” 


Fungicide and ee ee et J, Wo 
Concentration* Terminal Center Lower Top Center Bottom 


Diameter of zone of inhibition (mm) 


Benomyl 
1,500 18 19 22 13 15 15 
500 12 13 16 8 11 12 
Thiabendazole 
1,500 16 16 18 St 13 14 
500 9 9 11 5 6 6 
Bravo-6F 
1,500 0 0 0 0 0 0 
500 0 0 0 0 0 0 
Vitavax 
1,500 10 11 13 6 6 7 
500 8 10 10 5 6 6 
Control 0 0 0 0 0 0 


“Fungicides applied three times as a soil drench at the rate of 200 ml of aqueous suspension 
per pot. Plants bioassayed 30 days after last treatment. Concentration at ug/ml active ingre- 
dient-aqueous suspension. 

> Leaf disc (9 mm diameter) ; wood sections (10 mm long). 


¢Top (characterized by fully expanded terminal leaves); bottom (50 mm above severed 
base of stem). 


Aug., 1974 


not be detected in any plant tissue 
above ground. A higher concentration 
of the fungitoxicant accumulated in 
the sugar maple seedlings than in the 
Russian olive seedlings. The bioassay 
of foliage and wood from the sugar 
maple produced zones of inhibition ap- 
proximately twice as large as those 
from Russian olive seedlings. 


Foursar TREATMENTS.—A foliar ap- 
plication of benomy], dissolved in lactic 
acid or sulfuric acid, 2 weeks prior to 
soil infestation gave the best control 
(Fig. 18). Benomy] suspended in water 
gave less control than either applica- 
tion of benomyl dissolved in acid. 
When the application of benomyl was 
delayed for 2 weeks after soil infesta- 
tion, little control occurred. All foliar 
applications, regardless of formulations, 
gave better control if they were applied 
prior to soil infestation. 


Benomyl, or a benomyl derivative, 
was detected moving upward to areas 
of new growth after it had been ap- 
plied to localized areas at 5,000 pg/ml. 
After applications had been made to 
the top three leaves of sugar maple 


fo) 


SUGAR MAPLE 


NUMBER OF WILTED PLANTS 
O-NUEAUHYDOOCO_NUbUAHYDO 


A B c D A B c 


Treated 2 Weeks 
prior to soil Infestation 


Born: Roor INFection wiry V. albo-atrum 


SUGAR MAPLE 


243 


seedlings, a fungitoxic material could 
be detected in the treated leaves, but 
no fungitoxic material was found moy- 
ing downward in the wood. After a 
benomy! derivative was applied to the 
lower two branches and leaves, a fungi- 
toxic material was found in the foliage 
and vascular wood of the treated area, 
and also in the untreated foliage and 
wood above the point of application. 
When applications were made to leaves 
on one side of the plant, a fungitoxic ma- 
terial was found adjacent to the treated 
area and upward in the nontreated 
areas. Therefore, a benomy] derivative 
was translocated from the treated areas 
to adjacent nontreated areas above the 
point of application. No fungitoxic ma- 
terials were detected below the point of 
application. 


Roor TREATMENTS.—Root infection 
of sugar maple and Russian olive seed- 
lings can be reduced and symptom ex- 
pression delayed by dipping the roots 
with fungicides before placing them in 
infested soil (Table 17). Benomyl, 
TBZ, Bravo-6F, and Vitavax all gave 
some degree of control against Verticil- 


Fig. 18.—Degree of Verticillium 
wilt control with various foliar ap- 
plications of benomyl: A) Beno- 
myl/lactic acid/water; B) Beno- 
myl/sulfuric acid/water; C) Beno- 
myl/water; D) Control. 


D 


Treated 2 Weeks 
after soil Infestation 


244 


ItLinois Natura History Survey BULLETIN 


Table 17.—Effect of root treatments on symptom expression of sugar maple and Russian 
olive seedlings planted in Verticillium albo-atrum-infested soil. 


Fungicide* Plants Treated 


Days After Treatment 


for Initial Percentage of 


Sugar maple 


Benomyl 10 
Thiabendazole 10 
Bravo-6F 10 
Vitavax 10 
Control 10 
Russian olive 
Benomyl 10 
Thiabendazole 10 
Brayo-6F 10 
Vitavax 10 
Control 10 


Symptom Expression Plants Wilted 
28 40 
21 60 
14 70 
18 50 

1 80 
17 30 
14 50 
12 60 
13 50 

8 70 


8 Wungicides applied as a root dip for 5 minutes at the rate of 1,500 ywg/ml. 


lium wilt. On sugar maple and Russian 
olive seedlings, benomyl was much 
more prophylactic in protecting against 
root infection than the other fungicides 
tested. All fungicides used as a pro- 
phylactic delayed initial symptom de- 
velopment. Symptoms on sugar maple 
seedlings treated with benomyl de- 
veloped 21 days later than symptoms 
on the control. Initial wilt symptoms 
on benomyl-treated Russian olive seed- 
lings occurred 9 days later than those 
on the control. 


DISCUSSION AND CONCLUSIONS 


The control of a vascular wilt patho- 
gen is extremely difficult. Systemic 
fungicides which can be applied to the 
soil, taken up by the root system, and 
translocated throughout the plant are 
the most feasible. Fungicides applied 
prior to infection may serve as a bar- 
rier which will kill or arrest the fungus 
before it becomes established. The ap- 
plication of a systemic fungicide, which 
will inhibit growth of the fungus after 
symptoms appear, may be a more logi- 
cal control measure. 


The in vitro assay for fungicide 
toxicity appears to be quantitative 
using the agar diffusion method. There 
is a proportional increase in size of the 
zone of inhibition with increase in 


quantity of the fungicide. The differ- — 


ence in inhibition may not be related 
as much to differences in toxicity of 


the chemical as to solubility and dif- i 


fusibility. The in vitro data indicate 


that these fungicides are fungitoxic at 


low concentrations and that benomy] or 
its toxic breakdown product exists in 
plant tissue at a point beyond the place 
of application. 


Benomyl, TBZ, and Vitavax, when 
applied as soil drenches, reduced symp- 
tom development after plants had be- 
come infected. The inability of any 
fungicide to give 100 percent control 
may be due to the tyloses and gum- 
like materials which inhibit the fungi- 
toxicant from being translocated to the 
foliage. Recovery of the fungitoxicant 
from wilting plants that showed vascu- 
lar plugging was limited. If the fungi- 
cide was applied before vascular plug- 
ging took place, the fungitoxicant 
readily moved throughout the plant and 
could be assayed in the above-ground 
parts. Fungicides, such as Bravo-6F, 
which show no systemic action are of 
little value in controlling Verticillium 
wilt when applied as a soil drench. 


Benomy] dissolved in either an or- 
ganic or inorganic acid and applied to 
the foliage gave better control than 
a benomyl suspension in water. With 


Vol. 31, Art. 6 | 


: 


Aug., 1974 


the addition of acids, the fungitoxicant 
was water soluble, and could be taken 
up more readily and _ translocated 
throughout the plant. The fungitoxicant 
must be localized in the plant parts be- 
fore the host-pathogen interaction pro- 
duces gums and tyloses, blocking the 
upward movement of the fungitoxicant. 
Once wilt symptoms occurred and the 
vascular system was occluded, trans- 
location of the fungitoxicant was re- 
duced regardless of the formulation. 


The time of fungicide application is 
critical for the fungitoxicant to be dis- 
tributed throughout the plant before 
the fungus can become established. 
The critical time of application was 
similar for foliar treatments and soil 
drenches. 


When fungicides were tested as pro- 
phylactic root dips, each gave some 
degree of control. Initial wilt symp- 
toms were delayed as much as 3 weeks 
with benomyl. The delaying of root 
infection may allow wounds to be oc- 
cluded with wound material before 
the fungus can become established at 
the wound site. This method of control 
may be of value when used on bare- 
rooted nursery materials. 


More work is needed to determine 
how fast these systemic fungicides will 
move in the plant and how long they 
will remain active. Additional work is 
needed to determine the critical time of 
application and if higher concentrations 

- will be more effective but not phyto- 
toxic to the host. 


SUMMARY 


Verticillium albo-atrum Reinke and 
Berthold is a widespread and destruc- 
tive vascular pathogen. It is peculiar 
in that it does not confine its attack 
to one host, or a few closely related 
hosts, but attacks a large number of 
widely unrelated plants, many of which 
are of economic importance. 

The wound most conducive to in- 
fection was a vascular wound which 


Born: Root INFecrion wiru V. albo-atrum 


245 


allowed the pathogen to come in direct 
contact with the vessel members. No 
infection took place unless a wound 
was present on the root. Root wounds 
remained as infection courts up to 32 
days on redbud and 16 days on sugar 
maple seedlings. As the age of the 
wound increased, the number of plants 
infected through wounds decreased 
sharply. 

In the susceptible hosts, the patho- 
gen rapidly colonized the cortex, endo- 
dermis, and vessel members. Conidia 
were produced in abundance within 
8 days. The pathogen in the resistant 
hosts readily colonized the cortex, but 
few hyphae were found in the vessel 
members. Conidia were not present 
in the vascular system. Microsclerotia 
were found in both the cortex and 
vascular cylinder of the resistant hosts. 


Infection leads to a significant re- 
duction in dry-matter production, stem 
height, and leaf area of the plants. The 
nitrogen content was lower in infected 
redbud and green ash stems, but higher 
in leaves and roots. There was no 
definite pattern of water content be- 
tween infected and healthy redbud and 
green ash seedlings. Frequently the 
water content of the infected seedlings 
was above that of the healthy controls, 
but not consistently. 


Abundant microsclerotia were ob- 
served in roots after 14 days when 
incubated at 15, 20, 25, and 30 C. 
Microsclerotia were observed after 35 
days at 5 and 10 C, but no micro- 
sclerotia were observed at 35 C. Micro- 
sclerotia developed in dead roots incu- 
bated at 25 C in both steamed and 
nonsteamed soil. A moisture level near 
the field capacity of the soil was more 
favorable for microsclerotial develop- 
ment than was a lower soil moisture. 


The in vitro assay of the toxicity of 
the fungicides by the agar diffusion 
method appears to be quantitative. 
There is a proportional increase in the 
size of the zone of inhibition with in- 
crease in quantity of the fungicide. 


246 


Benomyl, TBZ, and Vitavax, when 
applied as soil drenches, reduced symp- 
tom development after plants had be- 
come infected. The inability of any 
fungicide to give 100-percent control 
may be due to the host-pathogen inter- 
action producing tyloses and gum-like 
material which prevents the fungitoxi- 
cant from being translocated to the 
foliage. Fungicides which show no 
systemic action are of little value in 
controlling Verticillium wilt when they 
are applied as a soil drench. 

Benomy] which had been solubilized 
in either an organic or inorganic acid 


Intinois NATURAL History SurRvEY BULLETIN 


Vol. 31, Art. 6 


and applied to the foliage gave better 
control than a benomyl suspension 
water. With the addition of acids, the 
fungitoxicant was water soluble, and 


translocated throughout the plant. 


When the fungicides were tested as 
prophylactic root dips, all delayed 
symptom expression and gave some 
degree of control. Benomyl delayed 
initial wilt symptoms as much as 3 
weeks. The delaying of root infection 
may allow wounds to be occluded with 
wound material before the fungus can 
become established at the wound site, 


8 ee 


LITERATURE CITED 


Anperson, M. E., and J. C. WALKER. 1935. 
Histological studies of Wisconsin Hollan- 
der and Wisconsin ballhead cabbage in 
relation to resistance to yellows. Journal 
of Agricultural Research 50:823-836. 


Armstronc, G. M., and J. K. ARMSTRONG. 
1958. Effect of cutting roots on the inci- 
dence of Fusarium wilt of cotton, toma- 
toes, cowpeas, and other plants. Phyto- 
pathology 48:341. (Abstr.). 


Arnpt, C. H. 1957. Temperature as a fac- 
tor in the infection of cotton seedlings 
by ten pathogens. Plant Disease Reporter 
Supplement 246:63-84. 


Banxkoutl, M. M., and..W. D. Tuomas, Jr. 
1964. Control of Fusarium and Verticil- 
lium wilt with defolatan. Phytopathology 
54:1431. (Abstr.). 


Beckman, C. H., S. HAmos, and M. E. Mace. 
1962. The interaction of host, pathogen, 
and soil temperature in relation to sus- 
ceptibility to Fusarium wilt of bananas. 
Phytopathology 52:134-140. 


Benken, A. A., and A. KHakimoy. 1964. 
Vertitsilleznaya infektsiya v list’-yakh 
Khlopchatnika (Verticillium infection in 
cotton leaves). In Review of Applied 
Mycology 44:292. 


Born, Geratp L. 1971. Heat treatment of 
‘soil enhances Verticillium wilt infection 
of barberry and redbud. Plant Disease 
Reporter 55:996-—997. 


BrinkerHorr, L. A. 1969. The influence 
of temperature, aeration, and soil micro- 
flora on microsclerotial development of 
Verticillium albo-atrum in abscised cot- 
ton leaves. Phytopathology 59:805—808. 


BucHENAUER, H., and D. C. Erwin. 1971. 
Control of Verticillium wilt of cotton by 
spraying foliage with benomyl and thia- 
bendazole solubilized with hydrochloric 
acid. Phytopathology 61:433-434. 


CAROSELLI, Nestor E. 1957. Verticillium 
wilt of maples. Rhode Island Agricul- 
tural Experiment Station Bulletin 335:5. 


Corpa, A. C. J. 1838. Icones Fungorum 
hucusgue cognitorum, 2 (Prague). 


Eperineton, L. V., and J. C. WALKER. 1957. 
Influence of soil and air temperature on 
Verticillium wilt of tomato. Phytopath- 
ology 47:594-598. 


Erwin, D. C., J. J. Stus, D. E. Borum, and 
J. R. Curtpers. 1971. Detection of the 
systemic fungicide, thiabendazole in cot- 
ton plants and soil by chemical analysis 
and bioassay. Phytopathology 61:964—967. 

= , and J. ParrrinGre. 1968. Evi- 
dence for the systemic fungitoxie activ- 
ity of 2-(4-thiazolyl) benzimidazole in the 


control of Verticillium wilt of cotton. 
Phytopathology 58:860—865. 


Evans, G., W. C. SnypEr, and S. WILHELM. 
1966. Inoculum increase of the Verticil- 
lium wilt fungus in cotton. Phytopath- 
ology 56:590-594. 


Futon, Ropert H. 1952. Studies on Ver- 
ticillium wilt of raspberry. Phytopath- 
ology 42:8 (Abstr.). 


GaLLecLy, M. E. 1949. Host nutrition in 
relation to development of Verticillium 
wilt of tomato. Phytopathology 39:7. 
(Abstr.). 


Garser, R. H. 1957. The penetration and 
development of Verticillium albo-atrum 
Reinke and Berthold in the cotton plant. 
Ph.D. Thesis. University of California. 
60 p. 

, and Byron R. Houston. 1966. Pen- 

etration and development of Verticillium 

albo-atrum in the cotton plant. Phyto- 

pathology 56:1121-1126. 


Ginrman, J. C. 1916. Cabbage yellows and 
the relation of temperature to its occur- 
rence. Annals of Missouri Botanical 
Garden 3:25-84. 


GREEN, RAaLtpH J. 1954. An investigation of 
the wilting phenomenon in Verticillium 
wilt of tomato Lycopersicum esculentum 
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916. 


HEate, J. B., and Ivor Isaac. 1963. Wilt 
of Lucerne caused by species of Verticil- 
lium. Annals of Applied Biology 52: 
439-451. 


HIME Ick, E. B. 1969. Tree and shrub hosts 
of Verticillium albo-atrum. Illinois Nat- 
ural History Survey Biological Notes 66. 
8 p. 

Isaac, Ivor. 1949. A comparative study of 
pathogenic isolates of Verticillium. Brit- 
ish Mycological Society Transactions 32: 
137-157. 

JoHANSEN, D. A. 1940. Plant microtech- 
nique. McGraw-Hill Book Co., Inc., New 
York. 523 p. 

KLeBAHN, H. 1913. Beitrage zur Kenntnis 
der Fungi Imperfecti. I. Eine Verticil- 
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Lupsrook, W. V. 1933. Pathogenicity and 
environal studies on Verticillium hadro- 
mycosis. Phytopathology 23:117—-154. 

McWuorter, Frank P. 1962. Disease symp- 
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349-353. 

NADAKAVUKAREN, M. J. 1960. The effect of 
soil moisture and temperature on survi- 


248 


val of Verticillium microsclerotia. Dis- 
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schen Buchhandlung, Wurzburg. 329 p. 


Netson, R. 1950. Verticillium wilt of pep- 
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Prestey, J. T. 1941. Saltants from a mono- 
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Phytopathology 31:1135-1139. 

Rankin, W.H. 1914. Thrombotic disease of 
maple. Phytopathology 4:395. 

Rawuins, T. E., and J. A. Bootu. 1968. 
Tween 20 as an adjuvant for systemic 
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Plant Disease Reporter 52:944—-945. 


Rep, J. 1958. Studies on the Fusaria which 
causes wilt in melons. Canadian Journal 
of Botany 36:394—410. 


REINKE, J., and G. BertHotp. 1879. Die 
Zersetzung der Kartoffel durch Pilze. 
Untersuchungen des Botanischen Labora- 
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mycosis. Hilgardia 5:197-353. 

ScunarHorst, W. C., J. T. Prestey, and 
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ScurerBer, L. R., W. K. Hock, and B. R. 
Roserts. 1971. Influence of planting 
media and soil sterilization on the uptake 
of benomyl by American elm seedlings. 
Phytopathology 61:1512-1515. 

SeLmay, I. W., and W. R. Bucktey. 1959. 
Factors affecting the invasion of tomato 
roots by Verticillium albo-atrum. British 
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234. 


, and G. F. Peee. 1957. An analysis 
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674-681. 


Ityinois NAturAL Hisrory SuRvEY BULLETIN 


Vol. 31, Art. 6 


SEWELL, G. W. F., and J. F. Wirson. 1964. 
Occurrence and dispersal of Verticillium 
conidia in xylem sap of the hop (Humu- 
lus lupulus L.). Nature 204:901. 


SmirH, Rose, and J. C. WALKER. 1930. A 


cytological study of cabbage plants in © 


strains susceptible or resistant to yel- 
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41:17-35. 

Sroucuton, R. W. 1930. Thionin and 
orange G. for the differential staining of 
bacteria and fungi in plant tissue. An- 
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TALBoys, P. W. 1958. Association of tylosis 
and hyperplasia of the xylem with vas- 
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albo-atrum. British Mycological Society 
Transactions 41: 249-260. 


1958. Some mechanisms contrib- 

uting to Verticillium-resistance in the 

hop root. British Mycological Society 

Transactions 41:227—241. 

1964. A concept of the host-para- 
‘site relationship 
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VAN DEN Enns, G. 1958. Untersuchungen 
liber den Pflanzen-parasiten Verticillium 
albo-atrum R. & B. Acta. Bontanica Neer- 
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VAN DER Meer, J. H. H. 1925. Verticillium 
wilt of herbaceous and woody plants. 
Meded. Landbouwhogeschool Wagenin- 
gen. 28:1-82. 

Van Hook, J. M. 1904. Disease of ginseng. 
Cornell Agricultural Experiment Station 
Bulletin 219:165-186. 

WitHerm, S. 1950. Verticillium wilt in 
acid soils. Phytopathology 40:776-777. 

, and J. B. Taytor. 1965. Control of 

Verticillium wilt of olive through nat- 

ural recovery and resistance. Phytopath- 

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WoLLENWEBER, H. W. 1929. Die Wirtelpilz- 
Welkekrankheit (Verticillose) von Ulme, 
Ahorn-and Linde usw. Arb. Biol. Reich- 
sanstalt Land-u. Forstwirtsch. Berlin- 
Dahlem. 17:273-299. 


in Verticillium wilt — 


A 
Acrostalagmus albus, 210 
Appressorium, 217-218, 224 


B 
Bananas 
Gros michel, 226 
Lactan, 226 
Benomyl, 237 
Bravo 6F, 237 
Budding, 224 


(e 
Cabbage, 224 
Code of Verticillum isolates, 212-213 


Conidia, 211, 216, 221, 224, 226, 240, 245 


Cortical invasion 
resistant hosts, 218 
susceptible hosts, 218 

Cotton, 224, 226 


D 
Disease control 
determination of, 238 
Dry weight 
reduction of, 227 


E 
Endodermis, 211, 221, 223, 226 


F 
Fungal penetration 
lateral root, 217 
intercellular, 217 
intracellular, 217 
root hairs, 217 
root tip, 217 
Fusarium species 
bulbigenum, 224 
conglutinans, 224 
oxysporium f. sp. lycopersici, 212 


G 
Green ash, 215, 226 


H 
Honey locust, 215 
Hyaline, 218, 224 


l 
Inoculum potential 


determination of, 237 


E 
Lateral root, 217 


Leaf area 
determination of, 226 
mean value of, 2382 
Leaf number 
mean value of, 227 


Maple, sugar, 213 
Microsclerotia, 211-212, 218, 224, 226, 
235, 245 


INDEX 


Microsclerotial formation of excised roots 


effect of temperature on, 235 
effect of heat on, 235 
Mycelium, 217 


N 
Nitrogen 
content in leaves, 281 
determination of, 227 


p 
Panax quinquafolium, 210 
Pelargonium, 211 
Peppermint, 224 
Periderm, components of 
phellem, 214 
phelloderm, 214 
phellogen, 214 
Pit, 221 
Plasmodesmata, 224 
Potato dextrose agar (PDA), 213 


R 
Ray parenchyma, 224 
Redbud, 213, 215, 226 


Relationship of root wounds on infection 


type of, 213 
age of, 213 
Root cap, 217 
Root elongatium, 217 


S 

Statistical analysis 

one way analysis of variance, 227 

student “T” test, 227 
Stem height 

mean value of, 227 
Sycamore, 215 
Systemic fungicide treatments 

foliar treatments, 243 

root treatments, 243 

soil drenches, 240 


T 
Thiabendazole, 212, 237 
Trachieds, 215 

Tyloses, 211, 214-215, 246 


Vv 
Vascular invasion 


resistant hosts, 221 

susceptible hosts, 218 
Verticillium dahliae, 210 
Vitavax, 237 


WwW 
Water content of leaves 
determination of, 231 
Wilt, symptoms of 
woody hosts, 209 
tomatoes, 211 


Wounds as sources of pathogen entry, 213 


Deo 
an 


a 


= 
(¢ He OU as 


2 
Re 


Some Publications of the ILLINOIS NATURAL HISTORY SURV; 


BULLETIN 


Volume 30, Article 7—A Comparative Study 
of Two Components of the Poinsettia 
Root Rot Complex, By Robert S. Perry. 
August, 1971. 35 p., index. 

Volume 30, Article 8—Dynamics of Condi- 
tion Parameters and Organ Measure- 
ments in Pheasants. By William L. An- 
derson. July, 1972. 44 p., index. 


Volume 31, Article 1—The Effects of Sup- 
plemental Feeding and Fall Drawdowns 
on the Largemouth Bass and Bluegills at 
Ridge Lake, Illinois. By George W. Ben- 
nett, H. Wickliffe Adkins, and William F. 
Childers. January, 1973. 28 p., index. 


Volume 31, Article 2—The Reproductive 
Cycle of the Raccoon in Illinois. By Glen 
C. Sanderson and A. V. Nalbandov. July, 
19738. 57 p., index. 


Volume 31, Article 3.—Nutritional Re- 
sponses of Pheasants to Corn, with Spe- 
cial Reference to High-Lysine Corn. By 
Ronald F. Labisky and William L. An- 
derson. July, 1973. 26 p., index. 


BIOLOGICAL NOTES 


75.—Illinois Birds: Turdidae. By Richard 
R. Graber, Jean W. Graber, and Ethelyn 
L. Kirk. November, 1971. 44 p. 


76.—Illinois Streams: A _ Classification 
Based on Their Fishes and an Analysis 
of Factors Responsible for Disappearance 
of Native Species. By Philip W. Smith. 
November, 1971. 14 p. & 

77—The Literature of Arthropods Asso- 
ciated with _Soybeans. I. A Bibliography 
of the Mexican Bean Beetle, Epilachna 
varivestis Mulsant (Coleoptera: Coc- 
cinellidae). By M. P. Nichols and M. 
Kogan. February, 1972. 20 p. 

78.—The Literature of Arthropods Associ- 
ated with Soybeans. II. A Bibliography 
of the Southern Green Stink Bug, Nezara 
viridula (Linneaus) (Hemiptera: Pen- 
tatomidae). By N. B. DeWitt and G. L. 
Godfrey. March, 1972. 23 p. 


79.—Combined Culture of Channel Catfish 
and Golden Shiners in Wading Pools. By 
D. Homer Buck, Richard J. Baur, Charles 
F, Thoits III, and C. Russell Rose. April, 
1972. 12 p. 


80.—Illinois Birds: Hirundinidae. By Rich- 


List of available publications mailed on request 


No charge is made for publications of the Ixt1nors Narurat History Survey. A 
copy of most publications will be sent free to anyone requesting it until the s 
comes low. Costly publications, more than one copy of a publication, and publica 


the writer and explain the use to be made of the publication or publications. 


Address orders and correspondence to the Chief, 
Illinois Natural History Survey 


ard R. Graber, Jean W. Gr 
Ethelyn L. Kirk. August, 19) 


81.—Annotated Checklist of the B 
of Illinois. By Roderick R. 
John C. Downey. May, 1973. 


82.—Lactate Dehydrogenase Is 
Darters and the Inclusivene: 
Genus Percina. By Lawrence 
and Gregory S. Whitt. May, 19 

83.—Illinois Birds: Laniidae. 
R. Graber, Jean W. Graber, and 
L, Kirk. June, 1973. 18 p. 


84.—Interactions of Intensive © 
Channel Catfish with Largemou 
in 1-Acre Ponds. By D. Homer 
Richard J. Baur, and C. Rus : 
February, 1974. 8 p. 


85.—The Literature of Arthropo 
ated with Soybeans. III. A Bibli 
of the Bean Leaf Beetles, Oe 
trifurcata (Forster) and @. 7 
(Olivier) (Coleoptera: Chryson 
By M. P. Nichols, M. Kogan, 
Waldbauer. February, 1974. 


86.—Illinois Birds: Tyrannidae. 
ard R. Graber, Jean W. i 
Ethelyn L. Kirk. February, 19 


87.—The Literature of Arthropod 
ated with Alfalfa. I. A Biblio 
the Spotted Alfalfa Aphid, 7 € 
maculata (Buckton) (Homopte 
dae). By D. W. Davis, M. P. Nich 
E. J. Armbrust. February, 1974. 
88.—The Literature of Arthropods 
ated with Alfalfa. II. A Bibliogra 
the Sitona Species, 
culionidae). 
Pass, M. P. Nichols, and E, J. A 
February, 1974. 24 p. 


CIRCULAR 


47.—Illinois Trees and Shrubs: 
sect Enemies. By L. L. Englis 
1970. (Fifth printing, with alte 
91 p. . 
51.—ITHinois Trees: Selection, Plantin 
Care. By J. Cedric Carter. Augus' 
1238p ae a 
52.—Fertilizing and Watering Trees. 
Dan Neely and E. B. Himelick. Dec 
' ber, 1971. (Third printing.) 20 D. 
53.—Dutch Elm Disease in Illinois. 
Cedric Carter. October, 1967. 19 


4 ILLINOIS 
itural History Survey 
| BULLETIN 


The Mecoptera, or Scorpionflies, 
of Illinois 


C. Marlin 
| NATURAL HISTORY SURVEY 
AUG 27 1975 
LIBRARY 


F ILLINOIS 
TI ENT OF REGISTRATION AND EDUCATION 


RAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 
ae VOLUME 31, ARTICLE 7 
AUGUST, 1975 _ : 


ILLINOIS 


BULLETIN 


The Mecoptera, or Scorpionflies, 
of Illinois 


OF ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


URAL HISTORY SURVEY DIVISION 
ANA, ILLINOIS 


AUGUST, 1975 


tural History Survey 


VOLUME 31, ARTICLE 7 


STATE OF ILLINOIS 


GurTowsky, Ph.D., Chemistry ; 


Illinois University. 


NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 
SCIENTIFIC AND TECHNICAL STAFF 
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Lu-Pine LEE, M.S., Technical Assistant 


Section of Botany and Plant Pathology 

Ciaus GRUNWALD, Ph.D., Plant Physiologist and Head 
Rospert A. Evers, Ph.D., Botanist 

EuGcENneE B. Himevick, Ph.D., Plant Pathologist 

R. Dan NEEty, Ph.D., Plant Pathologist 

D. F. ScHOENEWEIS8, Ph.D., Plant Pathologist 

J. LELAND CRANE, Ph.D., Associate Mycologist 
WALTER HarTSTIRN, Ph.D., Assistant Plant Pathologist 
Betty S. NELson, Junior Professional Scientist 

GENE E. ReE1D, Technical Assistant 


Section of Aquatic Biology 

D. Homer Buck, Ph.D., Aquatic Biologist 

WiuuiaM F. CuHILpers, Ph.D., Aquatic Biologist 

R. Wetpon Larimore, Ph.D., Aquatic Biologist 
Rogert C. HiLTIBRAN, Ph.D., Biochemist 

ALLISON BRIGHAM, Ph.D., Assistant Aquatic Biologist 
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DonaLp W. Durrorp, M.S., Research Associate 

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DEPARTMENT OF REGISTRATION AND EDUCATIO) 
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CONSULTANTS AND RESEARCH AFFILIATES: Systematic Entomo.ocy, Roperick R. Irwin, Chicago, Illi- 
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life Research, Southern Illinois University ; PARASITOLOGY, Norman D. Levine, Ph.D., Professor of Veterinary 
Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, University ofa 
Illinois; Entomouocy, Rosert L. Metcaur, Ph.D., Professor of Zoology and of Entomology, University of Illinois; 
and Giipert P, WALDBAUER, Ph.D., Professor of Entomology, University of Illinois; Statistics, HoRaAcH we 
Norton, Ph.D., Professor of Statistical Design and Analysis, University of Illinois. E 


CONTENTS 
PPS UAI ING NIE NIUS UME Wal crie ies cenieieret ats, ht eancieinreteueicy ehcliioie Gis, ave-viel x Psa eze vanes 252 
SPREE SATS LOR VAR rar ea nsec jor crante it cic ivere etsielelaie choles ep iva a echoes ar 252 
IPBSOHIYE? 2 tha Stent 8 Bi OIRO RPL IOI EY PAC IE Hr ES aI ca ee a ee 252 
Miatnomands@VvipOSibOn! 22 ages + 01a, soto ac cienbers vrelre 4 cal u ie aieeiaiee + 253 
Minteaatrine mS tA CCS MEN ayes 051s ses he cle sie sniaie SA eer omy & tpepare tneievesaie erate: andolare Here Bie 254 
TSIGNDTNE Saag Oo SH oe aE Cn eee 257 
PRESERIBURIONGANDE DISPERSAL: fyclo.5 5 c.0-6 fave sohusltue oicke weieleose olde als Ge slew naa 260 
COLLECTING AND PRESERVING MECOPTERA ..........0 000 0e ce ceeeeceeeuees 265 
“ TIRTETEGIL@VEN? ga chs ple ale cy cre nec ORG On OCC OEIC RECO na ee ae 266 
MoNoGRAPHS ON NEARCTIC MECOPTERA ..........0 00 ce cece cece eee eeeues 268 
RON OMIG MUR MATINIBN Telia Sicteyc)a cio tsieys ave oor avsusgeter's + svisiteres ey equip. Sie Suse Go Lesa Sie euars 268 
WrdermVlecopterates ds) ccwrac Sesleri ein ots!s eohiace wiSiei pele, pelos mie-hiee 268 
Hani Wap It AGI ACH a cye vis cea sieie Sinise tae Sf easike Ow ehdag eiiesuca, # Bip) 8 isele ve renee 269 
Haunt Va OLGA ie erie) otarera ketal els oie cue sie ole sore Gy oieltheway, oie ioue oS eace nie a a 277 
[Pareavihy IN Veti(ay oYE(6 EVO: erahtnn Gocco SILOS Se). 1O.as ES OR eC DIO emer 280 
er TAAT aw EAT OVISOCA CAC eters oli teeiel oe. epee eve eree Yer vafevs: © MM 03) 6) clove) ova lenele ia lier 281 
AIM VawPAMOLDICAG) exec ceatevr sue teioss ene vee aye creda diene sites eters «aye sinncle-wle chee 282 
_. TTELUANTIUMS TSE (GTA aD) th 5, entice MERCURE RES PRESSED Rc One cn ne ee So evil 
THESES gig Gtasha eusideel GR GIOIA CRONE PARE Ee ERCnr Once a een ne 315 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par, 58.12. 
It is a contribution from the Section of Faunistic Surveys and Insect Identification of the 
Illinois Natural History Survey. 


Donald W. Webdbd is an Assistant Taxonomist at the Illinois Natural History Survey. 
Norman D, Penny and John C. Marlin are former graduate research assistants at the 
Survey. 


(68937—3M—R—75) 


j 
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3 


i 
| 


Pay 


Frontispiece.—A hangingfly, Bittacus pilicornis, awaiting its prey, which includes mosqui- 
toes and other bottomland insects. (Photo by W. D. Zehr) 


THE ORDER MECOPTERA (scor- 
ionflies and hangingflies) is of ancient 
lineage. Fossils of this order are known 
from as far back as the Permian. Today 
relatively few species of Mecoptera 
exist; fewer than 500 are currently 
recorded for the world. They and 
their fossil relatives exhibit many primi- 
tive characteristics and are considered 
among the oldest and most primitive 
holometabolous insects. Eighteen spe- 
ies occur in Illinois. They live in mesic 
places, especially among dense _her- 
baceous vegetation in lowland woods. 
One species of Boreus occurs only on 
moss in woods and is a relict of the 
Arctotertiary forest. This species is 
found in the southwestern corner of the 
state. 


Twenty-one families of Mecoptera 
are recognized, a dozen of which are 
represented only by fossils. Of the nine 
extant families, the Bittacidae (hang- 
ingflies) are the most widespread, oc- 
curring on all continents in tropical and 
arm-temperate regions. The families 
Notiothaumidae (found only in South 
America ) and Meropeidae (one mono- 
typic genus in Australia and one in 
North America ) are considered the most 
primitive. Three families, Choristidae, 
\Nannochoristidae, and Apteropanorpi- 
jdae, are restricted to the southern 
‘hemisphere, occurring in Australia, Tas- 
mania, or New Zealand. The remaining 
|three families, Boreidae, Panorpodidae, 
and Panorpidae, are found in North 
‘America and Eurasia. 


The five families (Bittacidae, Borei- 
dae, Meropeidae, Panorpodidae, and 
Panorpidae ) occurring in North Amer- 
ica contain 80 species. The majority 
of these species are distributed through- 
out the eastern United States. Other 
species occur in Central America, 
Mexico, and the western coastal states. 
With the exception of the family 


The Mecoptera, or Scorpionflies, of Illinois 


Donald W. Webb, Norman D. Penny, and John C. Marlin 


Boreidae, no Mecoptera have been re- 
corded north of the 50th parallel in 
North America. 


The center of distribution of Mecop- 
tera in the United States is in the 
southern Appalachians (Byers 1969), 
from which area the various species 
have dispersed themselves northward 
and westward. Thirty-two species are 
recorded in the Midwest. Illinois, with 
its extensive north-to-south length and 
geological history, provides a wide 
variety of habitats for most groups of 
Mecoptera. The glaciated regions of 
northern Illinois, in particular the 
Northeast Morainal Division’, offer suit- 
able habitat for species, such as Panorpa 
subfurcata, P. mirabilis, and P. galerita, 
distributed primarily or wholly in pre- 
viously glaciated areas. The Coastal 
Plain Division (Austroriparian Divi- 
sion) at the southern tip of Illinois 
is attractive to those species, such as 
Panorpa nuptialis, distributed in the 
coastal plains of the southern Atlantic 
and Gulf states. The narrow strip of 
Ozark Division in southwestern Illinois 
is an extension of the Ozark uplift and 
provides habitats for species such as 
Panorpa braueri. Similarly, the Shawnee 
Hills Division of southern Illinois con- 
tains habitats similar to those in the 
southern Appalachians and in Kentucky 
and Tennessee for such species as Bit- 
tacus punctiger. The central part of 
Illinois has areas of deciduous forest 
along the eastern boundary and prairie 
and mixed woodland to the west that 
provide habitats for the other mid- 
western species. 

The objective of this study is to up- 
date our knowledge of the distribution 
and natural history of Mecoptera, par- 
ticularly in relation to the biogeographic 


1Terms from “The Natural Divisions of Illi- 
nois,” Illinois Nature Preserves Commission, 
1972. 


251 


252 


history of Illinois. Synoptic descrip- 
tions, keys, and illustrations have been 
prepared to provide an insight into this 
primitive and interesting group of in- 
sects. 


The emphasis of this study is on the 
fauna of Illinois, but other species oc- 
curring in the Midwest have been in- 


cluded. 


Collecting data are listed for those 
Illinois species known from fewer than 
ten localities. Records for other species 
are plotted on distribution maps. 


ACKNOWLEDGMENTS 


Considerable cooperation and advice 
have been required for this study, and 
the authors wish to express their sin- 
cere appreciation to colleagues who 
have supported and encouraged this 
work. We wish to thank L. J. Stannard, 
Illinois Natural History Survey, for his 
advice and guidance in collecting speci- 
mens and locating unique habitats in 
Illinois and H. H. Ross, University of 
Georgia, for his review of the manu- 
script. In particular, we wish to thank 
G. W. Byers, University of Kansas, for 
his advice on the identification of 
Mecoptera, comments during the prep- 
aration of this manuscript, and time 
spent in meticulously reviewing the 
final copy. 


Our sincere appreciation is offered to 
the following organizations and indi- 
viduals for the loan of material in their 
collections: American Museum of Natu- 
ral History, W. Brigham Collection 
(Mahomet, Illinois ), Canadian National 
Collection, Cornell University, Eastern 
Illinois University, Field Museum of 
Natural History, G. Finni Collection 
(West Lafayette, Indiana), Harvard 
University, Illinois State Museum, IIli- 
nois State University, Iowa State Uni- 
versity, H. R. Lawson Collection (West 
Lafayette, Indiana), Michigan State 
University, Naturhistoriska Riksmu- 
seum (Stockholm), Northern Illinois 
University, Ohio State University, Pur- 
due University, Southern Illinois Uni- 
versity, United States National Mu- 


Intrnois NaTurRAL History Survey BULLETIN 


Vol. 31, Art. 


seum, University of Arkansas, Unie 
versity of California (Davis), Univer-’ 
sity of Illinois, University of Indiana,\ 
University of Kansas, University of Ken-, 
tucky, University of Michigan, Uni- 
versity of Minnesota, University of Mis- 
souri, University of Wisconsin, Western; 
Illinois University, and Winona State: 
College. 4 


NATURAL HISTORY 


Feeding . 

In the Bittacidae, adults of Bitta 
and Apterobittacus are predaceous.; 
Hanging by their fore or, occasionally, 
middle legs from the underside of vege-' 
tation, they wait with outstretched hind’ 
legs for some unsuspecting prey. When) 
prey is within reach, it is seized by ther 
raptorial tarsi of the hind legs. Ther 
prey is brought to the mouth, and ther 
piercing mouthparts enter through thet 
intersegmental membranes. The soft’ 
body parts of the victim are withdrawn, | 
and the empty exoskeleton is discarded. | 
Bittacus feed on a wide variety of in- 
sects. In Illinois Bittacus apicalis, B.! 
strigosus, and B. pilicornis feed heavily 
on dolichopodids (Diptera). Setty; 
(1931 and 1940) and Newkirk (1957) | 
listed a wide range of insects that! 
Bittacus accept, noting a preference for | 
Diptera and Homoptera. 


i; 
iY 
f 
‘| 
fy 


The time required for feeding varies ‘ 
considerably. Setty (1931) reported’ 
the average time as 20 minutes although! 
feeding sometimes lasted as long as‘ 
40 or 50 minutes. Newkirk (1957) re- 
ported that the feeding of Bittacus: 
apicalis may last an hour. He gave a 
detailed account of B. apicalis feeding 
on aphids: 

The hangingfly regurgitates a dark- 
brown fluid, which resembles the + 
“tobacco juice” of a grasshopper,” 
and covers a part of the aphid with | 
it. Through this the hangingfly 
bites, and sucks out the aphid body 
fluid. Then the hangingfly injects 
saliva, kneads what is left in the 
aphid body cavity with its man- 
dibles, draws off the mixture; re-~ 


| Aug., 1975 


: 
j 
| 


peats this several times; and dis- 
cards the empty exoskeleton. 

Very young larvae of bittacids are 

relatively active, but older larvae move 


very little (Setty 1940) and can be 


found among ferns and moist leaf litter 


/ in humid lowland woods. They feed 


on dead or dying animal matter, and it 
is not known if they can catch live 
prey. 

Little is known of the feeding habits 
of Boreus. Withycombe (1922) ob- 
served the larvae and Fraser (1943) 
the adults of Boreus hyemalis feeding 
on moss. Other substances may also be 
consumed. In Illinois, Boreus lives in 
Atrichum angustatum and_ probably 
feeds on it. 

Nothing is known of the feeding 
habits of the family Meropeidae. 

The feeding habits of the Panorpidae, 
in particular Panorpa, have been vari- 
ously reported in the literature. Lyonnet 
(1742) initiated the misconception that 
Panorpa are predaceous when he saw 
a fly the size of a scorpionfly attack a 
damselfly and bring it to the ground. 
Kirby & Spence (1823) repeated Lyon- 
net’s description and asserted that the 
species involved was Panorpa com- 
munis. Since then, numerous authors 
(Brauer 1863; Byers 1963; Campion & 
Campion 1912; Felt 1895; Lucas 1910; 
Miyaké 1912; Shiperovitsh 1925; and 
Syms 1934) have published observa- 
tions on panorpids’ feeding, and none 
has been found to be predaceous. 
Panorpids feed primarily on dead or dy- 
ing insects although Carpenter (1931b) 
reported their feeding on the nectar of 
flowers, and Miyaké (1912) saw them 
feeding on the petals of sweet william. 

Larvae of Panorpa feed principally 
on dead or dying animal matter, but 
Felt (1895) reported larger larvae of 
Panorpa attacking and devouring 
smaller ones. 


Mating and Oviposition 

Setty (1940) and Newkirk (1957) 
gave detailed descriptions of the mating 
of Bittacus. The description here is 
a compilation of both. The male seizes 


Wess Et At.: MECOPTERA OF ILLINOIS 


253 


a prey and flies from leaf to leaf in 
search of a female. When at rest, he 
vibrates his hind wings, opens and 
closes his claspers, and bends his abdo- 
men vertically, everting and inverting 
his abdominal sacs. Both male and 
female hang by their fore legs facing 
each other, and the male offers the prey 
to the female, which she eats during 
mating. In some instances the female 
jabs with her mouth at the male ab- 
dominal tip, where the eversible sacs 
are located, or at the prey. The male 
secures her abdomen in his claspers, 
then moves along the ventral surface 
to the terminalia. Only the female feeds 
during copulation. The length of cop- 
ulation is proportional to the palatabil- 
ity of the prey and lasts from 1 to 25 
minutes. When copulation is completed, 
the abdominal tips separate, and the 
individuals jerk at each other to dis- 
entangle the legs. Both male and fe- 
male may mate more than once. 


During oviposition the female rests 
on the ground with her head bent down 
and legs sprawled outwards. The body 
is quite rigid and the tip of the ab- 
domen is inserted into cracks in the 
soil. Oviposition takes from 5 to 30 
minutes, and several eggs are laid at 
a time. The female may fly from place 
to place and lay a few eggs in each. 
Oviposition occurs during the day or 
night. In captivity females tend to lay 
eggs randomly on the soil surface rather 
than in some place of concealment. 

In the Boreidae the mating behav- 
ior of the European species Boreus 
hyemalis has been reported by several 
authors (Brauer 1855; Lestage 1920; 
Steiner 1937; Stitz 1908; Syms 1934; 
and Withycombe 1922). Cockle (1908) 
described the mating of B. californicus. 
Carpenter (1936), Crampton (1940), 
and Cooper (1940) described the mat- 
ing of B. brumalis. The description 
given here is based on the observations 
of Cooper (1940) on B. brumalis. 


The male approaches to within 10 
mm or so of the female, and both re- 
main momentarily stationary. The male 
may show his excitement by slowly 


254 Inuinois NaturaL History 
waving his antennae or twitching his 
claspers and wings. He springs at the 
female with his claspers in advance, 
seizing the antenna, tibia, or tarsi of 
the female. The female becomes im- 
mediately passive, and the male seizes 
her about the body with his modified 
wings. Once the female is securely 
gripped with his wings, the.male em- 
ploys his hind legs and claspers to right 
the female and move her venter across 
his back until his terminalia clasp her 
apical abdominal segments. The eighth 
sternum of the female is pried down 
by the male’s claspers, which are in- 
serted into a pair of pockets on the 
male’s ninth tergum. The male releases 
his wings from the female, and she then 
flexes her rostrum between her coxae, 
folds her antennae between her legs, 
and stretches her legs posteroventrally. 
Once the female is in this position, the 
male grips her profemora and rostrum 
with his clasping wings. This position 
is maintained throughout copulation of 
1-12 hours. The male may run about 
and feed during copulation, while the 
female remains motionless. This pat- 
tern of behavior follows closely ob- 
servations made on B. californicus and 
B. hyemalis. 


According to Carpenter (1931b), 
Boreus lays eggs one or two at a time 
at the bases of moss clumps. Nothing 
has been reported on the mating be- 
havior or oviposition of the Meropeidae. 


In Panorpa mating is relatively simple 
(Miyaké 1912). The male vibrates his 
wings as he approaches the female. The 
apex of the abdomen is extended with 
the claspers securing the abdomen of 
the female. The claspers are moved 
along the abdomen until the terminalia 
are reached and the individuals are at 
an acute angle to each other. In addi- 
tion, Mickoleit (1971b) noted the use 
by P. communis of the notal and post- 
notal organs as pincerlike devices for 
holding the costa of the female during 
copulation. Copulation lasts for 15 min- 
utes to several hours. Although the 
mating behavior of Panorpa is simple, 


SurvEY BULLETIN Vol. 31, Art. 7 
there is one peculiarity that has led 
to some controversy. Mercier (1915) 
noted that prior to copulation in P. 
germanica, P. alpina, and P. cognata the 
male was seen to emit from its mouth 
a drop of fluid that hardened into an 
opaline pellet, which it placed on the 
soil. The female then fed on the pellet 
during copulation. When the pellet 
was consumed, another was produced. 


Shiperovitsh (1925) observed males of © 
P. communis emitting cylindrical pellets — 
from their mouths, and Gassner (1963) 
noted that unfed specimens of P. nup-— 
tialis regurgitated a brownish secretion — 
on which the female fed during coitus. — 
Syms (1934) observed no pellets being 


released but noted that the female fed 
on a dead insect during mating. Car- 


penter (1931b) observed the mating of — 
several species of Panorpa but never — 


saw such feeding behavior. One of us 
(Penny) has observed the depositing 


of salivary pillars by P. speciosa, P.— 


nuptialis, P. anomala, and P. helena. 


| 
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4 
a 
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4 
F 
7 
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Byers (1963) observed no salivary se- — 
cretion being produced by the male — 


of P. nuptialis although Gassner (1963) 
did observe this phenomenon. In ob- 


serving the mating of P. sigmoides, — 
Webb saw no evidence of a salivary se- 


cretion or pellet being offered by the — 
male, nor did the female feed during — 


copulation. In the field P. sigmoides 
was also observed to mate during the 


hours of daylight. Most authors have — 


observed mating during the hours of 
darkness, but Byers (1963) found P. 
nuptialis to mate only during the day- 
light hours. 


During oviposition the female probes 


the surface of the soil for an appropriate — 


crevice, and the abdomen is extended 
and inserted deeply into the soil. The 


number of eggs laid at one time varies. — 


Immature Stages 


In Bittacus the size and shape of the 
eggs vary considerably among the spe- 
cies. The eggs range in length from 
0.56 to 0.72 mm and in width from 
0.41 to 0.65 mm (Setty 1940). 


Aug., 1975 


In B. apicalis the eggs are oval (Fig. 
1) or spherical and have a finely re- 
ticulated surface. In B. punctiger, B. 
strigosus, B. occidentis, B. stigmaterus, 
and B. pilicornis, the egg shape varies 
from cuboidal to heptahedral, and the 
egg has a shallow depression on each 
side (Fig. 2). The surface is rough 
and has numerous small protuberances. 

Prior to hatching, the eggs become 
spherical and increase in size (Setty 
1940). B. punctiger and B. pilicornis 
eggs hatch within 2 weeks, and the im- 
matures overwinter as larvae. B. strigo- 
sus, B. apicalis, and B. stigmaterus pass 
the winter in the egg stage. 

The newly hatched larva emerges 
through an irregular crack in the wall 
of the egg and feeds on the remnants 
of the egg shell. The larvae do not 
burrow through the soil in search of 
food, but the older larvae lie motion- 
less on the surface among the leaf litter 
and ground debris. The larvae pass 
through five instars before pupating 
(Setty 1940). 

The larvae of Bittacus (Fig. 3) are 
cylindrical and range in length from 
11 to 14 mm in the last instar. The 
heavily sclerotized head is broad an- 
teriorly. In lateral view the head is 
oval or elliptical. It is generally bent 
under the body so as to be completely 
hidden from above by the thorax. The 
antennae are short and stout and have 
only two segments. The single median 
ocellus is present as well as two large 
lateral eyes, which are not true com- 
pound eyes, according to Setty (1931 
and 1940), but simply a group of sev- 
eral ocelli. The mandibles are large 
and heavily sclerotized and bear several 


1.—B. apicalis. 


Fig. 1—2.—Bittaeus eggs. 
2.—B. strigosus. 


Wess Er At.: MECOPTERA OF ILLINOIS 


255 


Fig. 3.—Bittacus strigosus larva. 


large teeth. The labial and maxillary 
palps are short and stout and have two 
and four segments, respectively. The 
head bears numerous coarse setae and 
tubercles. Each of the three thoracic 
segments bears a pair of sharply pointed 
legs, and each of the first nine abdomi- 
nal segments possesses a pair of short 
ventral prolegs. The last abdominal 
segment bears a ventral protrusible 
sucker that aids in locomotion. The 
dorsal and lateral margins of the thorax 
and abdomen bear several simple or 
branched protuberances, each with a 
simple or clavate apical seta. Indi- 
viduals collected in the field usually 
are covered with soil which clings to 
these setae and protuberances. 


The larvae are negatively phototropic 
and prefer moist shaded areas. Prior 
to pupation the fourth instar larva bur- 
rows into the soil, forming a diagonal 
cylindrical chamber (Setty 1940). The 
larva constructs a collar around the 
opening with a thin layer of soil laid 
across it. At this time the larva molts 
to form a prepupa. The prepupa re- 
mains in the bottom of the chamber, 
for 9-18 days in the case of B. punctiger 
(Setty 1940), following which it meta- 
morphoses into a pupa. 

In the case of B. punctiger, the pupa 
remains in the chamber for 13-20 days 
(Setty 1940), after which the adult 
emerges through the opening that the 
larva had entered. 

Setty (1931, 1939, 1940, and 1941) 
has done extensive work on the mor- 
phology and behavior of the North 
American species of Bittacus, and much 
of the description of the immature 
stages presented here was extracted 
from his publications. 


256 


In North America the complete life 
history of Boreus has not been pub- 
lished for any species. The description 
presented here is for B. hyemalis, as 
described by Withycombe (1922 and 
1926). 


The eggs of Boreus are about 0.5 mm 
long and 0.3 mm wide. They are laid 
at the base of moss, and the larvae 
hatch in about 10 days, usually in late 
fall. The larvae pass through four in- 
stars, a mature larva (Fig. 4) being 
6-7 mm long. The head is pale yellow 
and heavily sclerotized. The eyes are 
small and composed of several small 
facets. Mandibles are large, dark brown, 
and heavily sclerotized. Antennae are 
small and have two segments and a 
fine apical bristle. Labial palps are 
small. The thorax is pale white and 
broad and has three pairs of ventro- 
laterally extended legs. The legs have 
three segments, the basal segment be- 
ing broad and the others tapering to 
a small, acute apical segment. The 
abdomen is pale white and without 
lateral appendages and has the apex 
rounded. Each segment has several 
fine setae. 


The larvae appear to aestivate 
throughout the summer in small cells 
made in compacted soil in which they 
pupate in late fall. The duration of 
the pupal stage is 4-8 weeks. 

Nothing is known of the immature 
stages of the Meropeidae. 

In Panorpa the size and characteris- 
tics of the egg vary considerably. In 
the lugubris group the eggs of P. nup- 
tialis are spherical or oval with a 
smooth surface and measure about 1.07 


Fig. 4.—Boreus brumalis larva. 


Inuino1is NaTurAL Hisrory SuRvVEY BULLETIN 


Vol. 31, Art. 7 


mm in length and 0.84 mm in width 
when laid (Byers 1963). In the rufes- 
cens group the eggs of P. helena are 
oval, have a fine network of depressions 
covering the surface, and measure about. 
1.10 mm in length and 0.65 mm in 
width. Felt (1895) described the eggs 
of P. debilis (as P. rufescens) as ellipti- 
cal and oval, 0.625 mm long, and 0.6 
mm wide. Numerous authors (Brauer 
1852; Byers 1963; Felt 1895; Syms 
1934; and Yie 1951) have observed that 
the color of the egg darkens before 
hatching. The duration of the egg pe- 
riod is about 8 days for P. nuptialis’ 
(Byers 1963) and 6-7 days for P. 
debilis (Felt 1895). 


Gassner (1963) observed an egg 
burster on the frons of the first instar 
of P. nuptialis. It is used in rupturing 
the chorion of the egg. According to 
Gassner, the larva assumes a flattened 
spiral position prior to hatching. It 
expands and forces the egg burster 
through the chorion. The larva then ~ 
makes a quarter turn and slices open 
the shell. 


The larva of Panorpa (Fig. 5) is” 
elongate and cylindrical. It passes 
through four larval instars before pupa- — 
tion (Boese 1973; Byers 1963; Mampe 
& Neunzig 1965; Shiperovitsh 1925; i 
Yie 1951). Based on measurements of © 
head width, Felt (1895) reported P. E 
debilis (as P. rufescens) as having 
seven larval instars, as Miyaké (1912) 
reported for P. klugi. Carpenter (1931la)_ 
also described Panorpa as having seven 
instars. The antennae are short and 
stout and have a scape, a pedicel, and 
one flagellar segment. The eyes are 
composed of 25 or more facets. The 
mandibles are large and heavily sclero-_ 
tized and have two to four mesal teeth. — 


The thorax bears a pair of short i 


Fig. 5.—Panorpa sp. larva. 


Aug., 1975 


pointed legs on each segment and a 
thick sclerotized pronotal shield. A 
single pair of spiracles is present on 
the pronotal segment. The thorax and 
the abdomen bear numerous setigerous 
prominences (pinacula) and unmodi- 
fied setae. The eighth and ninth ab- 
dominal segments each possess a pair 
of annulated setae borne on moderately 
sclerotized projections and a single an- 
nulated seta on segment 10. A pair of 
prolegs and a lateral spiracle are pres- 
ent on abdominal segments 1-8. Four 
translucent, retractible anal lobes and a 
basal fold of skin comprise the 11th 
segment. 

Byers (1963) reported in detail on 
the life history of P. nuptialis, from 
which much of the information pre- 
sented here has been taken. Boese 
(1973), Felt (1895), and Mampe & 
Neunzig (1965) have described other 
North American larvae. Several authors 
have described the immature stages of 
European and Asian panorpids (Brauer 
1863; Miyaké 1912; Shiperovitsh 1925; 
Steiner 1937; and Yie 1951). 

After hatching, the larvae burrow far- 
ther into the soil and feed primarily 


Wess Et Au.: MECOPTERA OF ILLINOIS 


257 


on decaying organic matter although 
Felt (1895) reported some larvae as 
being predaceous. 

The larvae spend 4-5 days in each 
of the first three instars and are active 
and feed for about 2 weeks in the 
fourth instar, following which the full- 
sized larvae become quiescent and con- 
struct prepupal cells. The prepupal cell 
is oblong with rounded ends and is 
formed in compacted soil. The cell is 
about as long as the larva but possesses 
no visible lid, like that noted by Yie 
(1951) in Formosan panorpids. The 
larvae then enter a prepupal or qui- 
escent stage, which carries them through 
the winter. 

The duration of the pupal stage varies 
from 6 to 21 days. Prior to emergence 
the pupal skin splits along the dorsal 
midline, and the adult emerges. The 
hour of emergence is dependent upon 
the species. Yie (1951) found that in 
Formosan panorpids emergence oc- 
curred most often in the early morning. 


Habitat 


In the Bittacidae most species are 
restricted to the humid, well-shaded 


Fig. 6.—Herbaceous vegetation in lowlands along the Illinois River, Starved Rock State 
Park, Illinois. (Photo by H. H. Ross, courtesy of Section of Botany and Plant Pathology, IIli- 
nois Natural History Survey) 


258 InLinois NaturaL History Survey BULLETIN Vol. 31, Art. 7 : 


Fig. 7.—Deciduous forest and herbaceous vegetation along creek bed at Trestle Hollow, 
Fountain Bluff, Jackson County, Illinois. (Photo by W. D. Zehr, courtesy of Section of Botany 
and Plant Pathology, Illinois Natural History Survey) 


Fig. 8.—Bittacus apicalis hanging from herbaceous vegetation. (Photo by W. D. Zehr) 


\ug., 1975 Wess Er At.: MECOPTERA OF ILLINOIS 259 


reas along streams and in bottomlands and 9) can be found hanging from the 
Fig. 6 and 7). Individuals (Fig. 8 undersides of leaves of jewelweed (Im- 


Fig. 9.—Bittacus pilicornis hanging from herbaceous vegetation. (Photo by W. D. Zehr) 


260 Inutinois NaturAL History SurvEY BULLETIN Vol. 31, a 


patiens sp.), stinging wood nettle (La- 
portea canadensis), gooseberry (Ribes 
sp.), and a variety of other bottomland 
plants. Bittacus strigosus has the widest 
range of habitats, extending from the 
moist bottomland areas to the drier hill- 
side areas and occurring predominantly 
on multiflora rose (Rosa multiflora). 
In western Illinois B. strigosus was col- 
lected abundantly in short pasture grass 
in the shade of poplars (Populus sp.). 
Little is known of the habitat for 
B. occidentis. Most of the individuals 
collected have been taken at lights. 

In the Boreidae the various species 
are highly restricted in habitat. Speci- 
mens are collected only in, or very 
close to, patches of moss on the ground 
(Fig. 10). In southern Illinois B. 
brumalis lives in Atrichum angustatum 
and Dicranella heteromalla. 

Of the habitat of the Meropeidae 
little is known. The majority of speci- 
mens have been collected in a variety 
of hardwood forests but mostly at lights 
or in Malaise traps. Occasionally indi- 


“¥ 


| AN gave aie 


a 


tata 


M4} 
AS 


; 
viduals have been found under stones ~ 


or rotting logs. 
The habitats of the Panorpidae are 
similar to those of Bittacus. Individuals 
of Panorpa (Fig. 11) are most com- 
monly collected as they rest on the 
leaves of stinging wood nettle, poison 
ivy (Rhus radicans), waterleaf (Hydro- 
phyllum appendiculatum), jewelweed, 
and a variety of other broad-leaved 
plants. Only members of the lugubris” 
group shun the shaded humid areas 
along streams and are found in the 
short grasses along roadside ditches or 
in cotton, tobacco, and soybean fields. 


DISTRIBUTION 
AND DISPERSAL 


The order Mecoptera is one of the 
most generalized groups of holometab- 
olous insects and has an abundant fossil 
record dating back to the early Permian 
(Tillyard 1935). 

The Bittacidae are the most highly 
specialized family of the Mecoptera. 


a 


Fig. 10.—Patches of moss on a hillside in Lake Murphysboro State Park. (Photo by L. J. 


Stannard) 


Aug., 1975 


Wess Et At.: MECOPTERA OF ILLINOIS 


261 


Fig. 11.—Panorpa sp. on herbaceous vegetation. (Photo by W. D. Zehr) 


Their tipulidlike appearance, single 
raptorial claw on the tarsus, and pre- 
daceous habit are three of the most sig- 
nificant specializations. Although bit- 
tacids have the bulbous basistyles of 
most of the Mecoptera, the presence 
of a four-branched sector vein and the 
absence of a notal organ suggest that 
this family’s specialization began at an 
early date. Jurassic fossils of Probit- 


tacus and Protobittacus (Tillyard 1935) 
also suggest early specialization. 


In the Nearctic Region the Bittacidae 
are represented by two genera, Bittacus 
and a wingless form, Apterobittacus. 
Apterobittacus is monotypic and found 
only in central California (Fig. 12) 
except for one doubtful record from 
southwestern Colorado. Bittacus, the 
most widespread genus of the Mecop- 


262 


Fig. 12.—General distribution of Bittacus (dots) and Apterobittacus (lines) in the Nearc- 


tic Region. 


tera (Fig. 12), extends from northern 
Florida to Quebec, west to eastern 
Montana, then south to Mexico, and 
an isolated species (B. chlorostigma) is 
restricted to California and Oregon. 


The spread of the Bittacidae into the 


Nearctic Region (Byers 1969) possibly - 


occurred during the late Mesozoic or 
early Tertiary, following the emergence 
of the Bittacidae prototype on the 
former southern land mass, Gondwana- 
land. All bittacid genera, except Bit- 
tacus and two apparently recent flight- 
less derivatives of Bittacus, are re- 
stricted to Australia and South and 
Central America. 


After the establishment of land con- 
nections between North and South 
America, Bittacus dispersed northward 
and is known from North American 
Eocene fossils (Carpenter 1955). Glaci- 
ations during the late Pliocene or early 
Pleistocene then forced the bittacids 
into the southern United States, Mexico, 
and South America (Byers 1969). After 
the glaciations the bittacids in the 
southeastern United States became sep- 


Inuino1is NaturAL History SurRvEY BULLETIN 


Vol. 31, Art. 7 


arated from the main bittacid stock in 
Central America by xeric conditions 
and the disappearance of mesic forests 
from northern Mexico and the South- 
west. Following the retreat of the gla- 
ciers, the southeastern bittacids spread 
northward and westward, and a second 
invasion from Mexico brought B. chlo- 
rostigma to California and B. texanus 
to the Southwest. 


Illinois forms the northwest border 
of the distribution of B. apicalis (Fig. 
43) and B. punctiger (Fig. 44). B. 
stigmaterus, B. pilicornis, and B. strigo- 
sus occur throughout Illinois and extend 
into the west-central states. B. occi- 
dentis has been collected only in central 
and northern IIlinois although it is wide- — 
spread from southern Ontario and New 
York southwestward to Arizona. Of 
the midwestern species, only B. texanus 
has not been recorded from Illinois 


The other three North American fam- 
ilies (Boreidae, Panorpodidae, and 
Panorpidae) are all confined to the 
temperate and boreal forests of the 
northern hemisphere. All have bulbous 


Aug., 1975 


Wess Ev Au.: MrEcopTerA OF ILLINOIS 


263 


Fig. 13.—General distribution of the Boreidae in the Nearctic Region. 


basistyles and pincerlike dististyles. 
Each has survived in a slightly different 
climatic zone. 


The Boreidae are found primarily in 
the colder regions of the northern 
hemisphere from St. Paul Island in 
the Bering Sea to 12,000 feet in the 
Colorado Rockies (Fig. 13). In eastern 
North America the family has spread 
northward from the southern Appa- 
lachians, leaving relict populations in 
marginal habitats in the southern por- 
tions of its range. Adaptations to cold 
environments include reduction in size, 
loss of flight and reduction in wing 
size, and loss of the notal (wing-clasp- 
ing) organ. 

The family Meropeidae is the most 
primitive family of the Mecoptera in 
North America. The broad wings with 
numerous costal crossveins, the short 
rostrum, and the elongate male ba- 
sistyles and dististyles indicate the prim- 
itive nature of this family. The recent 
distribution of Merope (Fig. 14) (Byers 
1973b) indicates the center of specia- 


Fig. 14.—General distribution of the Mero- 
peidae in the Nearctic Region. 


tion to be in the southern Appalachians, 
from which area this genus has dis- 
persed northward and to the east and 
west. Although widespread in the north- 
eastern United States, records of this 
genus are sparse. In Illinois Merope 
has been recorded only from Pine Hills 
Ecological Area and Urbana. 


264 Inyinois NaTurAL History 

The family Panorpodidae is found in 
boreal environments of montane areas 
of the southern Appalachian and the 
northwestern states (Fig. 15). Normally 
this family is distributed in cool areas 
from sea level to higher elevations in 
North America. Adaptations to such 
boreal environments include the flight- 
lessness of females and the loss of the 
male notal organ. 


The Panorpidae normally live at 
lower elevations than do the Boreidae 
and Panorpodidae, but ranges may 
broadly overlap. Species of the Panorpi- 
dae and Panorpodidae from Japan have 
almost identical wing venation; the 
North American Panorpodidae (Bra- 
chypanorpa ) have a reduced number of 
sector branches. The male genitalia of 
the Panorpidae and Panorpodidae are 
also very similar, indicating a close re- 
lationship between these two families. 
However, Oligocene Baltic amber has 
yielded specimens of both Panorpa and 
Panorpodes so different that these fam- 
ilies must have diverged before the 
Oligocene. 


bd 


SurveEY BULLETIN Vol. 31, Art.7 

The majority of Nearctic Panorpidae 
are distributed in the eastern United 
States, and several species are recorded 
from the Southwest and Mexico (Fig. 
16). Byers (1969) partitions the genus 
Panorpa north of Mexico into six dis- 
tributional groups: 

1. Those species occurring only in 
the southern Appalachians. This group 
contains five species found only at the 
middle to higher elevations. 


2. Those found in the southern Ap- 
palachians but also distributed widely 
to the northeast, northwest, and west. 
This group contains eight widely dis- 
tributed species. All extend into the 
Midwest, and four species occur in 
Illinois (P. banksi, P. debilis, P. helena, 
and P. nebulosa). 


3. Those occurring primarily in the 
Piedmont and sometimes up into the 
valleys of the Appalachians. Species in 
this group occur principally on the east- 
ern side of the Appalachians although 
both P. consuetudinis (= P. elaborata) 
and P. rufescens extend into the Mid- 
west. 


Fig. 15.—General distribution of the Panorpodidae in the Nearctic Region. 


| Aug., 1975 Wess Et At.: MEcopTEeRA OF ILLINOIS 265 


Fig. 16.—General distribution of the Panorpidae in the Nearctic Region. 


4, Those inhabiting the coastal plain 
from Texas eastward to Florida and 
northeastward approximately to New 
Jersey. This group contains seven 
species, only one of which (P. nup- 
tialis) extends northward into the Mid- 
west and is known from within 1 mile 
of Illinois near Cairo. 


5. Those occurring primarily or 
wholly in the formerly glaciated area of 
the northern Appalachians and west- 
ward. The five species in this group all 
occur in the Midwest to the north and 
east of Illinois. 


6. Those found only in the Midwest. 
This group contains six species, four of 
which (P. anomala, P. dubitans, P. 
speciosa, and P. sigmoides) occur in 
Illinois. 


Most species of Panorpa inhabit 
mesic temperate forests with humid, 
dense undergrowths of herbaceous veg- 
etation. During periods of glaciation 
in North America these species possibly 
sought areas of relatively stable climatic 
conditions (Byers 1969), such as those 
in the southern Appalachian and Ozark- 
Ouachita uplift. During interglacial 


periods the species spread northward. 
The southern Appalachian area has 
the greatest concentration of Panorpa 
species in North America. All the 
species in Byers’ groups one through 
five and some in group six appear to 
have arisen from a southern Appala- 
chian ancestral stock and migrated 
northward and westward. In group six 
Byers lists six species which occur only 
in the Midwest. Judging from their 
present distributions, one can infer that 
three of them (P. anomala, P. speciosa, 
and P. braueri) may have differentiated 
in the area of the Ozark-Ouachita 
uplift. 


COLLECTING AND 
PRESERVING MECOPTERA 


With the exception of the Boreidae, 
the Mecoptera are generally found on, 
or hanging from, low herbaceous vege- 
tation in shaded moist woodlands. Bit- 
tacus can be found by walking slowly 
through shaded weedy areas and brush- 
ing the vegetation back and forth with 
a net. When disturbed, bittacids will 
fly 10-20 feet ahead of the collector and 


266 


then hang from the vegetation again. 
The experienced collector may net spec- 
imens in flight or follow their flight and 
collect them as they hang from the 
vegetation. Some bittacids (B. apicalis 
and B. occidentis) have been collected 
at lights. 


The collecting of Boreus calls for a 
rather hardy, determined collector, be- 
cause these insects reach maturity dur- 
ing late fall and winter. They are asso- 
ciated with mosses on the ground, on 
bases of trees, and elsewhere. They can 
be collected by lying beside a patch 
of moss and waiting for the adults to 
move. They also move about on patches 
of snow, where they are easily seen and 
collected. Larvae of Boreus have been 
taken by Berlese funnel extraction from 
moss. 

The collecting of Merope has been 
accomplished more by chance than by 
skill. Most specimens have been col- 
lected at lights or in Malaise traps in 
heavily wooded areas. 


Panorpa can be collected individually 
from the surface vegetation. The col- 
lector must stalk slowly through the 
vegetation, particularly stinging wood 
nettle, until an individual is located. 
When disturbed, the somewhat seden- 
tary members of this genus will fly a 
short distance or drop to the ground 
and escape in the leaf litter. Panorpa 
is seldom taken at lights. 

Specimens of Mecoptera can be pre- 
served in 70-percent ethyl alcohol or 
mounted on insect pins. 

The taxonomic characters necessary 
to separate the genera and many of the 
species can be seen with a stereoscopic 
microscope. In the females of Panorpa, 
the genital plate is of taxonomic impor- 
tance. To observe this plate, one must 
cut off the tip of the abdomen basal to 
the eighth segment and boil the tip in 
10-percent KOH or leave it overnight 
in cold 10-percent KOH to remove the 
soft internal tissues. The tip is trans- 
ferred to 70-percent ethyl alcohol, and 
the abdominal terga and sterna are sep- 
arated with a pair of dissecting points, 


Initino1is NaturaL History SurvEy BULLETIN 


Vol. 31, Art. 7 


revealing the genital plate. In identi- 
fying males of some species of Panorpa, 
clearing the genital bulb in 10-percent 
KOH aids in species determination. : 


MORPHOLOGY ; 


Several excellent papers have been 
published on the external and internal _ 
anatomy of the Mecoptera (Crampton — 
1921 and 1931; Dohanian 1915; Grassé 
1951; Hepburn 1969 and 1970; Micko- 


Fig. 17.—Panorpa helena anterior view of 
head. 


Fig. 18-19.—Bittacus strigosus. 18.— 
Apical tarsal segments with claw. 19.— 
Apical tarsal segments with claw reflexed. 


| 
i 
| 
) 


Aug., 1975 


COMPOUND 


EN e 
OCELLI 


ANTENNA — % 


Ss 
MAXILLARY 
PALPS 


“~TARSUS 


leit 1971a; 
1938). 
The descriptions are supplemented 
with illustrations of the morphological 
characters of taxonomic importance. 
Fig. 17 presents an anterior view of 
the head of Panorpa, showing the dis- 
tinctive elongate rostrum of most of the 
Mecoptera. Ocelli are present in all 
genera of the North American Mecop- 
tera except Merope. In Boreus the 


Otanes 1922: and Potter 


Wess Et Au.: MECOPTERA OF ILLINOIS 


267 


FORE WING 


20 


Fig. 20.—Panorpa sp. lateral view of male adult. 


ocelli are indistinct, and numerous au- 
thors have reported them absent. In 
all genera the large, lateral compound 
eyes are widely separated, except those 
of Merope, which are reniform and al- 
most contiguous dorsally. 

The shape and venation of the wings 
vary from genus to genus. Fig. 24, 50, 
64, and 80 illustrate the wings of all 
midwestern genera. In Apterobittacus 
the wings are absent, and in certain 


Fig. 21A-B.—Panorpa sigmoides. A.—Ventral view of male terminalia. 


VP, ventral paramere. BS, basistyle. 
DS, dististyle. BS, basistyle. 9T, ninth tergum. 


HY, hypandrium. 


DS, dististyle. 
B.—Dorsal view of male terminalia. 


268 Inuinois NaTuRAL History 
species of Brachypanorpa the females 
have greatly reduced wings. 


The legs in all genera are elongate 
and cylindrical. In Panorpa the apical 
tarsal segment bears a pair of serrate 
claws. In Bittacus and Apterobittacus 
the tarsi have a single apical claw (Fig. 
18 and 19), which reflexes back into a 
groove in the fourth tarsal. segment. 
This claw is used in holding prey. 


Fig. 20 is a lateral view of a Panorpa 
male and illustrates the scorpionlike 
appearance of the genus. The abdomen 
is thick and rounded basally and tapers 
apically to the elongate seventh and 
eighth segments. The terminalia are 
bulbous and reflexed over the abdomen. 
In Panorpa the sixth abdominal tergum 
in males may or may not possess an 
anal horn. 


The terminalia of Panorpa are shown 
in Fig. 21A and 21B, and the morpho- 
logical characters of taxonomic impor- 
tance are identified. 


MONOGRAPHS ON 
NEARCTIC MECOPTERA 


Because of the small number of spe- 
cies of Nearctic Mecoptera, few major 
taxonomic revisions have been done on 
this group. Westwood (1846) in his 
monograph on the genus Panorpa de- 
scribed several Nearctic species. Walker 
(1853), Hagen (1861), Banks (1907), 
and Esben-Petersen (1915) catalogued 
the North American Mecoptera, and 
Hine (1898 and 1901) reviewed the 
Mecoptera north of Mexico. In 1908 
Sherman reported on the Panorpidae 
of North Carolina; Engelhardt (1915), 
the Mecoptera of the northeastern 
United States; and Esben-Petersen 
(1921), the North American species. 
The major revision of the Nearctic 
Mecoptera was published by Carpenter 
(193la) wherein he described many 
new species. Since then new species 
have been described and additional dis- 
tribution data have been reported by 
Carpenter (1932a, 1935, 1936, and 
1939) and Byers (1954, 1958, 1962a, 
1962b, and 1973a). 


SuRvEY BULLETIN Vol. 31, Art. 


TAXONOMIC TREATMENT 


Order MECOPTERA 


MECOPTERA Comstock & Comstock 
1895 : 


MECAPTERA Packard 1886 
PANORPATAE Brauer 1885 


The members of the order Mecoptera 
are moderately large, holometabolous 
insects, having biting mouthparts gen- 
erally extended ventrally to form a pro- 
longed rostrum. The antennae are 
elongate and filiform and have about 
20 flagellar segments. The large com- ~ 
pound eyes are dichoptic. Ocelli are — 
present or absent. The maxillary palps ; 
have five segments. 4 


The thorax is broad dorsally and 
tapered ventrally. The wings are usu-— 
ally elongate and narrow. The fore ¥ 
and hind wings are nearly equal in 
length and have numerous veins and 
crossveins. In several genera the wings — 
are greatly reduced or absent. In 
Merope and Notiothauma the wings are 
very broad and rounded apically. The 
legs are long and slender and have five 
tarsal segments, ending in one or two — 
claws. The coxae are large, and each © 
tibia bears a pair of long spurs. 


The first abdominal segment is fused 
to the thorax. The abdomen is gener- 
ally thick basally and tapered apically 
except in the Bittacidae. Cerci are 
present apically in females and subapi- 
cally in males. 


KEY TO THE NEARCTIC FAMILIES 
OF MECOPTERA 

1. Tarsi with single apical claw (Fig. 
19) Ore er eae Bittacidae 

Tarsi with two apical claws 

2. Male brachypterous. Female with 
OVipOSitOr ........e2+2--eeee Boreidae 

Male with elongate wings. Female 
without ovipositor 

3. Wings broad, rounded apically (Fig. — 
64), with numerous costal cross- 
veins. Ocelli absent ...... Meropeidae 


Wings narrow, elongate (Fig. 73), 
with few costal crossveins. Ocelli 
present 


4. Rostrum short 
Rostrum long (Fig. 17) 


abet ae Panorpodidae 
andoc Panorpidae 


' Aug., 1975 Wess Er At.: MECOPTERA OF ILLINOIS 269 


| BITTACIDAE Enderlein 1910 posed by Hine for the species B. 
/ strigosus and B. pilicornis. However, 


separate the bittacids from other fami- Fee ere oe ee 
lies of the Mecoptera. Twelve genera y ; : 
are distinguished, and their species are Thyridates Navas (1908:412). Synony- 
recorded from all continents although mized by Banks (1913). 
they are generally absent from the Diplostigma Navas (1908:413). Synony- 
northern parts of Europe, Asia, and mized by Banks (1913). 
North America. Bittacus is the most Haplodictyus Navas (1908:413). Syn- 
widespread genus, occurring in Europe, onymized by Banks (1913). 
Asia, Africa, and North and South Head small, pale to dark yellow, ta- 
America. Apterobittacus, found in Cali- pered ventrally to form distinctive ros- 
fornia, and Anomalobittacus from South trum. Eyes large. Ocelli large, amber, 
Africa are the only flightless genera. on raised subtriangular pad. Antennae 


Anabittacus, Nannobittacus, Neobitta- long, filiform with 14 flagellar segments. 
cus, Pazius, and Issikiella occur in 


South and Central America. Kalobit- 
tacus is recorded from Central America. 


The raptorial tarsi with a single claw 


Thorax broad, compressed laterally. 
Wings long, narrow, tapered basally. 


; ee Membranes clear or yellow, often with 
Austrobittacus, Edriobittacus, and Har- dark brown apex or crossveins, Sub- 


pobittacus occur only in Australia. costa ending in middle of wing. Sub- 


Of the two Nearctic genera, only costal crossvein (Fig. 24) usually basal 
Bittacus has been collected in Illinois. to first fork of radial sector. R, forked 


KEY TO THE NEARCTIC GENERA apically to form pterostigma, which 


OF BITTACIDAE has one or two pterostigmal crossveins. 

1. Wings present .............. Bittacus Pterostigma (Fig. 22) darker than sur- 
Wings absent ........... Apterobittacus rounding membrane. A whitish thyrid- 
ium (Fig. 26) around first fork of 

Apterobittacus MacLachlan media. Apical crossvein (Fig. 24) pres- 


Apterobittacus MacLachlan (1893: ent or absent. Legs elongate, slender, 
317). Type-species by monotypy. cylindrical. Coxae large, thick, tapered 
Apterobittacus apterus MacLachlan. apically. Femora generally slender al- 


though hind femora often swollen. 
ti ee ee ee Tibiae long, slender with two long 
a SNe se el ean asec cs . aenien “S SPUrS. Basal four tarsal segments cylin- 
8 Eyres : 8 drical with small apical enlargment; 
less. Legs similar to those of Bittacus. fifth segment fused to apical claw 
ae eee haw aoe eae which is reflexed to fit into groove in 
lateral view, broad, subrectangular, ex- fourth segment. 
tending well beyond apices of basi- Abdomen long, narrow basally. Male 
styles; in dorsal view, narrow, com- terminalia large (Fig. 29). Ninth ter- 
pressed laterally, apices converge. Ba- 8U™ modified to form two laterally flat- 
sistyles broad, thick, fused ventrally. tened claspers, often extending beyond 
Dististyles small. Aedeagus thick ba- @Pices of basistyles. Basistyles broad, 


Il P ‘call horn d_ fused ventrally, each with short, medi- 
a Sewanee ally extended dististyle. Aedeagus thick 


basally, tapering apically. Internal 
skeleton of female genitalia absent. 
Sternal region of eighth and ninth seg- 
: F ments fused to form subgenital plate. 
Bittacus Latreille Tenth segment bears pair of unseg- 

Bittacus Latreille (1805:20). Type- mented cerci. 
species: Bittacus italicus Miller. Seven species of Bittacus occur in 

Leptobittacus Hine (1898:108). Pro- the Midwest. 


This is a monotypic genus probably 
restricted to California. 


270 ItLinois NaturaAL History Survey BULLETIN 


Vol. 31, Art. 7 


Fig. 22-28.—Bittacus fore wings. 22.—B. apicalis. Pt, pterostigma. 23.—B. punctiger. 
24.—B. pilicornis. AC, apical crossvein. 25.—B. occidentis. ScC, subcostal crossvein. 26.— 
B. strigosus. Th, thyridium. 27.—B. stigmaterus. 28.—B. texanus. 


KEY TO THE MIDWESTERN 
SPECIES OF BITTACUS 


1. Apices of wings dark brown (Fig. 


CPD HATRED BIR T8 ARSE BORE ERS Oe apicalis 
Apices of wings not dark brown ... 2 
2. Apical crossvein present (Fig. 24).. 3 
Apical crossvein absent (Fig. 25)... 4 


3. Hind femora with brown spot sur- 
rounding base of setae ....punctiger 
Hind femora without brown spot sur- 
rounding base of setae ..... pilicornis 
4. Subcostal crossvein distal to first 
fork of radial sector (Fig. 25) .... 
ODO HORNS lo ei erenep tend Geel fee occidentis 
Subcostal crossvein basal to first fork 
of radial sector (Fig. 26) ........ 5 
5. Wing membranes colorless. Cross- 


veins margined (Fig. 26) ....strigosus 


Wing membranes yellow to pale 


brown. Crossveins usually not mar- 
gined (Big. 27) 2... J iin ve cee 6 


In males, lobe of ninth tergum in 
dorsal view with two medial prom- 
inences, each prominence bearing 
several black spines (Fig. 40). In 
females, wing color yellow to amber 
old DISS Rie tere etek ener stigmaterus 
In males, lobes of ninth tergum in 
dorsal view with one medial prom- 
inence bearing several black spines, 
and each lobe with a row of 10-15 
thick black spines basal to medial 
prominence (Fig. 42). In females, 
wing color brown to dark brown 
PITS A eI Scigu A te Se wine Osta texanus 


Aug., 1975 Wess Er AL.: MECOPTERA OF ILLINOIS 271 


42 


Fig. 29-42.—Bittacus male terminalia. 29.—B. apicalis. Lateral view of terminalia. 
St, ninth tergum. Ce, cerci. Bs, basistyle. Ds, dististyle. Ae, aedeagus. 30.—Dorsal view of 
ninth tergum. 31.—B. punctiger. Dorsal view of ninth tergum, 32.—Lateral view of ter- 
minalia. 33.—B. pilicornis, Lateral view of terminalia. 34.—Dorsal view of ninth tergum. 
35.—B. occidentis. Lateral view of terminalia. 36.—Dorsal view of ninth tergum. 37.—B. 
strigosus. Lateral view of terminalia. 38.—Dorsal view of ninth tergum. 39.—B. stigmaterus. 
Lateral view of terminalia. 40.—Dorsal view of ninth tergum. 41.—B. texanus. Lateral view 
of terminalia. 42.—Dorsal view of ninth tergum. 


272 


Bittacus apicalis Hagen 


Bittacus apicalis Hagen (1861:248). ¢, 
@. Type-locality: Southern Illinois. 
Haplodictyus incertus Navas (1926:59). 
8. Type-locality: Wilmerding, Penn- 
sylvania. Synonymized by Carpenter 

(1932b). 

Head and thorax pale glossy yellow 
to brown. 

Wings (Fig. 22) pale yellow, ptero- 
stigma and apex of wing dark brown. 
Subcostal crossvein basal to first fork 
of radial sector. One pterostigmal cross- 
vein. Apical crossvein absent. 

Legs pale yellow to brown. Hind 
femora slightly enlarged. 

Abdomen and terminalia pale yellow 
to brown, occasionally eighth tergum 
of males dark brown to black. In males, 
lobes of ninth tergum in lateral view 
(Fig. 29) extend slightly beyond apices 
of basistyles, dorsal margin with medial 
prominence; in dorsal view (Fig. 30) 
lobes diverge apically, curve ventrally, 
apex with 30 or more black spines. 
Basistyles broad, thick. Dististyles short, 
tapered apically. Aedeagus thick at 
base, tapered apically to slender coiled 
thread. Cerci short, slender, do not 
extend beyond middle of basistyles. 


The dark brown apices of the wings 
readily separate this species from all 
other Nearctic bittacids. When B. api- 
calis hangs from vegetation, the wings 
are extended laterally from the body 
(Fig. 8) rather than being folded over 
the abdomen. 


In Illinois B. apicalis was collected 
on jewelweed and stinging wood nettle 
in moist, shaded, bottomlands along 
streams. Very seldom were specimens 
collected on the drier hillsides. 


This species extends from North Car- 
olina to New York and west to Illinois, 
Missouri, and Oklahoma (Fig. 43). 


Illinois Records.—(Fig. 43). Col- 
lected from early May to mid-August 
in south and central Illinois. The north- 
western limit of distribution of B. api- 
calis is in Illinois. No specimens are 


Inurnois Naturat History SURVEY 


BULLETIN 


Fig. 43.—Distribution of Bittacus apicalis 
in Illinois and North America. 


recorded from northern Illinois, Iowa, 
or Wisconsin. 


Bittacus punctiger Westwood 


Bittacus punctiger Westwood \( 1846: 
195). ¢, 9. Type-locality: Georgia. 
Lectotype ¢ designated by Byers 
(1962b). 

Head and thorax pale yellow. 

Wings (Fig. 23) dark yellow; heav- 
ily patterned with dark brown mark- 
ings, particularly around crossveins; 
pterostigma dark brown. Subcostal 
crossvein basal to first fork of radial 
sector. Two pterostigmal crossveins. 
Apical crossvein present. 

Legs dark yellow with dark brown 
band at apices of femora and tibiae. 
Femora with dark brown spot at base 
of each seta, particularly on hind legs. 
Hind femora not noticeably swollen. 

Abdomen yellowish brown to dark 
brown with narrow dark brown strip 
along posterior margin of each tergum. 


Vol. 31, Art. q 


Aug., 1975 


Ninth tergum pale yellowish brown, 
basistyles dark brown. In males, lobes 
of ninth tergum in lateral view (Fig. 
32) rectangular, not extending beyond 
apices of basistyles, lobe apices emar- 
ginate; in dorsal view (Fig. 31) lobes 
diverge apically, sides straight with two 
medial prominences, each with several 
fine black spines. Basistyles broad. Dis- 
_tistyles short, projecting medially. Ae- 
deagus thickened at base, tapering api- 
cally to fine looped thread. Cerci elon- 
gate, extending slightly beyond apices 
of basistyles, bases swollen. 


This species resembles B. strigosus 
and B. pilicornis in the heavily mar- 
gined crossveins although it is readily 
separated from these species by the 
dark brown spot surrounding the base 
of each femoral seta and the dark brown 
maculation of the wings. 


This species was collected with indi- 
viduals of B. strigosus, B. apicalis, and 
B. pilicornis in a moist shaded woodland 
and among jewelweed. 


B. punctiger extends from Florida to 
Pennsylvania and west to Illinois and 
Texas (Fig. 44). 


Fig. 44.—Distribution of Bittacus punctiger 
in North America. 


Illinois Records.—ALEXANDER COUN- 
ty: 1 mile N of Olive Branch, D. W. 
Webb, 14-VI-1972, 42; 1 mile E of 
Olive Branch, Penny and Byers, 30-V- 
1972, ¢, @. FRANKLIN County: 3 
miles S of West Frankfort, 11-VI-1970, 
J. C. Marlin, 1¢. Itxrors: Belfrage 
Collection, Stockholm Museum, 12, 


Wess Er Au.: Mrecoprera OF ILLINOIS 


273 


1¢. Union County: Pine Hills Eco- 
logical Area, 14-VI-1972, D. W. Webb, 
49. 

Bittacus pilicornis Westwood 
Bittacus pilicornis Westwood (1846: 


196). ¢, 2. Type-locality: Amer- 
ica Septentrionali. Type-specimen 
missing. 


Head and thorax dark yellow to dark 
brown. 


Wings (Fig. 24) amber, pterostigma 
slightly darker than surrounding area, 
crossveins margined. Subcostal cross- 
vein basal to first fork of radial sector. 
Two pterostigmal crossveins. Apical 
crossvein present. 


Legs pale yellow to brown. Apices 
of tibiae and basistarsus dark brown. 
Hind femora not swollen. 


Abdomen pale yellow to dark brown. 
In males ninth tergum and basistyles 
brown. In males lobes of ninth tergum 
in lateral view (Fig. 33) broad, not ex- 
tending beyond apices of basistyles, 
lobe apices pointed; in dorsal view 
(Fig. 34) lobes thick, diverging api- 
cally, with 30 or more black spines 
across apical halves of lobes. Basistyles 
broad. Dististyles short, acute. Aedeagus 
with distinctive bilobed base ( penunci), 
tapering apically to ,slender coiled 
thread. Cerci elongate, slender, extend- 
ing slightly beyond apices of basistyles. 


This species is similar to B. punctiger 
in having wings with an apical cross- 
vein and margined crossveins although 
it differs markedly from B. punctiger 
in characters of the ninth tergum in 
males and the lack of a dark brown spot 
surrounding the base of each femoral 
seta. 


This species is the strongest flier of 
the midwestern bittacids and has been 
collected from damp, cool, shaded bot- 
tomlands and dry, shaded hillsides. In 
moist areas it is associated with jewel- 
weed and stinging wood nettle, while 
in drier areas it has been collected fre- 
quently on gooseberry and multiflora 
rose. 


274 


This species extends from Florida to 
Canada and west to Minnesota and 
Kansas (Fig. 45). 


Fig. 45.—Distribution of Bittacus pilicornis 
in Illinois and North America. 


Illinois Reeords.—(Fig. 45). Col- 
lected in Illinois from June to mid- 
August. 


Bittacus occidentis Walker 


Bittacus occidentis Walker (1853:469). 
$, 2. Type-locality: Erie, United 
States. Type-specimen missing. 

Bittacus arizonicus Banks (1911:350). 
8. Type-locality: Palmerlee, Ari- 
zona. Synonymized by Carpenter 
(1931a). 

Head and thorax dark yellowish 
brown to dark brown. 

Wings (Fig. 25) pale yellow, ptero- 
stigma slightly darker than surrounding 
membranes. Subcostal crossvein distal 
to first fork of radial sector. Two ptero- 
stigmal crossveins. Apical crossveins 


Intinois NatTuRAL History SuRVEY BULLETIN 


absent. Several specimens possess an 
apical crossvein on at least one of the 
fore wings. In one specimen the sub- 


Vol. 31, Art. 7 } 


. 


costal crossvein occurs at the first fork — 


of the radial sector although this cross- 


vein is normally found well beyond — 


the fork. 
Legs yellowish brown to brown, 


apices of tibiae dark brown. Hind fem- — 


ora swollen. 


Abdomen yellowish brown to brown. . 


In males ninth tergum and basistyles 
yellowish brown to dark brown. Ninth 
tergum in lateral view (Fig. 35) nar- 
row, rounded apically, extending to or 
slightly beyond apices of basistyles; in 
dorsal view (Fig. 36) ninth tergum di- 
verges apically, with 30 or more black 
spines along dorsal margins of lobes. 
Basistyles broad, curved dorsally. Dis- 
tistyles elongate, narrow. Aedeagus very 
thick at base, tapered apically to slender 
thread which curves anteriorly. Cerci 
short, slender, not extending beyond 
middle of basistyles. 


The wing’s subcostal crossvein, distal 
to the first fork of the radial sector, and 
the swollen hind femur readily distin- 
guish this species from other Nearctic 
bittacids. 


No specific habitat has been recorded 
for B. occidentis. All Illinois specimens 
were collected at lights or in light traps. 

This species extends from Alabama 
north into Canada and west to Kansas 
and Arizona (Fig. 46), with an isolated 
record from western North Dakota. 


Fig. 46.—Distribution of Bittacus occidentis 
in North America. 


Aug., 1975 


Illinois Records. — Collected infre- 
quently and in small numbers from mid- 
July to the end of September. Apams 
County: Quincy, Evers and Mills, 
9-IX-1951, 24, 29; Flint, 19-IX-1912, 
1g. CHampaicn County: Champaign, 
Hart, 18-VII-1889, 1¢; Hart, 22-VII- 
1889, 1?,1?; Urbana, 18-IX-1909, 1 ¢; 
Riegel, 19-VII-1938, 1¢; Riegel, 29- 
VIII-1938, 13; Woodworth, 12-IX- 
1898, 2¢, 49, 1?; Hart and Kahl, 22- 
IX-1892, 1 9. Cotes County: Charles- 
ton, Riegel, 12-IX-1961, 19. Coox 
County: Chicago, W. J. Gerhard, 6-IX, 
23-VII, 6 4,12. McDonoucu County: 
Macomb, 25-IX-1959, 14. SANGAMON 
County: Springfield, Frison, 16-IX- 
1932,1¢,29. 


Bittacus strigosus Hagen 


Bittacus strigosus Hagen (1861:246). 
$, 2. Type-locality: Chicago, Wash- 
ington, St. Louis. 

Head, thorax, and mouthparts dark 
yellow to dark brown. 

Wings (Fig. 26) clear, pterostigma 
pale brown, crossveins margined. Sub- 
costal crossvein basal to first fork of 
radial sector. Two pterostigmal cross- 
veins. Apical crossvein absent. 


Legs pale yellow. Hind femora cylin- 
drical. 


Abdomen dark yellow to dark brown. 
In males ninth tergum and basistyles 
brown. In males lobes of ninth tergum 
in lateral view (Fig. 37) broad basally, 
narrowed apically, apices of lobes 
rounded, extending well beyond apices 
of basistyles and having elongate medial 
prominences on ventral margins with 
several long black setae and spines; in 
dorsal view (Fig. 38) lobes broad ba- 
sally, apical third constricted, converg- 
ing medially at apex. Basistyles broad. 
Dististyles broad, elongate. Aedeagus 
thickened basally, extended apically in 
form of thin, tightly coiled thread. Cerci 
narrow, elongate, extending well be- 
yond apices of basistyles, bases of cerci 
enlarged. 


This species has margined crossveins 
like those of B. pilicornis and B. puncti- 


Wess Er Au.: MECOPTERA OF ILLINOIS 


275 


ger but lacks the apical crossvein. The 
lobes of the ninth tergum in dorsal view 
readily separate the males from other 
midwestern bittacid males. 


B. strigosus is found abundantly in 
Illinois in habitats ranging from moist 
shaded bottomlands to dry pastures. 
This species can be collected on a wide 
range of plants. 

B. strigosus extends from Louisiana 
and South Carolina to Canada and west 
to Manitoba and Montana (Fig. 47). 


Fig. 47.—Distribution of Bittacus strigosus 
in Illinois and North America. 


Illinois Records.—(Fig. 47). Col- 
lected from early June to early Septem- 
ber in almost every county in Illinois. 


Bittacus stigmaterus Say 
Bittacus stigmaterus Say (1823:164). 
Type-locality: Fort Osage, Missouri. 
Type-specimen missing. 
Bittacus pallidipennis Westwood (1846: 
195). ¢. Type-locality unknown. 
Synonymized by Hagen (1861). 


276 Intinois NaturaL History 

Head and thorax yellow to dark 
brown. 

Wings (Fig. 27) amber, pterostigma 
slightly darker than surrounding area, 
crossveins not margined except in spe- 
cimens found in western areas of Mis- 
souri and Arkansas. Subcostal crossvein 
basal to first fork of radial sector. Two 
pterostigmal crossveins. Apical cross- 
vein absent. 


In almost all specimens examined the 
wings were uniformly colored, and the 
crossveins were not margined although 
specimens collected in southwestern 
Missouri and Arkansas show some mar- 
gination of the crossveins. 

Legs dark yellowish brown. Femora 
slightly swollen. 

Abdomen pale yellow to dark brown. 
In males ninth tergum and basistyles 
brown. In males lobes of ninth tergum 
in lateral view (Fig. 39) narrow, sub- 
rectangular, extending well beyond api- 
ces of basistyles; in dorsal view (Fig. 
40) lobes converge apically, with two 
distinct medial prominences on each 
lobe, each prominence with several 
black spines; a small patch of spines 
present near ventral margin of lobes. 
Basistyles broad. Dististyles short, 
rounded apically. Aedeagus thickened 
basally, tapered apically. Cerci narrow, 
elongate, extending beyond apices of 
basistyles. 

This species closely resembles B. 
texanus. The females are separated on 
the basis of wing color, which is not 
always reliable. In places where these 
two species overlap, the wing crossveins 
in B. stigmaterus are often margined. 
The males of these two species can be 
separated by the arrangement of spines 
on the medial margin of the ninth 
tergum. 

This species has been collected in 
habitats similar to those of B. strigosus 
and B. apicalis although it is sometimes 
found in fairly dry woods. 

B, stigmaterus extends from Georgia 
to New York and west to Minnesota and 
Texas (Fig. 48). 


Fig. 48.—Distribution of Bittacus stigma- 
terus in Illinois and North America. 


Illinois Records.—(Fig. 48). 


Bittacus texanus Banks 


Bittacus texanus Banks (1908:261). 3. 

Type-locality: Plano, Texas. 

Head and thorax dark reddish brown. 

Wings (Fig. 28) pale brown, ptero- 
stigma concolor with membranes, cross- 
veins not margined. Subcostal cross- 
vein basal to first fork of radial sector. 
Two pterostigmal crossveins. Apical 
crossvein absent. 


Legs dark reddish brown. Hind fem- 
ora slightly swollen. 


Abdomen and terminalia dark red- 
dish brown. In males lobes of ninth 
tergum in lateral view (Fig. 41) nar- 


row, elongate, extending well beyond 


apices of basistyles; in dorsal view 
(Fig. 42) lobes converge apically, me- 
dial margin having a prominence bear- 
ing several short, thick spines; 10-15 


Col- — 
lected from late June to mid-September. _ 


Aug., 1975 


short, thick spines present along medial 
margin basal to this prominence, three 
to four short and thick medial spines 
occur near apices of lobes. Basistyles 
broad. Dististyles short, globular. Ae- 
deagus thickened basally, tapered api- 
cally to fine thread. Cerci narrow, elon- 
gate, extending well beyond apices of 
basistyles. 


B. texanus closely resembles B. stig- 
materus although B. texanus is much 
darker in color. The females are sepa- 
rated on the basis of wing color, which, 
as already noted, is not always reliable. 
The males of these two species can be 
separated by the arrangement of spines 
on the medial margin of the ninth 
tergum. 


Little is known of the habitat of this 
species. In Texas individuals were 
collected with B. stigmaterus along 
streams under cover of willows and 
elms. 


B. texanus has been recorded from 
Texas, Florida, Kansas, and New Mex- 
ico (Fig. 49). 


Fig. 49.—Distribution of Bittacus texanus 
in North America, 


BOREIDAE Stephens 1829 


The Boreidae are winter insects, the 
adults emerging from November until 
May. Adults and scarabaeiform larvae 
live in, and feed on, moss. The small 
size of these insects (varying in length 
from 2.5 to 5.0 mm), the presence of 
rudimentary wings, and the distinct 
Ovipositor in females readily define 
this family of Mecoptera. The family 


Wess Et Au.: MECOPTERA OF ILLINOIS 


277 


Boreidae has only one genus, Boreus, 
which occurs in Europe, Asia, and 
North America. Fifteen species are re- 
corded from North America, but only 
two species occur east of the Rocky 
Mountains. 


Boreus Latreille 


Boreus Latreille (1816:152). Type- 
species: Boreus hyemalis Linnaeus. 


Euboreus Lestage (1940:12). Synony- 
mized by Cooper (1972). 

Ateleptera Dalman (1823:34). Synony- 
mized by Esben-Petersen (1921). 
Small, stout insects. Coloration varies 

from reddish in B. elegans to olive green 
in some specimens of B. brevicaudus 
to brown and black in most species. 
Length 2.5-5.0 mm. Head broad, ta- 
pered apically to long rostrum. Ocelli 
present, but difficult to see. Compound 
eyes black, oval. Antennae brown to 
black, filiform, with 18-24 flagellar 
segments. 


Thorax reddish brown to olive to 
black. Pronotum broad, collarlike, an- 
terior margin smooth, rounded. Wings 
light brown to black. In males wings 
reduced to pair of thick, chitinous, 
coreaceous rudiments, broad basally, 
tapering apically to acute point, with 
coarse lateral and medial setae. Hind 
wings thin, membranous, covered by 
fore wings. In females fore wings re- 
duced to short, oval pads covering hind 
wing pads, except for extremely re- 
duced wing pads of B. reductus. Legs 
dark yellow to black, elongate, with 
simple claws. 


Abdomen short, thick, pale brown to 
black. In males ninth tergum short, 
broad, apex truncate or emarginate, 
with numerous short black spines; in 
some species a concave medial depres- 
sion receives apices of dististyles. Ninth 
sternum (hypandrium ) broad, rounded; 
apex rounded, truncate, or emarginate. 
Basistyles thick, broad. Dististyles nar- 
row, elongate, each with mesal lobe and 
several thick spines along dorsal mar- 
gin. In females ovipositor composed of 


278 


50 


Intinois NaruraL History SuRvVEY BULLETIN 


Vol. 31, Art. 7 


Fig. 50-55.—Boreus brumalis. 50.—Dorsal view of male fore wing. 51.—Lateral view 


of male terminalia. 


52.—Ventral view of male ninth sternum 


(hypandrium) . 


view of male ninth tergum. 54.—Lateral view of female ovipositor. 8t, eighth tergum. 8s, 
eighth sternum, 9t, ninth tergum. 10t, tenth tergum. Ce, cerci. 55.—Ventral view of female 


ovipositor. 


eighth through tenth segments and 
cerci. Eighth sternum formed by two 
elongate sclerotized plates. Tenth ter- 
gum subrectangular, elongate, apex 
emarginate. Cerci short, triangular, 
apex acute. Eleventh segment hidden 
beneath cerci. 


Only one species of Boreus occurs in 
Illinois. 


KEY TO THE MIDWESTERN 
SPECIES OF BOREUS 


MALES 


1. Specimens pale to dark reddish 
brown. Fore wing curved smoothly 
to apex (Fig. 57). Ninth sternum 
(Fig. 59) rounded apically. Ninth 
tergum (Fig. 60) rounded apically 
nivoriundus 
Specimens dark brown to black. Fore 
wing constricted near middle (Fig. 
50). Ninth sternum (Fig. 52) emar- 
ginate apically. Ninth tergum (Fig. 
53) with medial fissure ...... brumalis 


FEMALES 


1. Specimens pale to dark reddish 
brown. Ovipositor 1.20 mm in length 
(measured from base of eighth ster- 
num to apices of cerci) ..nivoriundus 


Specimens dark brown to black. Ovi- 
positor 0.53 mm in length ....brumalis 


Boreus brumalis Fitch 
Boreus brumalis Fitch (1847:278). 2, 
9. Type-locality: eastern New York. 


Head and thorax dark brown to 
black. 


Fore wings (Fig. 50) in males dark 
brown to black, slender, apical half nar- 
rowed, apex acute with numerous 
coarse black setae along lateral and 
medial margins. Hind wings with sin- 
gle apical spur. Fore wings of females 
dark brown to black, rudimentary, re- 
duced to small suboval pads. 


Legs elongate, dark brown to black. 


Abdomen and terminalia (Fig. 51) 
dark brown to black. In males ninth 
tergum (Fig. 53) short, broad basally, 
apex truncate with narrow medial fis- 
sure, lateral areas of apex with numer- 
ous short black spines; shallow medial 
depression receives tips of dististyles. 
Dististyles elongate, curved dorsally, 
apices acute; numerous small spines 
along dorsal margins of the dististyles, 
with narrow elongate lobes on mesal 
margins. At rest dististyles curve dor- 
sally to fit into dorsomedial depression 
of ninth tergum. Ninth sternum (hy- 
pandrium )- oval, apical margin emar- 
ginate (Fig. 52). In ventral view (Fig. 
55) eighth sternum of female formed 
by two narrow, elongate plates, 6.0 
times longer than wide, rounded api- 
cally, with numerous short apical 
spines, bases and apices separated. In 
lateral view (Fig. 54) eighth sternum 
broad basally, apical three-fourths 
thick, extending beyond apex of tenth 
tergum. Tenth tergum elongate, thick, 


53.—Dorsal’ 


Aug., 1975 


3.6 times longer than wide. Tenth 
sternum hidden. Cerci short, triangu- 
lar, apices acute. 


This species is related to B. nivoriun- 
dus, the other eastern North American 
species of Boreus. The dark brown to 
black coloring generally separates B. 
brumalis from B. nivoriundus in addi- 
tion to the constricted wing pads and 
emarginate apical margin of the ninth 
sternum (hypandrium) in males. 


In Illinois individuals of B. brumalis 
have been collected primarily on moss 
in the beach-maple-tulip forest of 
southwestern Illinois along the escarp- 
ment of the Mississippi River. 

B. brumalis extends from Tennessee 
to Massachusetts and west to Ohio and 
Michigan with isolated populations in 
Illinois, Wisconsin, and Minnesota 
(Fig. 56). 

Illinois Records.— (Fig. 56). The first 
record of Boreus in Illinois was re- 


Fig. 56.—Distribution of Boreus brumalis 
in Illinois and North America. 


Wess Er Au.: MECOpTERA OF ILLINOIS 


279 


ported from Fountain Bluff in Jackson 
County by Stannard (1957). Individ- 
uals have since been collected from 
mid-October to mid-April only in the 
Ozark uplift of Illinois. 


Boreus nivoriundus Fitch 


Boreus nivoriundus Fitch (1847:277). 
8, 2. Type-locality: eastern New 
York. 

Head and thorax light to dark red- 
dish brown. 

Fore wings in males (Fig. 57) pale 
brown, broad basally, tapering apically, 
with numerous strong black setae along 
lateral and medial margins. Hind wings 
with single apical spur. In females fore 
wings pale brown, rudimentary, re- 
duced to small suboval pads. 

Legs elongate, pale brown. 

Abdomen and terminalia (Fig. 58) 
pale brown. In male ninth tergum 
(Fig. 60) short, broad basally, apex 
broadly rounded, with numerous short, 
black spines; medial depression re- 
ceives apices of dististyles. Dististyles 
elongate, curved dorsally, apex acute, 
dorsal margin with numerous small dark 
spines; at rest dististyles curved dor- 
sally to rest in dorsomedial depression 
of ninth tergite. Ninth sternum (hy- 
pandrium ) broad, entire, oval, rounded 
apically (Fig. 59). In ventral view 
(Fig. 62) eighth sternum of female 
formed by two narrow, elongate plates, 
6.3 times longer than wide, rounded 
apically, with numerous short, apical 
spines, bases and apices of plates sepa- 
rated. In lateral view (Fig. 61) eighth 
sternum broad basally, apical three- 
fourths flattened dorsoventrally, ex- 
tending beyond apex of tenth tergum. 
Tenth tergum elongate, thick, 3.1 times 
longer than wide. Cerci short, fused, 
triangular, apex acute. Tenth and elev- 
enth sterna hidden. 


B. nivoriundus is one of two eastern 
species and differs from B. brumalis 
in its pale to reddish brown coloration, 
the longer length of the female oviposi- 
tor, and the rounded apices of the ninth 
tergum and sternum in males. 


280 


57 


6| 


Intino1is NATuRAL History SurRvEY BULLETIN 


Vol. 31, Art.7 — 


Fig. 57—62.—Boreus nivoriundus. 57.—Dorsal view of male fore wing. 58.—Lateral 
view of male terminalia. 59.—Veritral view of male ninth sternum (hypandrium), 60,— 
Dorsal view of ninth tergum of male. 61.—Lateral view of female ovipositor. 62.—Dorsal 


view of female ovipositor. 


Often collected with B. brumalis in 
the deciduous forests of eastern North 
America. 

B. nivoriundus extends from Massa- 
chusetts to Maine and _ southwest 
through New York and on to Ohio, 
Kentucky, and Tennessee (Fig. 63). 


Fig. 63.—Distribution of Bareus nivoriun- 
dus in North America. 


MEROPEIDAE Esben-Petersen 1921 


This family name is emended from 
Meropidae Esben-Petersen (1921) by 
Opinion 140 of the International Com- 
mission of Zoological Nomenclature. 

The family Meropeidae is the most 
primitive group of extant Mecoptera 
in North America. The broadly rounded 
wings with their dense venation asso- 
ciate the Meropeidae with the South 
American family Notiothaumidae al- 
though current knowledge of morphol- 


ogy (Mikoleit 197la) indicates that 
these two families are not as closely re- 
lated as was previously thought. The 
Meropeidae differ from the Notiothau- 
midae in the absence of ocelli, the non- 
coalescing radial and medial veins at 
the bases of the wings, and the absence 
of a notal organ. 


Two genera are recorded for the 
Meropeidae. Merope is found in east- 
ern and north-central North America, 
and Austromerope in western Australia. 


Merope Newman 


Merope Newman (1838:180). Type- 
species: Merope tuber Newman by 
monotypy. The description of the 
type-species will characterize the 
genus. 


Merope tuber Newman 


Merope tuber Newman (1838:180). 2, 
6. Type-locality: Trenton Falls, 
New York. 
Head pale yellow to brown. Ocelli 
absent. 


Thorax pale yellow to pale brown. 
Pronotum shieldlike, extending anteri- 
orly over vertex of head, with distinct 
dorsomesal suture. 

Fore wing length 11.0-13.0 mm. 
Membranes (Fig. 64) pale whitish yel- 
low; wing broad, apex rounded. Costa 
circumambient, broader along anterior 


Aug., 1975 


Fig. 64.—Merope tuber fore wing. 


margin. Veins and crossveins numer- 
ous and variable. Pterostigma not dis- 
tinct. Thyridium absent. Small brown 
basal lobe near apex of A,. Hind wings 
slightly smaller than fore wings. The 
fore wings contain numerous veins and 
crossveins which show considerable 
variation in their number, branching, 
and origins. 

Legs pale yellow. 
paired, simple. 

Abdomen pale yellow to brown, seg- 
ments subrectangular, flattened dorso- 
ventrally. Male terminalia (Fig. 65) 
pale yellow, elongate, equal in length 
to or longer than abdomen. Ninth ter- 
gum short, emarginate apically, form- 
ing two pointed lobes. Anus mesoven- 
trally beneath ninth tergum. Basistyles 
elongate, broad basally. Dististyles 
elongate, shorter than basistyles, apex 
of each dististyle flattened laterally, 
emarginate, forming two black clawlike 
lobes; small apical concave disc in 
mesal margin of each dististyle (Fig. 
66). Cerci present as short clavate 


Tarsal claws 


65 
66 
Fig. 65-66.—Merope tuber. 65.—Dorsal 


view of male terminalia. 66.—Male dististyle. 


Wess Er At.: MECOPTERA OF ILLINOIS 


281 


lobes posterior to ninth tergum. Female 
terminalia lack sclerotized genital bulb. 

Specimens of M. tuber are rare in 
collections but have been collected 
from a variety of habitats. Illinois 
specimens have been collected in Mal- 
aise and picric acid traps. Indiana 
specimens have been collected by bait 
traps in a hickory woods near Lafay- 
ette. Most specimens recorded have 
been taken at lights, under stones, in 
rotting logs, and in European chafer 
traps. Merope appears to spend a great 
deal of time on the ground. 

Nothing is known of the immature 
stages of this insect. 

M. tuber extends from northern 
Georgia to Maine and west to Missouri 
and Minnesota (Fig. 67). 


Fig. 67.—Distribution of Merope tuber in 
North America. 


Illinois Records.—Collected during 
August in southern Illinois and during 
May in east-central Illinois. CHam- 
PAIGN County: Urbana,  Trelease 
Woods, K. H. Leim, 1-7-V-1972, 1 2 . 
Union County: Pine Hills, H. S. Dy- 
bas, 28-VIII-1963, 29-VIII-1963, 5-VIII- 
1963, 28,49. 


PANORPODIDAE Issiki 1933 


Byers (1965) first used the family 
name Panorpodidae but has recom- 
mended that Issiki (1933) be credited 
with the name because Issiki first sug- 
gested that the genus Panorpodes be 
raised to subfamily rank. 


282 


The short rostrum of Brachypanorpa 
with the gena bearing a distinct tooth 
separates the panorpodids from other 
families of North American Mecoptera. 
Two genera are distinguished, with 
Panorpodes restricted to eastern Asia 
and Brachypanorpa found in southeast- 
ern and northwestern North America. 
The family Panorpodidae is very closely 
associated taxonomically with Panorpi- 
dae, and Byers (1965) erected the fam- 
ily Panorpodidae on the basis of their 
being phytophagous and because of 
the differences between the larvae of 
the two groups. 

Only Brachypanorpa occurs in North 
America, but this genus does not occur 
in the Midwest. 


Brachypanorpa Carpenter 


Brachypanorpa Carpenter (1931a:209). 
Type-species: Panorpodes carolinen- 
sis Banks, 

Three ocelli present. Antennae fili- 
form, 30-40 flagellar segments, genae 
with distinct acute lobes. Thorax yel- 
lowish brown. Wings yellowish brown 
to amber, crossveins not margined. 
Pterostigma concolor with rest of wing. 
Thyridium absent. Wings reduced in 
some females. Legs elongate, dark yel- 
lowish brown, with pair of simple 
claws. Body light yellowish brown. 
Abdomen and terminalia dark yellow- 
ish brown, oval. Ninth tergum of males 
oval, emarginate apically, forming two 
thick lateral lobes. Hypovalves thick, 
fused near middles of basistyles, sepa- 
rate apically. Basistyles oval, elongate, 
longer than dististyles. 

Three species of Brachypanorpa are 
recorded from North America: B. 
carolinensis in the southern Appala- 
chians and B. oregonensis and B. mon- 


tana in the northwestern states (Fig. 
13). 


PANORPIDAE Stephens 1835 


The paired, serrate claws; the elon- 
gate rostrum; the presence of a thyrid- 
ium; and the narrow, elongate wings 
with the cubital vein not fused to the 


Ittivo1is NATuRAL History SuRvEY BULLETIN 


Vol. 31, Art. 7 


medial vein separate the panorpids 
from other families of Mecoptera. 
Three genera are recognized. Lepto- 
panorpa and Neopanorpa are found in 
Asia, and Panorpa occurs in North 
America and Eurasia. 


In North America Panorpa contains 
the greatest number of species of any 
genus of Mecoptera. Twenty-three spe- 
cies occur in the Midwest, eight in 
Illinois. 


Panorpa Linnaeus 
Panorpa Linnaeus (1758:551). Type- 
species: 
naeus. 
Aulops Enderlein (1910:390). Synony- 
mized by Esben-Petersen (1915). 


Estenalla Navas (1912:356). Synony- 
mized by Esben-Petersen (1915). 


Head pale yellow to dark reddish 
brown. Ocelli amber on raised subtri- 
angular pad. Antennae filiform with 
more than 30 flagellar segments. Ros- 
trum elongate, tapered. Mandibles 
large, heavily sclerotized, with two or 
three lateral teeth. Labial and maxil- 
lary palps have two and four segments, 
respectively. 


Thorax pale yellow to dark reddish 
brown. Wings colorless to amber, cross- 
veins often margined. Membranes pat- 
terned with dark brown spots or bands. 
Thyridium at base of first fork of me- 
dial vein. Legs pale yellow to dark 
reddish brown, with serrate claws. 


Abdomen and terminalia yellow to 
dark reddish brown. The sixth abdom- 
inal tergum of males may possess an 
anal horn. Apex of tergum in males 
tapered, truncate, or emarginate. Hy- 
povalves generally fused near bases of 
basistyles except in lugubris group. 
Basistyles broad, oval, usually longer 
than dististyles. Dististyles simple or 
with large mesal lobes. Ventral para- 
meres variable. In females the genital 
plate, usually heavily sclerotized, con- 
sists of a distal plate, often a basal 
plate, and a medial spermathecal 
apodeme. 


Panorpa communis Lin- 


$ 


4 


x 


é 


Wess Er At.: MECOPTERA OF ILLINOIS 


Aug., 1975 


..(lugubris group) nuptialis 


Hypovalves long, fused near bases of 


OF PANORPA 
Mates (Modified from Carpenter 1931a) 


KEY TO MIDWESTERN SPECIES 
Hypovalves fused near middles of 


Nn 


hasistyles: ((ties98)i |e eee 
2. Anal horn absent ........:..%. 


Anal horn (Fig. 20) present ........ 


Basistyles’ (Wig: 91)) 2. cc cei enes 


il 


y-% 
3 | 
“ . 
- @ 
WL oe 
ants 
[8 
aa a4 
ro) 
R39 
ov 
a E,: 
a 
a 
oF] 
ps 
oo 
eae 
‘5 
Ts 
oN a} 
OLD 
.a3 
22 
Toe 
eoo 
e | | 
Emo 
~~ 
a 
fe 9) = 
ogs 
ia? 
g|e 
Sait 
~ 
ees 
Qu. 
ca 
aes 
= Vip 
2 2'0 
ce 
ao-ێ 
larg: 
eae 
—o 
co 
ole 
ogg 
2s 
32 
oo 
Ec 


284 Ittinois NATuRAL History Survey BULLETIN Vol. 31, Art. 7 


3. Aedeagus extending posteriorly be- 4, Dististyles slender, smoothly curved 
tween dististyles (Fig. 93) ........ 4 (Fig. 93) Oi. wisi cee maculosa 

Aedeagus not extending posteriorly Dististyles broad, falcate apically 
between dististyles .............65 5 (BIg 95)) cis crashes anes submaculosa 


Fig. 80-90.—Panorpa fore wings. 80.—P. helena. 81.—P. insolens, 82.—P. debilis. 
83.—P. claripennis, 84.—P. rufescens. 85.—P. dubitans. 86.—P. braueri. MS, marginal 
spots. BB, basal band. PB, pterostigmal band. AB, apical band. FBS, first basal spot. 87.— 
P. speciosa, 88.—P. bifida. 89.—P. anomala. 90.—P. consuetudinis. 


Aug., 1975 Wess Et At.: MECOPTERA OF ILLINOIS 285 


5. Dististyles with small fingerlike Dististyles simple, without lobes (Fig. 
Mes CRA G8) sco specars pocoie ce eine latipennis MDE Vitys oie occa seccun te eens teas ahha: » ae 6 


Fig. 91-99.—Panorpa male terminalia. 91.—P. nuptialis. Ventral view of terminalia. 
92.—Dorsal view of ninth tergum. 93.—P. maculosa. Ventral view of terminalia. AE, aedea- 
gus. 94.—Ventral paramere. 95.—P. submaculosa. Ventral view of terminalia. 96.—Ventral 
paramere. 97.—Dorsal view of ninth tergum. 98.—P. latipennis. Ventral view of terminalia. 
99.—Ventral paramere. 


286 Inurivno1is NaturAL History SuRvEY BULLETIN Vol. 31, Art. 7 
6. Ninth tergum truncate apically (Figs 1108) 6 04 . oS Soe 7 
CRE, TOO) ter ayess otc ate seis a eee Reem acuta 7, Ventral parameres slender, straight 
Ninth tergum emarginate apically (Fig, 105) ye). cc cee banksi 


Fig. 100-109.—Panorpa male terminalia. 100.—P. acuta. Dorsal view of ninth tergum. 
101.—Ventral view of terminalia. 102.—Ventral paramere. 103.—P. banksi. Dorsal view 
of ninth tergum. 104.—Ventral view of terminalia. 105.—Ventral paramere. 106.—P. sig- 
moides. Ventral view of terminalia. 107.—Ventral paramere. 108.—P. nebulosa. Ventral 
view of terminalia. 109.—Ventral paramere. 


Aug., 1975 


Ventral parameres thick, curved medi- 


Pan sO ET NOT, bse coie biovettualene octane ee 8 

8. Ventral parameres sigmoidally 

curved, with barbs covering apices 
(TE UO} ceetio et acole hae rien sigmoides 

Ventral parameres not sigmoidally 

curved, with apices constricted and 

bare (Fig. 109). A small patch of 

setae on tubercle near bases of dis- 
BREE VOSS: F terstataiars~\sbet 5,6 0% oe are nebulosa 

9. Dististyles with large lobes (Fig. 
Ap Sty er cyafira © bts seen woss,o:shel dverevers a} are 10 

Dististyles simple, without lobes (Fig. 
LEAL) ee es, eS ee ea ese 13 

10. Hypovalves thick, divergent apically 
(AR TG DE Aare oe eee cieiceIceiote 11 

Hypovalves slender, elongate (Fig. 
ASLEY ays cesta. a: = Shao deloratetsisi sive cisiei ee wiece 12 

11. Lobes of dististyles large, covering 

all but tips of dististyles (Fig. 111) 
=i 0 CE GOS COB Oe PaO aerate mirabilis 

Lobes of dististyles small (Fig. 113) 
oo ett ASG A SAB eee meron oe galerita 

12. Ventral parameres with barbs (Fig. 
1) Sa oo.6 SC RIScE IIOIE es hungerfordi 

Ventral parameres without barbs 
(0a Fo le lacteley Bicker subfurcata 

13. Ventral parameres unbranched 
CEPA Sen bio 0 Pree IEE iC Ie Ren 14 

Ventral parameres branched (Fig. 
UST So 23. BSI, be Aue ae ne enon 19 

14. Each basistyle with one to three 

dark thick setae near base of each 
dististyle (Fig. 120) .......... helena 

Basistyles without dark thick setae 
near bases of dististyles .......... 15 

15. Hypovalves narrow, reaching to 

bases of dististyles (Fig. 122).... 
=o EC OL sa otigS, “ee POC ER eR rufescens 

Hypovalves not reaching bases of 
EISUISCVICS) Fy vicysiet sie aici cs ses clarsenvels 16 

16. Ventral parameres with basal tuft 
of barbs (Fig! 125) ........:60... 17 

Ventral parameres without basal tuft 
SP DAES: (Pigs teOy! ae ce cc an ce 18 

17. Ventral parameres with apical tuft 
of barbs (Fig. 125) .......... dubitans 

Ventral parameres without apical tuft 
Gc panos: CWigt: E27)! Aces. s caer insolens 


18. Basistyles with apical tubercle 
bearing tuft of setae (Fig. 128)... 
debilis 

Basistyles without apical tubercle and 
setae (Fig. 130) .......... claripennis 


19. Hypovalves thick (Fig. 132). Ven- 
tral parameres as in Fig. 133 and 


SSO tee cat kas te eye a epctey oh se eadind~vaxelny at coye loys 20 
Hypovalves thin (Fig. 140). Ventral 

parameres as in Fig. 137, 139, and 

TS eres Sieicrss aie Miatelsqersisheseyeress ve selone 21 


Wess Et At.: MEcOPTERA OF ILLINOIS 


20 


21 


22. 


ile 


287 


. Basal band of wing broken (Fig. 
SU). polos nleleaigiek wake ete end speciosa 
Basal band continuous (Fig. 86) 
BOT LOd Rein tO git. Jn AOC EO HOO e: braueri 


. Ventral parameres (Fig. 136) ex- 
tending well beyond bases of disti- 
styles 

Ventral parameres (Fig. 140) short, 
reaching at most only slightly be- 
yond bases of dististyles 


Hypovalves very short, not reach- 
ing to bases of dististyles (Fig. 
140). Ventral parameres (Fig. 141) 
with thick lateral branch, curved 
GOrSalliys cysts siereecsnevayeoses sa anomala 
Hypovalves long, extending to or 
slightly beyond bases of dististyles 
(Fig. 138). Ventral parameres (Fig. 
139) with two narrow, thin branches 
consuetudinis 


FEMALES 


Wings with very broad bands (Fig. 
68). Apex of genital plate truncate 
(Cota a OD ee ee chia nuptialis 
Wings with narrow bands or spots 
(Fig. 73 and 80). Apex of genital 
plate emarginate 


Pterostigmal band not continuous 
from anterior to posterior margin 
Of wing (Mig! 73) e.cmetae oe erence o's 
(nebulosa group).. 3 


Pterostigmal band continuous from 
anterior to posterior margin of wing 
(Fig. 80) (rufescens group).. 8 


Spermathecal apodeme (Fig. 146) 
extending beyond base of distal 
plate. Genital plate greater than 1.0 


INM AMALOM ETM, cf oteperers s Cyeve ele )si5) s16) crepe 5 
Spermathecal apodeme (Fig. 143) not 
extending beyond base of distal 
plate. Genital plate less than 1.0 
mim. in: len eth. capetreie save oteraatoe. cies 4 
Genital plate (Fig. 143) about 0.44 
mm in length; lateral lobes of 
apical emargination of distal plate 
moderately broad .......... maculosa 


Genital plate (Fig. 144) about 0.57 mm 
in length; lateral lobes of apical 
emargination of distal plate nar- 
TOR Rite toneeota meth preheat ese a submaculosa 


Spermathecal apodeme (Fig. 146) 
reaches to or beyond apical emar- 
gination of distal plate 
Spermathecal apodeme (Fig. 145) 
does not reach apical emargination 
of distal(plate..<*.......... latipennis 


First marginal spot present (Fig. 
73). Genital plate (Fig. 146) nar- 
row, elongate, over 1.5 mm in 
Giri Wo aeeeadnieconnocoonncoeon banksi 


288 InLinois NaturAL History SuRVEY BULLETIN Vol. 31, Art. 7 
First marginal spot absent. Genital 7. Genital plate (Fig. 149) oblong, con- 

plate (Fig. 147 and 149) oblong or stricted basally ............. nebulosa 

oval, 1.5 mm or less in length .... 7 acuta 


119 


Fig. 110-119.—Panorpa male terminalia. 110.—P. mirabilis. Ventral paramere. 111.— 
Ventral view of terminalia. 112.—P. galerita. Dorsal view of ninth tergum. 113.—Ventral 
view of terminalia. 114.—Ventral paramere. 115.—P. subfurcata. Ventral view of terminalia. 
116.—Ventral paramere. 117.—Dorsal view of ninth tergum. 118.—P. hungerfordi. Ventral 
view of terminalia. 119.—Ventral paramere. 


q 


Aug., 1975 Wess Er Au.: MEcoprera OF ILLINOIS 289 


Genital plate (Fig. 147) oval, basal 8. Marginal spot(s) present ......... 9 
two-thirds of plate broad ...sigmoides Marginal spot(s) absent 


12| 123 
125 P27 
29 13] 


Fig. 120-131.—Panorpa male terminalia. 120.—P. helena. Ventral view of terminalia. 
121.—Ventral paramere. 122—P. rufescens. Ventral view of terminalia. 123.—Ventral 
Paramere. 124.—P. dubitans. Ventral view of terminalia. 125.—Ventral paramere. 126.— 
P. insolens. Ventral view of terminalia. 127.—Ventral paramere. 128.—P. debilis. Ventral 
view of terminalia. 129.—Ventral paramere. 130.—P. claripennis. Ventral view of terminalia. 
131.—Ventral paramere. 


290 Inuivois NaturaL History Survey BULLETIN Vol. 31, Art. 7 

9. Spermathecal apodeme extends be- WG4) eo ee ef 10 
yond base of distal plate one or Spermathecal apodeme extends beyond 
more times length of plate (Fig. base of distal plate by less than 


14 


138 


133.—Ventral paramere. 134.—P. braueri. Ventral view of terminalia. 135.—Ventral para- 
mere. 136.—P. bifida. Ventral view of terminalia. 137.—Ventral paramere. 138.—P. con- 
suetudinis. Ventral view of terminalia. 139.—Ventral paramere. 140.—P. anomala. Ventral 


Fig. 132-141.—Panorpa male terminalia. 132.—P. speciosa. Ventral view of terminalia. : 
view of terminalia. 141.—Ventral paramere. 


Aug., 1975 Wess Er Au.: MecopTera OF ILLINOIS 291 


rectangular basal membrane ..... 


length of plate (Fig. 160) ......... 12 
consuetudinis 


ieGenital plate (Hig. 1164)) with sub-  kcieccccueeeeas 


if 


143 


Y) 
ale) 


| 

Tt 
ia 
p Py: 


142.—Panorpa nuptialis. 143.—Panorpa macu- 
losa. 144.—Panorpa submaculosa. 145.—Panorpa latipennis. 146.—Panorpa banksi. 147.— 
Panorpa sigmoides. 148.—Panorpa acuta. 149.—Panorpa nebulosa. 150.—Panorpa mirabilis. 
151.—Panorpa galerita. 152.—Panorpa hungerfordi. 153.——Panorpa subfurcata. 154.— 
Panorpa helena. 155.—Panorpa rufescens. 156.—Panorpa dubitans. 157.—Panorpa insolens. 
158.—Panorpa debilis. 159.—Panorpa claripennis. 160.—Panorpa speciosa. 161.—Panorpa 
braueri. 162.—Panorpa bifida. 163.—Panorpa anomala. 164.—Panorpa consuetudinis. 


Fig. 142-164.—Female genital plate. 


292 


Genital plate without subrectangular 
basal membrane 


11. Genital plate (Fig. 156) about 0.85 
gy Yh TOMET yas! <jniene ane erect dubitans 

Genital plate (Fig. 153) about 1.76 mm 
Sei LON ECI ic orn. o ate siieta eee subfurcata 


12. Genital plate (Fig. 161) short, 
broad, about 0.69 mm in length. 
Crossveins margined; basal band 
CONTINUOUS A. ts. dele Bie ose es braueri 

Genital plate about 1 mm or more in 
length. Basal band broken 


13. Genital plate (Fig. 155) about 0.98 
mm in length, with lateral lobes of 
apical emargination short and thick. 
Distal plate oval. Spermathecal 
apodeme extends beyond base of 
distal plate 0.41 times length of 
LLG 7 sierate ere cota eis alte etsy oes rufescens 

Genital plate subcircular. Spermathe- 
cal apodeme extends beyond base 
of distal plate more than 0.60 times 
lengthof: plate: <% cto o.0:cts.cckeeeeie 3 14 

14. Genital plate (Fig. 160) deeply 
emarginate apically, reaching al- 
most to apex of spermathecal apo- 
(YE) 11 2 OPO OSS 3. ANG ccearatrye, SP Tee speciosa 

Genital plate (Fig. 163) with mod- 
erate emargination apically 

15. Inner margins of apical emargina- 
tion of genital plate (Fig. 162) par- 
allel. Genital plate about 0.99 mm 
pO tag! Fc) cf4 eins 5 ee IPR RRR Cc yciea cy oe bifida 

Inner margins of apical emargination 
of distal plate (Fig. 163) converg- 
ing. Genital plate about 1.15 mm 
in Lene thy: 53k te cfelsc.cscheiete eta anomala 


16. Crossveins margined ............. 17 
Crossveins not margined 
17. Genital plate (Fig. 159) very broad 
basally, over 1 mm in length ..... 
claripennis 
Genital plate (Fig. 158) constricted 
basally, 1 mm or less in length. .debilis 


18. Wing membranes colorless ...... 19 
Wing membranes pale yellow to 
ATIMIIOL Oy cabaret eho) ctitkeliclise horse os ctecweeptine 3 21 


19. First basal spot fused with anterior 
part of basal band (Fig. 76)..mirabilis 

First basal spot not fused with basal 
band 

20. Genital plate (Fig. 151) with shal- 

low emargination apically, about 
0:97 mm in length! <: sc 6 sacs galerita 

Genital plate (Fig. 153) deeply emar- 

ginate apically, about 1.76 mm in 
ROMS re ccic.s verccrcrae core. subfurcata 

21. Genital plate (Fig. 152) elongate, 
about 1.50 mm in length ..hungerfordi 


Intrivois NaTurAL History SURVEY BULLETIN 


Vol. 31, Art. 7 


Genital plate less than 1.30 mm in 
length 


22. Genital plate (Fig. 154) about 1.07 
mm in length |~../.). > seit helena 


Genital plate (Fig. 157) about 0.98 mm 
in length) (7Gs..0\ic, dc eho insolens 


Lugubris Group 


The lugubris group consists of three 
Nearctic species, P. lugubris, P. rufa, 
and P. nuptialis, which are dark red- 
dish brown to black with dark, broad 
wing bands. The sixth abdominal ter- 
gum of males lacks an anal horn. The 
seventh and eighth abdominal segments | 
are elongate and slender. The ninth 
tergum of males is tapered apically, 
and the hypovalves, or ninth sternum, 
are fused near the mid length of the 
basistyles. 


Panorpa nuptialis Gerstaecker 


Panorpa nuptialis Gerstaecker (1863: 
187). 9, 8. Type-locality: Texas. 


Head and thorax reddish brown. 


Fore wing length 14.0-17.9 mm. 
Membranes (Fig. 68) amber, cross- 
veins not margined. Apical band dark 
brown, broad, entire. Pterostigmal band 
dark brown, broad, entire, not forked. 
Basal band dark brown, broad, entire. 
Marginal and first basal spots fused. 
Second basal spot large, extending 
along posterior margin of wing from 
base to posterior fourth of basal band. 

Legs dark reddish brown. 


Abdomen reddish brown. Male ter- 
minalia reddish brown. Ninth tergum 
(Fig. 92) broad and rounded basally, 
tapering apically to narrow truncate 
apex. Hypovalves (Fig. 91) broad ba- 
sally, fused well beyond bases of basi- 
styles, separated apically to form two 
narrow, lateral lobes, ending well be- 
fore -bases of dfstistyles. Basistyles 
large, oval. Dististyles equal in length 
to basistyles. Ventral parameres nar- 
row, elongate, branched, extending to 
bases of dististyles. Female genital 
plate large (Fig. 142), elongate, 1.35 
mm in length. Distal plate broadened 
laterally, apex truncate. Basal plate 
narrow, elongate. Spermathecal apo- 


Aug., 1975 


deme elongate, bifurcate basally, ex- 
tending slightly beyond apex of distal 
plate. 


The broad dark bands on the wings 
and the elongate seventh and eighth 
abdominal segments associate P. nup- 
tialis with P. lugubris and P. rufa. Both 
sexes of P. nuptialis are readily distin- 
guished by the broad, unforked ptero- 
stigmal band and the large second basal 
spot which extends along the posterior 


margin of the wing from the base to 
the basal band. 


Individuals of P. nuptialis have been 
collected in dense vegetation along a 
drainage ditch in Missouri, in short 
grass of roadside ditches, and in cotton 
and soybean fields. The general habi- 
tat of this and other species of the P. 
lugubris group differs markedly from 
that of most panorpids. 

P. nuptialis is a south-central species 
recorded from Louisiana to Missouri 
and southwest into Mexico (Fig. 165). 


Fig. 165.—Distribution of Panorpa nuptialis 
in North America. 


It is the only species of this group to 
extend up the Mississippi valley, and 
it has been collected within a mile of 
Illinois. 


Nebulosa Group 


The nebulosa group consists of eight 
species of Panorpa, seven of which 
occur in the Midwest. The wing mem- 
branes are usually clear, and the wing 
bands are generally reduced to numer- 
ous small spots. In males the sixth ab- 
dominal tergum lacks an anal horn. 


Wess Er Au.: MECOPTERA OF ILLINOIS 


293 


The seventh and eighth abdominal seg- 
ments are short. The ninth tergum is 
truncate or emarginate apically, and 
the hypovalves are fused at the bases 
of the basistyles. 


Panorpa maculosa Hagen 


Panorpa maculosa Hagen (1861:245). 
8, 2. Type-locality: Pennsylvania. 

Panorpa utahensis Gurmey (1937:223). 
@ 9. Synonymized by Gurney 
(1938), and now placed in P. sub- 
maculosa by Webb, Penny, and 
Marlin. 


Head and thorax dark to reddish 
yellow. 


Fore wing length 11.6-11.8 mm. 
Membranes (Fig. 69) clear to pale yel- 
low, crossveins margined. Apical band 
pale brown, broken into numerous small 
brown spots. Pterostigmal band broad 
anteriorly, broken into small brown 
spots posteriorly. Basal band broken 
into two small spots. Marginal and sec- 
ond basal spots absent. First basal 
spot small. 


Legs pale to dark yellow. 


Abdomen dark yellow to reddish 
brown. Male terminalia dark yellow. 
Ninth tergum, as in Fig. 97, oval, broad 
basally, apex deeply emarginate. Hypo- 
valves (Fig. 93) slender, elongate, end- 
ing before bases of dististyles. Basi- 
styles broad, with medial patch of fine 
setae at bases of dististyles. Dististyles 
slender, each with large basal lobe, 
shorter than basistyles. Ventral para- 
meres (Fig. 94) short, slender, un- 
branched, barbed along one side, ex- 
tending beyond bases of dististyles. 
Aedeagus extending between  disti- 
styles. Female genital plate (Fig. 143) 
small, 0.44 mm in length. Distal plate 
short, rounded basally, apex deeply 
emarginate, forming two broad lateral 
lobes. Basal plate absent. Spermathe- 
cal apodeme short, not extending be- 
yond base of distal plate and not reach- 
ing apical emargination of distal plate. 

This species is closely associated with 
P. submaculosa. In both species the 


294 


aedeagus extends between the disti- 
syles. In both P. maculosa and P. sub- 
maculosa the female genital plate is 
small, and the spermathecal apodeme 
does not extend beyond the base of the 
distal plate. 


Individuals of P. maculosa have been 
collected on tall herbaceous vegetation 
in swampy woods of ash, oak, and yel- 
low birch (Byers 1954). 


P. maculosa extends from Georgia to 
Vermont and west to Michigan (Fig. 
166). 


Fig. 166.—Distribution of Panorpa maculosa 
in North America. 


Panorpa submaculosa Carpenter 


Panorpa submaculosa Carpenter 
(1931a:255). 38, 9. Type-locality: 
Ann Arbor, Michigan. 


Panorpa utahensis Gurney (1937:223). 
68. Synonymized by Gurney 
(1938 ). 


Panorpa utahensis Gurney (1937:223). 
? ¢. New synonymy. Gurney (1938) 
synonymized the females of P. utah- 
ensis with those of P. maculosa. 
Head and thorax pale yellow to red- 

dish brown. 


Fore wing length 10.4-12.1 mm. 
Membranes (Fig. 70) clear, crossveins 
margined. Apical band dark brown, 
broad, with numerous large clear spots. 
Pterostigmal band dark brown, broad 
anteriorly, narrow and broken poster- 
iorly. Basal band broken, forming two 
small dark brown spots. Marginal and 


Ittrwvois NaturAL History Survey BULLETIN 


Vol. 31, Art.7 — 


second basal spots lacking. First basal 
spot small. 


Legs pale to dark yellow. 


Abdomen dark yellow to reddish 
brown. Male terminalia dark yellow. 
Ninth tergum (Fig. 97) elongate, 
broad basally, tapered toward emar- 
ginate apex. Hypovalves (Fig. 95) 
moderately broad, extending three- 
fourths length of basistyles. Basistyles 
broad. Dististyles short, each with 
large basi-mesal lobe. Ventral para- 
meres (Fig. 96) narrow, barbed, un- 
branched, elongate, extending well be- 
yond bases of dististyles. Aedeagus ex- 
tends posteriorly between dististyles. 
Female genital plate (Fig. 144) short, 
0.57 mm in length. Distal plate short, 
rounded, deeply emarginate apically, 
forming two moderately narrow lateral 
lobes. Basal plate absent. Spermathe- 
cal apodeme very short, not extending 
beyond base of distal plate and not 
reaching apical emargination of distal 
plate. 


The posterior extension of the aedea- 
gus between the dististyles associates 
P. submaculosa with P. maculosa. The 
two species differ in the shape of the 
dististyles. 


Individuals of P. submaculosa are 
found in drier, less dense habitats than 
are most species of Panorpa. 


P. submaculosa is an eastern species, 
extending from Georgia to Maine and 
west to Wisconsin (Fig. 167), with an 


Fig. 
maculosa in North America. 


167.—Distribution of Panorpa sub- 


Aug., 1975 


isolated record from Utah (Gumey 
1937, described as P. utahensis). 


Panorpa latipennis Hine 


Panorpa latipennis Hine (1901:248). 
6, 2. Type-locality: Detroit, Mich- 
igan; Sea Cliff, Long Island, New 
York. 

Panorpa longipennis Banks (1911:349). 
2. Type-locality: Black Mountain, 
North Carolina. Synonymized by 
Carpenter (1931a). 

Head and thorax dark reddish brown. 


Fore wing length 13.0-14.0 mm. 
Membranes (Fig. 71) clear to faint 
brown, crossveins margined. Apical 
band pale brown, broken, with numer- 
ous clear spots. Pterostigmal band pale 
brown, broad anteriorly, but broken 
posteriorly. Basal band reduced to two 
small pale brown spots. Marginal and 
second basal spots absent. First basal 
spot small, pale brown. The continuity 
of the apical and pterostigmal bands 
varies considerably. In females the 
banding is broader and darker than it 
is in males. 

Legs pale brown, apical segments 
darker. 


Abdomen dark reddish brown. Male 
terminalia reddish brown. Ninth ter- 
gum broad, elongate, apex emarginate, 
forming two broad lateral lobes. Hypo- 
valves (Fig. 98) moderately broad, ex- 
tending three-fourths length of basi- 
styles, apical one-fourth narrowed. 
Basistyles broad, longer than dististyles. 
Dististyles falcate, each with slender 
fingerlike lobe. Ventral parameres 
(Fig. 99) narrow, elongate, barbed, 
unbranched, extending slightly beyond 
bases of dististyles. Female genital 
plate (Fig. 145) large, elongate, 1.37 
mm in length. Distal plate broad, apex 
deeply emarginate, forming two broad 
lateral lobes. Basal plate oval, tapered 
basally. A broad sclerotized membrane 
extends laterally over basal plate. Sper- 
mathecal apodeme elongate, widely 
bifurcate basally, not reaching apical 
emargination of distal plate. 


Wess Et Au.: MEcOpTERA OF ILLINOIS 


295 


In P. latipennis the general appear- 
ance of the male terminalia resembles 
those of P. banksi and P. claripennis 
with the fingerlike lobe of each disti- 
style readily separating P. latipennis 
from these two species. 

In Wisconsin individuals have been 


collected among ferns in a red oak- 
white pine forest. 


P. latipennis is an eastern species 
which extends from North Carolina to 
Vermont and west to Michigan and 
Wisconsin (Fig. 168). 


Fig. 168.—Distribution of Panorpa latipen- 
nis in North America. 


Panorpa acuta Carpenter 


Panorpa acuta Carpenter (1931a:253). 
$. Type-locality: Smoky Mountains, 
Tennessee, near Newfound Gap. 


Head pale to dark yellow, thorax 
pale yellow to dark reddish brown. 


Fore wing length 10.2-13.4 mm. 
Membranes (Fig. 72) clear, crossveins 
margined. Apical band broken into 
numerous pale brown spots. Pterostig- 
mal band indistinct, broken into numer- 
ous pale brown spots. Basal band re- 
duced to two small brown spots. Mar- 
ginal and second basal spots absent. 
First basal spot very small, pale brown. 

Legs pale yellow. 

Abdomen dark yellow to dark red- 
dish brown. Male terminalia dark yel- 
low. Ninth tergum (Fig. 100) narrow, 
elongate, apex truncate. Hypovalves 
(Fig. 101) moderately broad, apical 


296 Intinois NatTuraAL Hisrory 
third narrowed, ending before bases of 
dististyles. Basistyles broad, each with 
medial patch of thick setae at bases of 
dististyles. Dististyles shorter than ba- 
sistyles. Ventral parameres (Fig. 102) 
narrow, barbed, unbranched, extending 
slightly beyond bases of dististyles. 
Female genital plate (Fig. 148) elon- 
gate, 1.47 mm in length. Distal plate 
deeply emarginate apically, forming 
two moderately broad lateral lobes. 
Basal lobe narrowed basally. Sper- 
mathecal apodeme elongate, extending 
beyond apical emargination of distal 
plate. 


The truncate ninth tergum of males 
separates P. acuta from other species 
in the nebulosa group although the fe- 
male genital plate is identical with that 
of P. nebulosa. 


P. acuta has been collected in the 
same habitat as that of P. nebulosa 
along cool shaded ravines and at high 
elevations. 


P. acuta is an eastern species extend- 
ing from Georgia to Vermont along the 
Appalachian Mountains with an iso- 
lated record from Michigan (Fig. 169). 


Fig. 169.—Distribution of Panorpa acuta in 
North America. 


Panorpa banksi Hine 


Panorpa banksii Hine (1901: 247). ¢. 
Type-locality: Sea Cliff, New York. 

Panorpa affinis Banks (1895:315). 3. 
Type-locality: Sea Cliff, New York. 
Original name preoccupied. Re- 
named by Hine (1901). 


Survey BULLETIN Vol. 31, Art.7 
Panorpa chelata Carpenter (193la: 
251). 8, 2. Type-locality: Wol- 
laston, Massachusetts. Synonymized 
by Byers (1974). 
Head and thorax pale to dark yellow. 


Fore wing length 10.4-12.5 mm. 
Membranes (Fig. 73) faintly yellow, 
several crossveins margined. Apical 
band dark brown, separated into a nar- 
row band across apex and several dark 
brown subapical spots. Pterostigmal 
band dark brown, broad anteriorly, 
broken into several dark brown spots 
posteriorly. Basal band broken into 
two large spots. First marginal and 
first basal spots dark brown. Second 
marginal and second basal spots absent. 
The wing bands show considerable 
variation in the size and arrangement of 
spots. The first marginal spot is usu- 
ally present, but in several specimens 
no marginal spots were evident. 


Legs dark yellow. 


Abdomen dark yellow to reddish 
yellow. Male terminalia reddish yel- 
low. Ninth tergum (Fig. 103) elon- 
gate, emarginate apically, forming two 
narrow lateral lobes. Hypovalves (Fig. 
104) elongate, narrow, tapered poste- 
riorly, ending near bases of dististyles. 
Dististyles about one-half length of 
basistyles. Ventral parameres (Fig. 
105) elongate, unbranched, barbed, ex- 
tending well beyond bases of disti- 
styles. Female genital plate (Fig. 146) 
elongate, 1.61 mm in length. Distal 
plate short, deeply emarginate apically, 
forming two narrow lateral lobes. Basal 
plate oval, elongate. Spermathecal 
apodeme elongate, widely divergent 
basally, extending beyond apical emar- 
gination of distal plate. 


The male terminalia of P. banksi 
closely resemble those of P. neglecta 
although the hypovalves are broader 
than those of P. neglecta and the ven- 
tral parameres are barbed. 


In Illinois individuals of P. banksi 
were collected in relatively dry areas 
away from the humid bottomlands. 
Near Chicago individuals were col- 


Aug., 1975 


lected on a dry gravel hillside among 
wild roses and in narrow steep ravines 
in cultivated areas. 

P. banksi is a northeastern species 
extending from Georgia to Maine and 
west to Illinois, Iowa, and Wisconsin. 

Illinois Records.—(Fig. 170). Col- 
lected from mid-May until early Au- 


TNGSTON 


EFFINGHAM 
CLAY 


Fig. 170.—Distribution of Panorpa banksi 
in Illinois and North America. 


gust. Restricted to the hilly areas of 
northern, western, and southern Illinois. 


Panorpa sigmoides Carpenter 


Panorpa sigmoides Carpenter (1931a: 
250). 6, 9. Type-locality: Turkey 
Run [State Park], Indiana. 

Head and thorax pale yellow to dark 
yellowish brown. 

Fore wing length 10.7-11.7 mm. 
Membranes (Fig. 74) clear to pale yel- 
low, crossveins margined. Apical band 
dark brown, broken into a narrow api- 
cal and subapical band. Pterostigmal 


Wess Ev Au.: MEcopTeRA OF ILLINOIS 


297 


band dark brown, broken but forked. 
Basal band reduced to two dark brown 
spots. Marginal and second basal spots 
absent. First basal spot dark brown. 
Considerable variation occurs in the 
arrangement of the apical and ptero- 
stigmal bands. 


Legs pale to dark yellow. 


Abdomen dark yellow. Male termina- 
lia pale to dark yellow. Ninth tergum, 
as in Fig. 103, broad basally, apex 
emarginate, forming two narrow, lat- 
eral lobes. Hypovalves (Fig. 106) en- 
larged medially, tapering apically, end- 
ing before bases of dististyles. Disti- 
styles shorter than basistyles. Ventral 
parameres (Fig. 107) thick, un- 
branched, with barbs on both margins; 
parameres sigmoidally curved, extend- 
ing beyond bases of dististyles, each 
apex smoothly tapered to acute point. 
Female genital plate elongate (Fig. 
147), 1.39 mm in length, oval. Distal 


Se 
= 


171.—Distribution of Panorpa sig- 


Fig. 
moides in Illinois and North America. 


298 Inuivois NaTuRAL History 
plate broad, apex emarginate, forming 
two narrow lateral lobes. Basal plate 
broad, oval. Spermathecal apodeme 
elongate, widely bifurcate basally; apex 
swollen, extending beyond apical emar- 
gination of distal plate. In the female 
genital plate, significant variation was 
noted, making the separation of female 
specimens from P. banksi and P. nebu- 
losa very subjective. 

The male terminalia of P. sigmoides 
resemble those of P. nebulosa, but the 
middle third of the hypovalves is 
broader in P. sigmoides, and the ven- 
tral valves are sigmoidally curved. 


This species was collected on sting- 
ing wood nettle and jewelweed along 
heavily wooded streams throughout 
Illinois. It appears to require a moister 
habitat than most species of Panorpa. 

P. sigmoides is a midwestern species 
extending from Tennessee to Ohio and 
west to Minnesota. 

Illinois Records.—(Fig. 171). Col- 
lected abundantly from the end of 
April to early August. 


Panorpa nebulosa Westwood 


Panorpa nebulosa Westwood (1846: 
188). 2. Type-locality: America 
boreali. Byers (1962b) reported that 
the female holotype bears the local- 
ity Trenton Falls, New York. 

Head and thorax dark yellow to dark 
reddish brown. 


Fore wing length 10.2-12.9 mm. 
Membranes (Fig. 75) clear, crossveins 
faintly margined. Apical band broken 
into numerous pale brown spots. Ptero- 
stigmal band pale brown, broad ante- 
riorly, forked but broken posteriorly. 
Basal band reduced to two small brown 
spots. Marginal and second basal spots 
absent. First basal spot pale brown. 


Legs pale to dark yellow. 


Abdomen pale yellow to dark brown. 
Male terminalia pale yellowish brown. 
Ninth tergum, as in Fig. 103, broad ba- 
sally, constricted at apical third, apex 
emarginate, forming two broad lateral 
lobes. Hypovalves (Fig. 108) narrow, 


SurveEY BULLETIN Vol. 31, Art. 7 
elongate, tapered apically, extending 
three-fourths length of basistyles. Basi- 
styles broad, each with patch of elongate 
setae at base of dististyle. Dististyles 
shorter than basistyles. Ventral para- 
meres (Fig. 109) elongate, sinuate, 
crossing medially, barbed, unbranched, 
apex narrowed and bare. Female geni- 
tal plate (Fig. 149) elongate, 1.40 mm 
in length. Distal plate large, apex emar- 
ginate, forming two broad lateral lobes. 
Basal plate narrowed basally. Sper- 
mathical apodeme elongate, bifurcate 
basally, apex swollen, reaching slightly 
beyond apical emargination of distal 
plate. Considerable variation is evi- 
dent in the female genital plate, mak- 
ing the separation of P. nebulosa from 
P. sigmoides difficult. 


The male terminalia of P. nebulosa 
resemble those of P. sigmoides although 
differing in the shape of the ventral 
parameres. The female of P. nebulosa 
cannot be separated from the P. acuta 
female on the basis of the genital plate. 


This species occurs in a wide range 
of habitats, both wet and dry, always 
in wooded situations. 


P. nebulosa is a wide-ranging east- 
em species extending from Georgia to 
Quebec and west to Wisconsin and 
Missouri (Fig. 172). 


Fig. 172.—Distribution of Panorpa nebu- 
losa in North America. 


Illinois Records. — Collected from 
early May to late July. Du Pace 
County: Wayne, 19-VII-1947, R. Mit- 


H 


3 


Aug., 1975 


chell, 1 9. Harpin County: Elizabeth- 
town, 22-VI-1932, H. H. Ross,13,19. 
LakE County: Lake Forest, 6-V-1906, 
J. G. Needham, 28. Wooprorp 
County: 4 miles W of Cazenovia, 10- 
VI-1969, Webb and Marlin, 29. Ici 
nots: Belfrage Collection, Stockholm 
Museum, 13,19. 


Rufescens Group 


The rufescens group is the largest 
species-group of Panorpa, having 30 
species, of which 15 occur in the Mid- 
west. The wing membranes vary from 
clear to dark yellow and usually have 
broad apical and pterostigmal bands. 
The pterostigmal band is generally con- 
tinuous from the anterior to the poste- 
rior margin of the wing. The sixth 
abdominal tergum of males possesses 
an anal horn. The ninth tergum of 
males is emarginate apically, often 
forming two narrow lateral lobes. The 
hypovalves (ninth sternum) are fused 
near the bases of the basistyles. 


Panorpa mirabilis Carpenter 


Panorpa mirabilis Carpenter (1931a: 
229). ¢, @. Type-locality: An- 
dover, New Jersey. 

Head and thorax dark reddish brown. 


Fore wing length 13.3-13.8 mm. 
Membranes (Fig. 76) clear to pale 
grey, crossveins not margined. Apical 
band pale brown, entire, with one or 
two small clear spots. Pterostigmal 
band pale brown, continuous, apical 
fork broken. Basal band pale brown, 
usually entire, fused with first basal 
spot along anterior margin. Both mar- 
ginal and second basal spots absent. 
First basal spot pale brown. 

Legs pale to dark yellow. 

Abdomen pale to dark yellow. Male 
terminalia pale to dark yellow. Ninth 
tergum, as in Fig. 112, large, broad ba- 
sally, tapered to shallow apical emar- 
gination. Hypovalves (Fig. 111) broad, 
divergent apically, ending before bases 
of dististyles. Basistyles narrow, each 
with small patch of setae near base of 
dististyle. Dististyles longer than basi- 


Wess Et Au.: MECOPTERA OF ILLINOIS 


299 


styles each with large lobe nearly 
covering dististyle and with pair of 
large basi-medial lobes. Ventral para- 
meres (Fig. 110) narrow, elongate, un- 
branched, barbed, extending almost to 
apices of dististyles. Female genital 
plate (Fig. 150) large, elongate, 1.58 
mm in length. Distal plate deeply 
emarginate apically. Basal plate ab- 
sent. Spermathecal apodeme long, ex- 
tending well beyond base of distal plate 
but not reaching apical emargination. 


The shapes of the hypovalves and 
dististyles readily associate P. mirabilis 
with P. galerita, but the narrow elon- 
gate ventral parameres of P. mirabilis 
separate the two species. In females 
the long spermathecal apodeme and the 
overall length of the genital plate of 
P. mirabilis readily separate this spe- 
cies from P. galerita. 

Nothing has been recorded on the 
habitat of P. mirabilis. 

P. mirabilis is a northeastern species, 
recorded from New Jersey, New York, 
Pennsylvania, and Michigan (Fig. 173). 


Fig. 173.—Distribution of Panorpa mirabilis 
in North America. 


Panorpa galerita Byers 

Panorpa galerita Byers (1962b:472). 

6, @. Type-locality: Lake Jean, 

Ricketts Glen State Park, Sullivan 

County, Pennsylvania. 

Head and thorax pale to reddish 
yellow. 

Fore wing length 12.5-13.7 mm. 
Membranes (Fig. 77) clear, crossveins 
not margined. Apical band pale brown, 


300 


entire or with few clear spots. Ptero- 
stigmal band pale brown, entire, pos- 
terior fork usually broken. Basal band 
pale brown, entire, occasionally fused 
anteriorly with first basal spot. Mar- 
ginal and second basal spots absent. 
First basal spot pale brown. 


Legs dark to reddish yellow. 


Abdomen reddish brown. «Male ter- 
minalia reddish brown. Ninth tergum, 
as in Fig. 112, oval, narrowed apically, 
apical margin with shallow emargina- 
tion. Hypovalves (Fig. 113) broad, 
divergent apically, ending before bases 
of dististyles. Basistyles broad, each 
with small medial patch of setae near 
base of dististyle. Dististyles large, 
shorter than basistyles, with broad dor- 
sal lobe covering two-thirds of each dis- 
tistyle and two sinuate basi-medial 
lobes. Ventral parameres (Fig. 114) 
thick, sinuate, unbranched, barbed, ex- 
tending well beyond bases of disti- 
styles. Female genital plate (Fig. 151) 
short, 0.97 mm in length. Distal plate 
subtriangular,' tapered basally, with 
concave apical emargination. Basal 
plate absent. Spermathecal apodeme 
extending beyond base of distal plate 
but not reaching apical emargination. 


The large lobes of the dististyles, the 
divergent apices of the hypovalves, and 
the shape of the ninth tergum readily 
associate P. galerita and P. mirabilis. 
The males of P. galerita differ from 
those of P. mirabilis in the thick barbed 
ventral parameres, the dististyles being 
shorter than the basistyles, and the 


Fig. 174.—LDistribution of Panorpa galerita 
in North America. 


Inutinois NatTurAL History SurvEY BULLETIN 


Vol. 31, Art.7 


lobes of the dististyles covering only 
two-thirds of the dististyles. In females 
the genital plate of P. galerita is con- 
siderably smaller in length than that of 
P. mirabilis. 


Individuals of P. galerita have been 
collected among ferns at the edge of 
a beech, maple, and hemlock forest. 


P. galerita is a northeastern species 
extending from Quebec and Vermont 
west to Ohio with a disjunct distribu- 
tion in Wisconsin (Fig. 174). 


Panorpa subfurcata Westwood 


Panorpa subfurcata Westwood (1846: 
191). 8, 9%. Type-locality: Nova 
Scotia. 

Panorpa modesta Carpenter (1931a: 
233). 8. Type-locality: Douglas 
Lake, Michigan. Synonymized by 
Byers (1974). 

Panorpa signifer Banks (1900:251). 6, 
2. Type-locality: Gaylord, Michi- 
gan. Synonymized by Byers (1962b). 
Head and thorax reddish to dark red- 

dish brown. 

Fore wing length 11.1-144 mm. 
Membranes (Fig. 79) clear, crossveins 
not margined. Apical band dark brown, 
broad, with several small clear spots. 
Pterostigmal band dark brown, broad 
anteriorly, forked, apical branch may 
or may not be continuous. Basal band 
broad, entire. Marginal spots variable. 
First basal spot dark brown, second 
basal spot present or absent. Byers 
(1962b) reported that the marginal 
spot was absent in all specimens of the 
type series, as is the case in most of 
the specimens we examined. However, 
material examined from Minnesota 
showed as many as four marginal spots. 


Legs reddish to dark reddish brown. 


Abdomen reddish brown. Male ter- 
minalia reddish brown. Ninth tergum 
(Fig. 117) long, broad basally, con- 
stricted three-fourths way from base; 
apex emarginate, forming two broad 
lateral lobes. Hypovalves (Fig. 115) 
slender, elongate, ending before bases 
of dististyles. Basistyles broad, each 


Aug., 1975 


with patch of elongate setae near base 
of dististyle. Dististyles large, almost 
equal in length to basistyles, with large 
medial lobe. Ventral parameres (Fig. 
116) slender, elongate, unbranched, 
bare, extending well beyond bases of 
dististyles. Female genital plate (Fig. 
153) long, 1.76 mm in length. Distal 
plate oval, apex emarginate. Basal 
plate absent. Spermathecal apodeme 
long, widely divergent basally, not 
reaching apical emargination of distal 
plate. 

The large lobes of the dististyles re- 
late P. subfurcata to P. mirabilis and 
P. galerita, but the narrow hypovalves 
and the elongate, bare ventral para- 
meres readily separate P. subfurcata 
from the latter two species. 

Collected in the dense undergrowth 
of birch-maple woodlands. 

P. subfurcata is a northeastern spe- 
cies, extending from North Carolina to 
Nova Scotia and west to Minnesota and 
western Ontario (Fig. 175). 


Kes 


Fig. 175.—Distribution of Panorpa subfur- 
cata in North America. 


Panorpa hungerfordi Byers 


Panorpa hungerfordi Byers (1973a: 
367). ¢, 2. Type-locality: 4 miles 
west of Pellston, Emmet County, 
Michigan. 

Head and thorax dark reddish brown. 


Fore wing length 11.3-12.0 mm. 
Membranes (Fig. 78) pale yellow, 
crossveins not margined. Apical band 
entire, pale brown, with two to four 
small clear spots. Pterostigmal band 
pale brown, continuous, forked, with 


Wess Et Au.: MECOPTERA OF ILLINOIS 


301 


apical branch broken. Basal band 
broken into two large pale brown spots. 
Marginal and second basal spots ab- 
sent. First basal spot small. 


Legs dark yellowish brown. 


Abdomen dark reddish brown. Male 
terminalia reddish brown. Ninth ter- 
gum, as in Fig. 117, large, broad ba- 
sally, tapered to deep apical emargina- 
tion. Hypovalves (Fig. 118) slender, 
elongate, extending to base of disti- 
styles. Basistyles broad. Dististyles 
shorter than basistyles, large, falcate, 
each with large mesal lobe. Ventral 
parameres (Fig. 119) slender, un- 
branched, barbed, extending to middle 
of dististyles. Female genital plate 
(Fig. 152) elongate, 0.87 mm in length. 
Distal plate broad apically, narrowed 
basally, apex having moderately shal- 
low emargination. Spermathecal apo- 
deme elongate, extending beyond base 
of distal plate but not reaching apical 
emargination. 

This species was intially identified by 
authors as P. virginica, which it resem- 
bles in the shape of the dististyles and 
the ventral parameres. On closer ex- 
amination P. hungerfordi differs (Byers 
1973a) in the absence of a small tooth 
on each dististyle present in P. virgin- 
ica; these species also differ in the 
shape of the lobes on the dististyles 
and in the lengths of the ventral para- 
meres. 


Nothing has been reported on the 
habitat of this species. 


Fig. 176.—Distribution of Panorpa hunger- 
fordi in North America. 


302 


Panorpa hungerfordi is distributed 


through Wisconsin, Michigan, and 
Ohio (Fig. 176). 
Panorpa helena Byers 


Panorpa helena Byers (1962b:474). 2, 
2. Type-locality: Swampy woods 
south of Hopewell Lake, French 
Creek State Park, Berks County, 
Pennsylvania. 


Panorpa venosa (Authors). 
mized by Byers (1962b). 


Head dark yellow, thorax reddish 
brown. 


Fore wing length 10.9-12.7 mm. 
Membranes (Fig. 80) clear to amber, 
crossveins not margined. Apical band 
dark brown, broad, entire, occasionally 
having few small clear spots. Ptero- 
stigmal band dark brown, broad, apical 
branch generally separated, forming 
small spot. Basal band broad, entire. 
Marginal and second basal spots ab- 
sent. First basal spot small. 

Legs pale yellow, fourth and fifth 
tarsal segments dark brown to black. 

Abdomen dark yellow. Male termi- 
nalia dark yellow. Ninth tergum, as in 
Fig. 117, oblong, rounded basally, ta- 
pered apically, deeply emarginate apex 


Synony- 


forming two broad lateral lobes. Hypo- 


valves (Fig. 120) moderately thick, ex- 
tending to bases of dististyles. Basi- 
styles broad, each with one to three 
dark black setae near bases of disti- 
styles. Dististyles about two-thirds 
length of basistyles. Ventral parameres 
(Fig. 121) narrow, elongate, barbed, 
unbranched, extending to bases of dis- 
tistyles. Female genital plate (Fig. 
154) oval, 1.07 mm in length. Distal 
plate oval, tapered basally, apex emar- 
ginate. Basal plate absent. Spermathe- 
cal apodeme elongate, base bifurcate, 
apex not reaching apical emargination 
of distal plate. 


The dark setae at the bases of the 
dististyles relate P. helena with P. amer- 
icana, but they differ in the shapes of 
the hypovalves and the ventral para- 
meres. If the dark setae at the bases 


Ittinois NaturaAL History Survey BULLETIN 


Vol. 31, Art. A 

§ 
of the dististyles were absent, the male — 
terminalia of P. helena would resemble — 
closely those of P. insolens. ; 


P. helena is probably the most abun- — 
dant and widely distributed species of — 
Panorpa in North America. It is col- 
lected readily in a moist shady woods — 
with a thick herbaceous undergrowth — 
of jewelweed, stinging wood nettle, and — 
poison ivy. 

P. helena extends from Georgia to 
Massachusetts and west to Manitoba, 
with an isolated record from Utah. 


Fig. 177.—Distribution of Panorpa helena 
in Illinois and North America. 


Illinois Records.—(Fig. 177). Col- 
lected abundantly from early May to 
mid-October throughout the state. 


Panorpa insolens Carpenter 
Panorpa insolens Carpenter (1935: 
106). 2. Type-locality: Cincinnati, 
Ohio. 
Head and thorax reddish brown. 


Fore wing length 10.9-12.4 mm. 
Membranes (Fig. 81) pale yellow, 


Aug., 1975 


crossveins not margined. Apical band 
dark brown, entire. Pterostigmal band 
dark brown to black, entire, broad 
along anterior margin, forked with api- 
cal branch broken. Basal band dark 
brown, broad, continuous. Marginal 
and second basal spots absent. First 
basal spot dark brown, small. 


Legs yellowish to reddish brown. 


Abdomen reddish brown. Male ter- 
minalia dark yellowish brown. Ninth 
tergum, as in Fig. 117, broad basally, 
tapering apically to deep emargination 
forming two thick lateral lobes. Hypo- 
valves (Fig. 126) moderately thick, 
ending slightly before bases of disti- 
styles. Basistyles broad. Dististyles 
shorter than basistyles. Ventral para- 
meres (Fig. 127) unbranched, barbed, 
but bare on apical half, extending to 
middles of dististyles. Female genital 
plate (Fig. 157) 0.98 mm in length. 
Distal plate narrowed basally, wider 
apically, with deep apical emargination. 
Basal plate absent. Spermathecal apo- 
deme elongate, extending beyond base 
of distal plate but not reaching apical 
emargination. 


P. insolens was described by Carpen- 
ter on the basis of a single female, 
which had the basal band of the right 
fore wing broken at the middle and the 
upper and lower portions fused with 
the first basal spot, a condition not pres- 
ent in the left fore wing. The sperma- 
thecal apodeme was confined to the 
distal plate. In the holotype, the broken 
end of the spermathecal apodeme is 
evident, and when the portion that was 
broken off is added to the remainder 
of the apodeme retained in the distal 
plate, the apodeme extends beyond the 
base of the distal plate. We concluded, 
after comparing the wing patterns and 
female genital plate of the holotype 
with specimens collected near the type- 
locality, that Carpenter based his de- 
scription on an aberrant specimen. 
Byers (1973a) has also discussed this 
variation in Carpenter's holotype of 
P. insolens. 


The male terminalia of P. insolens 


Wess Et Au.: MECOPTERA OF ILLINOIS 


303 


resemble closely those of P. helena, 
though lacking the dark setae near the 
bases of the dististyles. These species 
also have differences in the shapes of 
the ventral parameres. 


Collected on stinging wood nettle 
along shaded streams in northern Ken- 
tucky. 


P. insolens is known only from south- 
ern Ohio and northern Kentucky (Fig. 
178). 


Fig. 178.—Distribution of Panorpa insolens 
in North America. 


Panorpa debilis Westwood 


Panorpa debilis Westwood (1846:191). 
2, &. Type-locality: America Sep- 
tentrionali. Byers (1962b) desig- 
nated the lectotype ¢ and reported 
the type-locality as Trenton Falls, 
New York. 


Panorpa canadensis Banks (1895:315). 
8. Type-locality: Sherbrooke, Que- 
bec. Synonymized by Byers (1962b). 


Head and thorax dark yellow to red- 
dish brown. 


Fore wing length 10.4-11.4 mm. 
Membranes (Fig. 82) colorless, cross- 
veins faintly margined. Apical band 
dark brown, broad, almost entire except 
for few pale spots. Pterostigmal band 
dark brown, apical branch broken, 
leaving small spot. Basal band brown, 
separated into two large spots. Mar- 
ginal and second basal spots absent. 
First basal spot small. 


Legs dark yellow to reddish brown. 
Abdomen dark yellow to dark red- 


304 


dish brown. Male terminalia reddish 
brown. Ninth tergum, as in Fig. 117, 
broad basally, apex deeply emarginate, 
forming two lateral lobes. Hypovalves 
(Fig. 128) moderately broad, rounded 
apically, ending well before bases of 
basistyles. Basistyles broad, each with 
cluster of long setae near bases of dis- 
tistyles. Dististyles shorter than basi- 
styles. Ventral parameres (Fig. 129) 
elongate, curved, barbed, converging 
apically, extending beyond bases of dis- 
tistyles. Female genital plate (Fig. 
158) short, 0.79 mm in length. Distal 
plate narrowed basally, expanded api- 
cally, with deep emargination forming 
two broad lateral lobes. Basal plate 
absent. Spermathecal apodeme short, 
not reaching apical emargination of 
distal plate. 

The male terminalia of P. debilis re- 
semble those of P. rufescens, but the 
ventral parameres of P. debilis con- 
verge apically and the hypovalves are 
broader. 

Byers (1954) reported P. debilis in- 
habiting a wide variety of habitats. In 
southern Illinois it was collected on 
jewelweed in the Pine Hills area. In 
central Wisconsin individuals were col- 
lected in upland raspberry patches. 

P. debilis is an eastern species, ex- 
tending from North Carolina to Quebec 
and west to Illinois and Wisconsin, 
with a doubtful record in Colorado 
(Fig. 179). 


Fig. 179.—Distribution of Panorpa debilis 
in North America. 


Illinois Records. — Collected only 
twice in Illinois in mid-May and early 


ILttinois NaturAL History SurvEY BULLETIN 


Vol. 31, Art.7_ 


July. Octe County: Grand Detour, 
2-VII-1932, Dozier and Mohr, 134,19. 
Union County: Pine Hills, 18-V-1963, 
W. Brigham, 1 ¢. 


Panorpa claripennis Hine 


Panorpa claripennis Hine (1901:252), 
3. Type-locality: Sherbrooke, Que- 
bec. 


Head and thorax dark reddish brown. 


Fore wing length 12.0-14.0 mm. 
Membranes (Fig. 83) colorless, cross- 
veins faintly margined. Apical band 
dark brown, broad, broken posteriorly. 
Pterostigmal band dark brown, broad 
anteriorly, tapered posteriorly, with 
apical branch of fork broken. Basal 
band broken, forming two large dark 
brown spots. Marginal and second basal 
spots absent. First basal spot small. 


Legs dark yellow. 


Abdomen dark reddish brown. Male 
terminalia reddish brown. Ninth ter- 
gum, as in Fig. 117, elongate, deeply 
emarginate apically, forming two broad 
lateral lobes. Hypovalves (Fig. 130) 
moderately broad, tapered apically, 
ending before bases of dististyles. Basi- 
styles broad. Dististyles shorter than 
basistyles. Ventral parameres (Fig. 
131) elongate, barbed, extending well 
beyond bases of dististyles. Female 
genital plate (Fig. 159) broad, 1.30 
mm in length. Distal plate oval, broad, 
deeply emarginate apically, forming 
two narrow lateral lobes. Basal plate 
absent. Spermathecal apodeme broad, 
bifurcate basally, apex not reaching 
apical emargination of distal plate. 


The male terminalia of P. claripennis 
resemble those of P. latipennis, differ- 
ing in the absence of the basal lobes 
on the dististyles. 


Individuals collected at Otter Creek, 
Wisconsin, were abundant on jewel- 
weed on a shaded hillside of a steep 
ravine. 


P. claripennis is a northeastern spe- 
cies, extending from Maine and Quebec 
to Wisconsin with an isolated record 
from western Florida (Fig. 180). 


Fig. 180.—Distribution of Panorpa claripen- 
nis in North America. 


Panorpa rufescens Rambur 


Panorpa rufescens Rambur (1842:330). 
8, 2. Type-locality: Amerique 
septentrionale. 

Panorpa venosa Westwood (1846:190). 
Type-locality: Georgia. Lectotype 
2 designated by Byers (1962b). 
Synonymized by Byers (1962). 

Panorpa confusa Westwood (1846: 
190). &, @. Type-locality: Massa- 
chusetts. Lectotype 3 designated 
by Byers (1962b). Synonymized by 
Carpenter (1931a). 

Head and thorax pale to dark yellow. 


Fore wing length 11.4-12.4 mm. 
Membranes (Fig. 84) clear to pale yel- 
low, crossveins not margined. Apical 
band dark brown, entire, with few clear 
spots. Pterostigmal band dark brown, 
entire, posterior fork broken. Basal 
band broken, forming two large spots. 
Marginal and first basal spot small. 
Second basal spot absent. 


Legs pale to dark yellow. 


Abdomen dark yellow. Male termi- 
nalia pale yellow. Ninth tergum, as in 
Fig. 117, large, broad basally, apex 
emarginate, forming two broad lateral 
lobes. Hypovalves (Fig. 122) slender, 
extending to or just below bases of dis- 
tistyles. Basistyles broad. Dististyles 
falcate, with row of coarse setae along 
mesal margin. Ventral parameres (Fig. 
123) slender, barbed, unbranched. Fe- 
male genital plate (Fig. 155) broad, 
0.98 mm in length. Distal plate oblong, 
broad basally, with apical emargination 


Wess Et At.: MECOPTERA OF ILLINOIS 


305 


forming two broad lateral lobes. Basal 
plate absent. Spermathecal apodeme 
extending beyond base of distal plate 
but not reaching apical emargination. 


The male terminalia of P. rufescens 
resemble those of P. debilis. However, 
the hypovalves of P. rufescens are nar- 
rower and much longer, and the shape 
of the ventral parameres is different. 


Nothing has been reported on the 
habitat of this species. 


Panorpa rufescens is an eastern spe- 
cies extending from Florida to Canada 
and west to Michigan, Illinois, and Ala- 
bama (Fig. 181). 


Fig. 181.—Distribution of Panorpa rufescens 
in North America. 


Illinois Records. — Cook County: 
North Evanston, 20-VIII-1905, W. J. 
Gerhard, 12; Bowmanville, 3-VIII- 
1904, A. B. Wolcott, 1 2. 


Panorpa dubitans Carpenter 


Panorpa dubitans Carpenter (1931a: 
243). ¢. Type-locality: Hessville, 
Indiana. 


Head and thorax reddish brown. 


Fore wing length 9.9-11.8 mm. Mem- 
branes (Fig. 85) pale yellow to amber, 
crossveins margined. Apical band dark 
brown, broad, with several basal white 
spots. Pterostigmal band dark brown, 
broad anteriorly, forked posteriorly, 
apical fork broken. Basal band broken, 
forming two dark brown spots. Mar- 
ginal and first basal spots dark brown. 
Second basal spot absent. Some varia- 
tion was noted in the color of the fore 


306 Intivois NaturaL History 
wings and in the size and number of 
clear spots in the apical band. In 50 
percent of the specimens examined, 
the second marginal spot was absent. 


Legs reddish brown. 


Abdomen dark yellowish brown to 
reddish brown. Male terminalia dark 
yellowish brown. Ninth tergum elon- 
gate, base broad, apex emarginate, 
forming two slender lateral lobes. Hy- 
povalves (Fig. 124) elongate, moder- 
ately broad, ending well before bases 
of dististyles. Basistyles broad, with 
projection along mesal margin. Disti- 
styles shorter than basistyles. Ventral 
parameres (Fig. 125) elongate, barbed, 
unbranched, extending to bases of dis- 
tistyles. Female genital plate (Fig. 156) 
short, 0.85 mm in length. Distal plate 
oval, apex emarginate, forming two 
broad lateral lobes. Basal plate absent. 
Spermathecal apodeme elongate, ex- 
tending well beyond base of distal plate 
although not reaching apical emar- 
gination. 

Superficially the male terminalia of 
P. dubitans resemble those of P. speci- 
osa, especially in the shapes of the ven- 
tral parameres and hypovalves. The 
males of P. dubitans are distinguished 
from those of P. speciosa in having nar- 
rower hypovalves, longer basistyles, 
and fewer and broader barbs, tending 
to occur in tufts, on the ventral para- 
meres. 


In northern Illinois P. dubitans was 
collected on stinging wood nettle along 


Fig, 182.—Distribution of Panorpa dubitans 
in North America. 


SurvEY BULLETIN 


the bottomlands of Sugar Creek in the 


Vol. 31, Art. 7 ' 


a. Sak 


Macktown Forest Preserve, Winnebago — 


County. 
P. dubitans is a north-central species, 


occurring in Illinois, Indiana, and Wis- — 


consin (Fig. 182). 


Illinois Records. — Collected abun- — 


dantly from mid-May to early Septem- 
ber in northern Illinois. Coox County: 
Thornton, 22-VI-1949, Ross and Stan- 
nard, 14; Thornton, Glenwood Forest 
Preserve, 3-VI-1970, L. J. Stannard, 1 ¢. 
Lake County: Waukegan, 7-VII-1932, 
T. H. Frison, 1 ¢. WinNEBAGO County: 
Macktown Forest Preserve, J. C. Mar- 
lin, 16-VII-1969, 13, 17-VI-1970, 44, 
39, 4-IX-1971, 18; D. W. Webb, 10- 
VII-1970, 23. 


Panorpa speciosa Carpenter 


Panorpa speciosa Carpenter (193la: 
243). $. Type-locality: Heyworth, 
Illinois. 


Head and thorax pale yellow to dark 
brown. 


Fore wing length 10.7-12.0 mm. 
Membranes (Fig. 87) clear to amber, 
crossveins faintly margined. Apical 
band dark brown, entire, with one or 
two posterior clear spots. Pterostigmal 
band dark brown, broad anteriorly, 
forked, apical fork broken. Basal band 
broken, forming two large dark brown 
spots. Marginal and first basal spots 
small. Second basal spot absent. 


Considerable variation was noted in 
the pattern of the apical and pterostig- 
mal bands. In certain specimens the 
pterostigmal band is continuous and 
has both posterior branches. The num- 
ber of marginal spots varies from one 
to four. In a few specimens the basal 
band_is weakly continuous. 


Legs pale to dark yellow. 
Abdomen pale yellow to dark yellow- 


ish brown. Male terminalia pale to 


dark yellow. Ninth tergum, as in Fig. 
117, elongate, broad basally, apex emar- 
ginate, forming two broad lateral lobes. 
Hypovalves (Fig. 132) broad, ex- 
panded medially, apices rounded, ex- 


Aug,, 1975 


tending three-fourths length of basi- 
styles. Basistyles broad. Dististyles 
about two-thirds length of basistyles, 
each dististyle with small patch of elon- 
gate setae near base. Ventral parameres 
(Fig. 133) branched, elongate, barbed, 
each with apical branch extending 
slightly beyond base of dististyle. Fe- 
male genital plate (Fig. 160) short, 
oval, 1.17 mm in length. Distal plate 
oval, broad basally, emarginate api- 
cally, forming two lateral lobes. Basal 
plate absent. Spermathecal apodeme 
elongate, widely bifurcate basally, not 
reaching apical emargination. 

The male terminalia of P. speciosa 
are indistinguishable from those of P. 
braueri although these species can be 
separated by the characters of the basal 
band. In the holotype of P. braueri, the 
ventral parameres are very similar to 
those of P. speciosa in being branched, 
but the mesal branch in P. braueri is 


Fig. 183.—LDistribution of Panorpa speciosa 
in Illinois and North America. 


Wess Et Au.: MECOPTERA OF ILLINOIS 


307 


somewhat thicker and larger than that 
in P. speciosa. In females the genital 
plate of P. speciosa is much longer than 
it is in P. braueri. 

This species has been collected abun- 
dantly in Illinois on stinging wood net- 
tle, jewelweed, and poison ivy in hu- 
mid shaded areas along slow-moving 
streams. 

P. speciosa is a north-central species 
extending from Arkansas and Tennessee 


to Minnesota and Wisconsin (Fig. 
183). 
Illinois Records.—(Fig. 183). Col- 


lected frequently from late April until 
early November. The prolonged collec- 
tion period indicates the possibility of 
two generations per year. 


Panorpa braueri Carpenter 
Panorpa braueri Carpenter (1931a: 

242). ¢, 2. Type-locality: Wash- 

ington County, Arkansas. 

Head and thorax dark yellowish 
brown. 

Fore wing length 10.0-11.4 mm. 
Membranes (Fig. 86) pale yellow, 
crossveins margined. Apical band dark 
brown, entire, with several small clear 
spots. Pterostigmal band dark brown, 
broad from anterior margin to poste- 
rior, apical fork broken, small. Basal 
band dark brown, broad, continuous. 
Two marginal spots and first basal spot 
dark brown. Second basal spot absent. 

Legs dark yellowish brown. 

Abdomen dark reddish brown. Male 
terminalia dark yellowish brown. Ninth 
tergum, as in Fig. 117, elongate, base 
broad, apex emarginate, forming two 
broad lateral lobes. Hypovalves (Fig. 
134) broad, expanded medially, apices 
rounded, extending three-fourths length 
of basistyles. Basistyles broad. Disti- 
styles about two-thirds length of basi- 
styles, each with small patch of elon- 
gate setae at base of inner basal cusp. 
Ventral parameres (Fig. 135) narrow, 
elongate, each with broad mesal branch 
and slender apical branch extending 
beyond base of dististyle. In ventral 
view the slender apical branch is often 


308 


hidden, giving the paramere the ap- 
pearance of having a single, broad, 
bulbous apex. Female genital plate 
(Fig. 161) small, broad, 0.69 mm in 
length. Distal plate broad, deeply emar- 
ginate apically, forming two broad lat- 
eral lobes. Basal plate absent. Sperma- 
thecal apodeme short, not reaching api- 
cal emargination of distal plate. 

P. braueri is very closely related to 
P. speciosa, and little difference exists 
in the characters of the male terminalia. 
These species may be separated by the 
broad, continuous basal band in the 
wing of P. braueri. In females of P. 
braueri the genital plate is much 
shorter than that of P. speciosa. 

Byers (1954) reported collecting 
Missouri specimens of P. braueri on 
small patches of Impatiens in a shaded 
swale. 


P. braueri seems restricted to north- 
western Arkansas and southern Mis- 
souri (Fig. 184). 


Fig. 184.—Distribution of Panorpa braueri 
in North America. 


Panorpa bifida Carpenter 


Panorpa bifida Carpenter (1935:107). 
$, 2. Type-locality: Rector, Penn- 
sylvania. 


Head and thorax dark yellowish 
brown. 


Fore wing length 12 mm. Membranes 
(Fig. 88) pale yellow, crossveins not 
margined. Apical band pale brown, 
entire, with one or two clear spots. 
Pterostigmal band pale brown, contin- 
uous, apical work broken. Basal band 
pale brown, broken into two large 


Inurnois NatrurAL History SurvEY BULLETIN 


Vol. 31, Art.7 


spots. Two marginal spots and large 
first basal spot present. Second basal 
spot absent. 

Legs pale yellow. 

Abdomen dark yellowish brown. 
Male terminalia dark yellowish brown. 
Ninth tergum elongate, deeply emar- 
ginate apically, forming two narrow 
lateral lobes. Hypovalves (Fig. 136) 
broad, extending almost to bases of dis- 
tistyles. Basistyles broad. Dististyles 
each with small patch of elongate setae 
near base. Ventral parameres (Fig. 
137) narrow, elongate, each with two 
thin, barbed, branches extending be- 
yond base of dististyle, united basally 
to form Y-shaped projection. Female 
genital plate (Fig. 162) broad, 0.99 mm 
in length. Distal plate broad, deeply 
emarginate apically to form two broad 
lateral lobes. Basal plate absent. Sper- 
mathecal apodeme elongate, not reach- 
ing apical emargination of distal plate. 

P. bifida is related to P. anomala, but 
it is easily distinguished from P. anom- 
ala by the narrow elongate branches 
of the ventral parameres. 


Nothing has been reported on the 
habitat of this species. 

P. bifida is known only from Pennsyl- 
vania and Ohio (Fig. 185). 


Fig. 185.—Distribution of Panorpa bifida 
in North America. 


Panorpa anomala Carpenter 
Panorpa anomala Carpenter (1931a: 
245). 8, 2. Type-locality: Leaven- 
worth County, Kansas. 
Panorpa proximata Carpenter (1931a: 
247). 8. Type-locality: Washington 


Aug., 1975 


County, Arkansas. Synonymized by 

Byers (1974). 

Head and thorax dark yellow to dark 
reddish brown. 

Fore wing length 10.6-12.4 mm. 
Membranes (Fig. 89) pale yellow, 
crossveins faintly margined. Apical 
band dark brown to black, broad, usu- 
ally entire. Pterostigmal band dark 
brown, broad anteriorly, broken pos- 
teriorly. Basal band broken, forming 
two dark brown spots. Two marginal 
spots and first basal spot small. Sec- 
ond basal spot absent. In most speci- 
mens the apical band of the fore wing 
is broad and entire although several 
specimens showed the apical band 
broken into several small dark brown 
spots and a narrow subapical band. 
In some specimens a second basal spot 
was present and the second marginal 
spot absent. 

Legs dark yellow. 

Abdomen dark yellow to dark red- 
dish brown. Male terminalia dark red- 
ish brown, oval. Ninth tergum, as in 
Fig. 117, emarginate apically, forming 
two broad lateral lobes. Hypovalves 
(Fig. 140) broad apically, ending well 
before bases of distisyles. Basistyles 
broad. ODististyles shorter than ba- 
sistyles. Ventral parameres (Fig. 141) 
elongate, barbed, with one branch ex- 
tending posteriorly beyond bases of 
dististyles and a mesal branch curved 
dorsally. Female genital plate (Fig. 
163) oval, 1.15 mm in length. Distal 
plate short, deeply emarginate apically, 
forming two acute lateral lobes. Basal 
plate absent. Spermathecal apodeme 
elongate, bifurcate basally, not reach- 
ing apical emargination of distal plate. 


The shape of the male terminalia of 
P. anomala resemble somewhat those 
of P. elaborata but differ markedly 
in the short hypovalves and heavily 
barbed ventral parameres. 


In Illinois P. anomala was initially 
collected along the bottomlands of the 
Illinois River at Morris in a dense 
growth of stinging wood nettle. Indi- 


Wess Et At.: MECOPTERA OF ILLINOIS 


309 


viduals in southern Illinois were col- 
lected on jewelweed in shaded areas 
along small creeks. 

P. anomala is a western species, oc- 
curring from southeastern Tennessee 
and northwestern Georgia west to Wis- 


consin, Kansas, and Arkansas (Fig. 
186). 


Fig. 186,—Distribution of Panorpa anomala 
in Illinois and North America. 


Illinois Records.—(Fig. 186). Col- 
lected from late May until mid-August 
in northern, western, and southern IIli- 
nois. Carpenter (193la) erroneously 
recorded P. anomala in Illinois from 
Starved Rock State Park on the basis 
of an imperfect female. This specimen 
has since been identified as a female of 
P. speciosa. 


Panorpa consuetudinis Snodgrass 


Panorpa consuetudinis Snodgrass 
(1927:77). ¢. Type-locality: Ta- 
koma Park, Maryland. Neotype ¢ 
designated by Byers (1974). 


310 


Panorpa elaborata Carpenter (1931a: 
239). &, 9. Type-locality: Falls 
Church, Virginia. Synonymized by 
Byers (1974). 

Head and thorax dark yellowish 
brown. 


Fore wing length 10.0-11.0 mm. 
Membranes (Fig. 90) amber, cross- 
veins margined. Apical band dark 
brown, broad, with several subapical 
clear spots. Pterostigmal band dark 
brown, broad anteriorly, forked pos- 
teriorly. Basal band continuous or 
broken. Marginal and first basal spots 
small. Second basal spot lacking. 


Legs dark yellow. 


Abdomen dark yellow. Male termi- 
nalia dark yellow. Ninth tergum elon- 
gate; base broad, tergum constricted 
beyond middle, apex deeply emargin- 
ate, forming two narrow lateral lobes. 
Hypovalves (Fig. 138) narrow, elon- 
gate, extending to bases of dististyles. 
Basistyles broad. Dististyles shorter 
than _basistyles. Ventral parameres 
(Fig. 139) extend beyond bases of 
dististyles, each paramere with two 
branches, mesal branch barbed, apical 
branch with two tufts of barbs. Female 
genital plate (Fig. 164) short, 0.85 mm 
in length. Distal plate concave apically, 
not deeply emarginate, sides parallel. 


Itutinois NaturaL History Survey BULLETIN 


Vol. 31, Art. 7 


Basal plate absent. Large, subrectangu-— 
lar, sclerotized membrane covers most | 
of distal plate. Spermathecal apodeme 
elongate, bifurcate basally, not reach- 
ing apical emargination of distal plate. 

The male terminalia of P. consue- 
tudinis are similar to those of P. dubi-— 
tans although differing in the longer 
hypovalves and the branched ventral 
parameres. : 

Little is known of the specific habitat — 
of P. consuetudinis. In Kentucky indi-— 
viduals were collected with specimens — 
of P. insolens in densely shaded vegeta- 
tion along a slow-moving stream. : 

P. consuetudinis is an eastern species, — 
extending from South Carolina to New 
York and west to Indiana and Mis-— 
sissippi (Fig. 187). 


Fig. 187.—Distribution of Panorpa consue- 
tudinis in North America. 


LITERATURE CITED 


Banks, N. 1895. New neuropteroid insects. 
American Entomological Society Trans- 
actions 22:313-316. 


1900. New genera and species of 
Nearctic neuropteroid insects. American 
Entomological Society Transactions 26: 
239-259. 


. 1907. Catalogue of the neuropteroid 
insects of the United States. American 
Entomological Society, Philadelphia. 53 p. 


. 1908. Neuropteroid insects — notes 
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INDEX 


A 
Anabittacus, 269 
Anomalobittacus, 269 
Apterobittacus, 252, 261, 267-269 
apterus, 269 
Apteropanorpidae, 251 
Arctotertiary, 251 


_ Ateleptera, 277 
_ Atrichum angustatum, 258, 260 


Aulops, 282 

Austrobittacus, 269 
Austromerope, 280 
Austroriparian Division, 251 


B 
Bait trap, 281 
Baltic amber, 264 
Berlese funnel, 266 
Bittacidae, 251-252, 257, 260-262, 268-269 
Bittacus, 252-255, 260-262, 265, 268-271 
apicalis, 252, 255, 258, 262, 266, 270-273, 
276 
arizonicus, 274 
chlorostigma, 262 
italicus, 269 
occidentis, 255, 260, 262, 266, 270-271, 
274 
pallidipennis, 275 
pilicornis, 252, 255, 259, 262, 269-271, 
273-275 
punctiger, 251, 255, 262, 270-273, 275 
stigmaterus, 255, 262, 270-271, 275-277 
strigosus, 252, 255, 260, 262, 266, 269-271, 
273, 275-276 
texanus, 262, 270-271, 276-277 
Boreidae, 251, 253, 260, 262-265, 268, 277 
Boreus, 251, 253-254, 256, 266-267, 277-279 
brevicaudus, 277 
brumalis, 253, 256, 260, 278-280 
californicus, 253-254 
elegans, 277 
hyemalis, 253-254, 256, 277 
nivoriundus, 278-280 
reductus, 277 
Brachypanorpa, 264, 268, 282 
carolinensis, 282 
montana, 282 
oregonensis, 282 


Cc 
Chafer trap, 281 
Choristidae, 251 
Coastal Plain Division, 251 
Cotton, 260, 293 


D 


Dicranella heteromalla, 260 


| Diplostigma, 269 


Diptera, 252 
Dolichopodids, 252 


E 
Edriobittacus, 269 
Elms, 277 
Eocene, 262 
Estanella, 282 
Euboreus, 277 


G 


Gondwanaland, 262 
Gooseberry, 260, 273 


H 

Hangingflies, 251 
Haplodictyus, 269 

incertus, 272 
Harpobittacus, 269 
Homoptera, 252 
Hydrophyllum appendiculatum (see water- 

leaf) 

Hypandrium, 277, 279 


Impatiens, 259-260, 308 (see jewelweed) 
Issikiella, 269 


J 
Jewelweed, 259, 272, 273, 298, 302, 304, 307, 
309 
Jurassic, 261 


K 
Kalobittacus, 269 


L 
Laportea canadensis (see stinging wood 
nettle) 
Leptobittacus, 269 
Leptopanorpa, 282 


M 
Malaise trap, 260, 266, 281 
Mecaptera, 268 
Mecoptera, 251, 260-261, 263, 265-269, 277, 
280, 282 
Meropeidae, 251, 253, 256, 260, 268, 268, 280 
Merope, 263, 266-268, 280-281 
tuber, 280-281 
Meropidae, 280 
Mesozoic, 262 
Multiflora rose, 260, 273 


N 
Nannobittacus, 269 
Nannochoristidae, 251 
Neobittacus, 269 
Neopanorpa, 282 
Northeast Morainal Division, 251 
Notal organ, 261, 263-264, 280 
Notiothaumidae, 251, 280 
Notiothauma, 268 


315 


316 TIuuinois NaTurRAL History SuRvEY BULLETIN Vol. 31, Art. 7 


re) neglecta, 296 
Oligocene, 264 nuptialis, 251, 254, 256, 265, 283, 285, 
Ozark Division, 251 287, 291-293 
Ozark-Ouachita uplift, 265 Pe ane dae 

k uplift, 251, 279 rufa, 292- 

CAneeEE rufescens, 256, 264, 284, 287, 289, 291- 

Pp 292, 304-305 

rufescens group, 256, 287, 299 

Panorpatae, 268 : j 
Panorpidae, 251, 253, 260, 262, 264, 268, 282 cate Gees” 267, 283, 286-287, 
Panorpa, 253-254, 256, 260-261, 264-268, 282— Beebe 7: 

286, 288-290, 293-294, 298-299 Ha Nan RA: seiae 
acuta, 283, 286, 288, 291, 295-296, 298 oa 4, 265, 284, 287, 290-292, 306— 
ee a subfurcata, 251, 283, 287-288, 291-299, 

a, 300-301 
Pe EO rae Ug ty waa ae submaculosa, 283-285, 287, 291, 293-294 
308-309 ¢ a : : utahensis, 293-295 
banksi, 264, 283, 286-287, 291, 295-298 Mensa S021 305 
banksii, 296 virginica, 301 
5 Ge tee tue. Be Panorpodidae, 251, 262, 264, 268, 281-282 
Dare d! 2 z Panorpodes, 264, 281-282 
braweri, 251, 265, 284, 287, 290-292, 307— f 
308 Pazius, 269 


Permian, 251, 260 
Picric acid trap, 281 
Pinacula, 257 


canadensis, 303 
chelata, 296 
claripennis, 284, 287, 289, 291-292, 295, 


Pleistocene, 262 
caer Pliocene, 262 
communis, 253-254, 282 Poison ivy, 260, 302, 307 


confusa, 305 ome Fas oplars) 
consuetudinis, 264, 284, 287, 290-291, Pp cePon 


Prepupa, 255 
309-310 
debilis, 256, 264, 284, 287, 289, 291-292, Prepupal cell, 257 
Probittacus, 262 
303-305 Protobittacus, 262 
dubitans, 265, 284, 287, 289, 291-292, , 


305-306, 310 R 
elaborata, 264, 309-310 
galerita, 251, 283, 287-288, 292, 299-301 
germanica, 254 
helena, 254, 256, 264, 266, 284, 287, 289, 
291-292, 302-303 5 Ss 
hungerfordi, 288, 287-288, 291-292, 301- Scorpionflies, 251 


302 = Shawnee Hills Division, 251 
insolens, 284, 287, 289, 291-292, 302-303, Soybeans, 260, 293 


Rhus radicans (see poison ivy) 
Ribes (see gooseberry) 
Rosa multifiora (see multiflora rose) 


310 Stinging wood nettle, 260, 272-273, 298, 302, 
klugi, 256 306-307, 309 
latipennis, 288, 285, 287, 291, 295, 304 Sweet william, 253 
longipennis, 295 
lugubris, 292-293 T 
lugubris group, 256, 260, 282-283, 292 Tertiary, 262 
maculosa, 283-285, 287, 291, 293-294 Thyridates, 269 
mirabilis, 251, 288, 287-288, 291-292, Tobacco, 260 
299-301 = 
modesta, 300 W 


nebulosa, 264, 283, 286-288, 291, 296, 298  Waterleaf, 260 
nebulosa group, 287, 293, 296 Willows, 277 


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and the Western Corn Rootworm, 
brotica virgifera LeConte (Colec 
Chrysomelidae). By W. H. Lue 
H. C. Chiang, E. E. Ortman, and 
P. Nichols. April, 1974. 15 p. 


91.—The Distribution of Periodical 
in Illinois. By Lewis J. Stanni 
February, 1975. 12 p. 


92.—The Literature of Arthropods 
ated with Soybeans. IV. A Bib 
of the Velvetbean Caterpillar A7 
gemmatalis Hubner (Lepidopte 
tuidae). By B. J. Ford, J. R. § 
Reid, and G. L. Godfrey. Febru: 
15 p. - 
93——The Life History of the St 
Darter, Etheostoma kennicotti, 
Creek, Illinois. By Lawrence M. 
February, 1975. 15 p. 


94.—Illinois Pheasants: Their Dist1 
and Abundance, 1958-1973. By Ro 
Labisky. February, 1975. All Dp. 


95.—The Nest Biology of the Bee A 
(Ptilandrena) erigeniae Robertson 
menoptera: Andrenidae). By 0 
Davis, Jr. and Wallace E. LaBergi 
1975. 16 p. 76 


CIRCULAR 

51.—Illinois Trees: Selection, Plantin; 
Care. By J. Cedric Carter. August, 
123 p. 

52.—Fertilizing and Watering Trees. | 
Dan Neely and E. B. Himelick. 
ber, 1971. (Third printing.) 20 PB 


54—Corn Rootworm Pest Manageme 
Canning Sweet Corn. By W. H. 
mann, J. T. Shaw, D. E. Kuhlma 
Randell, and C. D. LeSar. March, 
10 p. 


e 
a 


| ILLINOIS 
itural History Survey 
" BULLETIN 


An Electrofishing Survey 
of the Illinois River, 
1959-1974 


rd E. Sparks 
m C. Starrett 


MENT OF REGISTRATION AND EDUCATION 


IRA L HISTORY SURVEY DIVISION 
INA, ILLINOIS 

q é VOLUME 31, ARTICLE 8 
AUGUST, 1975 - 


| US LSSN UU O—4915 


ILLINOIS 
atural History Survey 
BULLETIN 


An Electrofishing Survey 
of the Illinois River, 
1959-1974 


rd E. Sparks 
m C. Starrett 


| 
f 


ILLINOIS 
MENT OF REGISTRATION AND EDUCATION 


IRAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 


VOLUME 31, ARTICLE 8 


STATE OF ILLINOIS 


GuTowsky, Ph.D., Chemistry; Robert H. ANDERSON, 


Illinois University. 


NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 
SCIENTIFIC AND TECHNICAL STAFF 
GEORGE SPRUGEL, JR., Ph.D., Chief 
AuicE K, Apams, Secretary to the Chief 


Section of Economic Entomology 

Wituiam H,. Luckmann, Ph.D., Entomologist and Head 

Wiis N. Bruce, Ph.D., Entomologist 

Wayne L. Hown, Ph.D., Entomologist 

STEVENSON Moore, III, Ph.D., Entomologist, Extension 

JAMES E. APPLEBY, Ph.D., Associate Entomologist 

Epwarp J. ARMBRUST, Ph.D., Associate Entomologist 

Marcos Koean, Ph.D., Associate Entomologist 

JosEPH V. Mappox, Ph.D., Associate Entomologist 

Ronaup H. MEYER, Ph.D., Associate Entomologist 

RoBeErT D. PauscuH, Ph.D., Associate Entomologist 

RALPH E. SECHRIEST, Ph.D., Associate Entomologist 

JoHN K. BousEMan, M.S., Assistant Entomologist 

GeorGE L, GopFREY, Ph.D., Assistant Entomologist 

MicHagEL E, Irwin, Ph.D., Assistant Entomologist 

DonaLp E, KuHuman, Ph.D., Assistant Professor, 
Extension 

Roscoe RANDELL, Ph.D., Assistant Professor, Extension 

Witiiam G. Ruesink, Ph.D., Assistant Entomologist 

JAMES R. SANBORN, Ph.D., Assistant Entomologist 

Dovueuas K. SELL, Ph.D., Assistant Entomologist 

C. Ropert Taytor, Ph.D., Assistant Entomologist 

JouN L. WEDBERG, Ph.D., Assistant Entomologist 

CLARENCE E. WHITE, B.S., Assistant Entomologist 

Tim Coo.ey, M.A., Assistant Specialist, Extension 

Kurt E. RepporG, M.S., Assistant Specialist 

Joun F. Watt, M.S., Assistant Specialist, Extension 

JEAN G. WILSON, B.A., Supervisory Assistant 

STEPHEN Rosexts, B.S., Junior Professional Scientist 

JouNn T. SHaw, B.S., Junior Professional Scientist 

DaNIEL P. BARTELL, Ph.D., Research Associate 

Bettina Francis, Ph.D., Research Associate 

MARGARET ANDERSON, B.S., Research Assistant 

Rozert J. BARNEY, B.S., Research Assistant 

Tzu-Suan Cuu, M.S., Research Assistant 

STEPHEN D. Cowan, B.S., Research Assistant 

STEPHEN K. EvrarD, B.S., Research Assistant 

Marion Farris, M.S., Research Assistant 

Bonnie Irwin, M.S., Research Assistant 

JENNY Koaan, M.S., Research Assistant 

GuEeNN Levinson, B.S., Research Assistant 

RosE ANN MECCOLI, B.S., Research Assistant 

Brian MELIN, B.S., Research Assistant 

Ceuia SHIH, M.S., Research Assistant 

Katuy Woop, M.S., Research Assistant 

Jo ANN AUBLE, Technical Assistant 

LowELi Davis, Technical Assistant 

CuHaRLes G. Hew, M.S., Technical Assistant 

Linpa IsENHOWER, Technical Assistant 

Lu-Pine LEE, M-S., Technical Assistant 


Section of Botany and Plant Pathology 

Cuaus GRuNWALD, Ph.D., Plant Physiologist and Head 
Rosert A, Evers, Ph.D., Botanist 

EvuGeENE B, Himevick, Ph.D., Plant Pathologist 

R. Dan NEELY, Ph.D., Plant Pathologist 

D. F. SCHOENEWEISS, Ph.D., Plant Pathologist 

J. LELAND CRANE, Ph.D., Associate Mycologist 
WALTER HaRTSTIRN, Ph.D., Assistant Plant Pathologist 
Betty S. NELson, Junior Professional Scientist 

GENE E. REID, Technical Assistant 


Section of Aquatic Biology 


D. Homer Buck, Ph.D., Aquatic Biologist 

WiuutaM F, Cuinpers, Ph.D., Aquatic Biologist 

R. WeLpon Larimore, Ph.D., Aquatic Biologist 
RoBert C. HiLTIBRAN, Ph.D., Biochemist 

ALLISON BRIGHAM, Ph.D., Assistant Aquatic Biologist 
WarRREN U. BriGHAM, Ph.D., Assistant Aquatic Biologist 
Ricuarp E, Sparks, Ph.D., Assistant Aquatic Biologist 
Tep W. Storck, Ph.D., Assistant Aquatic Biologist 
JOHN TRANQUILLI, Ph.D., Assistant Aquatic Biologist 
Mary FraNcES BiaL, Junior Professional Scientist 
Cart M. THompson, Junior Professional Scientist 
RicHarp J. Baur, M.S., Research Associate 

Donautp W. Durrorp, M.S., Research Associate 

Joun M. McNorney, M.S., Research Associate 

Harry W. BeraMann, B.S., Research Assistant 


CONSULTANTS AND RESEARCH AFFILIATES: 


life Research, Southern Illinois University ; 


DEPARTMENT OF REGISTRATION AND EDUCATION 
BOARD OF NATURAL RESOURCES AND CONSERVATION 


Ronaup E. Stacker, J.D., Chairman; THomas Park, Ph.D., Biology; L. L. Stoss, Ph.D., Geology; HERBERT S® 
B.S.C.E., Engineering ; 
senting the President of the University of Illinois; JoHn C. Guyon, Ph.D., Representing the President of Southe 


Systematic EntomMoLocy, RopEricK R. Irwin, Chicago, i fe 
nois ; WILDLIFE RESEARCH, WILLARD D. Kuimstra, Ph.D., Professor of Zoology and Director of Cooperative Wi ‘ 
PARASITOLOGY, Norman D. Levine, Ph.D., Professor of Veterin 
Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, Unive of 
Illinois; ENTomouocy, Ropert L. Metcaur, Ph.D., Professor of Zoology and of Entomology, University of Illinois; 
and GILBERT P. WALDBAUER, Ph.D., Professor of Entomology, University of Illinois; 
Norton, Ph.D., Professor of Statistical Design and Analysis, University of Illinois. 


W. L, Everitt, E.E., Ph.D., Reprew 


Kurt T. CLEMENT, B.S., Research Assistant 
Larry W. Courant, M.S., Research Assistant 
HersBert M. Dreigr, M.S., Research Assistant 
MicHakt A. Frakes, M.S., Research Assistant 
Tuomas EK. HILL, M.S., Research Assistant 
Earu THomas Joy, JR., M.S., Research Assistant 
RicuarD Kocuer, B.S., Research Assistant 
Ropert Moran, M.S., Research Assistant 
Katuryn Ewine, B.S., Technical Assistant 
Susan Moore, Technical Assistant 

FLORENCE PARTENHEIMER, B.A., Technical Assistant 
C. Russevt Rose, Field Assistant 


Section of Faunistic Surveys and 
Insect Identification | 


Puiuie W. Smita, Ph.D., Taxonomist and Head 
Wa acer E. LaBerGe, Ph.D., Taronomist ; 
MILTON W. SANDERSON, Ph.D., Tazonomist 

Lewis J. STANNARD, JR., Ph.D., Tazonomist 

Larry M. Pace, Ph.D., Assistant Taxonomist 
JoHN D, Unzicker, Ph.D., Assistant Taxonomist 
DonaLp W. WEBB, M.S., Assistant Taronomist 
BERNICE P. SWEENEY, Junior Professional Scientist 
Craig W. Ronto, Technical Assistant 


Section of Wildlife Research 


GurEn C. SanpErson, Ph.D., Wildlife Specialist and Head: 
Frank C, BELLROSE, B.S., Wildlife Specialist 
JEAN W. GRABER, Ph.D.,. Wildlife Specialist 
RicHarD R, GraBer, Ph.D., Wildlife Specialist 
Haroutp C, Hanson, Ph.D., Wildlife Specialist 
Ronaup F. LaBisky, Ph.D., Wildlife Specialist 
Wiwiram L, ANDERSON, M.A., Associate Wildlife 
Specialist 
W. W. Cocuran, JR., B.S., Associate Wildlife Specialist) 
Wiuuiam R. Epwarps, Ph.D., Associate Wildlife 
Specialist % 
G. Buarr JoseELyN, M.S., Associate Wildlife Specialist 
Cuarues M. Nixon, M. S., Associate Wildlife Specialist — 
KENNETH E. SMitH, Ph.D., Associate Chemist 
RicHarD E, WaRNER, M.S., Associate Wildlife Speci 
RonaLp L. WESTEMEIER, M.S., Associate Wildlife 
Specialist ¢ 
STEPHEN P. Havera, M.S., Assistant Wildlife Specialis 
Davip R. Vance, M.S., Assistant Wildlife Specialist 
Ronaup E. Duzan, Junior Professional Scientist ‘i 
HELEN C. ScuuLtz, M.A., Junior Professional Scientist 
ELEANORE WILSON, Junior Professional Scientist 
SHARON FRADENBURGH, B.A., Laboratory Technician 
Rosert D. Crompton, Field Assistant 
James W. SEETs, Laboratory Assistant 


ac 


Paiva 


Section of Administrative Services ; 
RoBert O. Watson, B.S., Administrator and Head 


Supporting Services 

Witma G. Dittman, Property Control and Trust 

Accounts 
Patty L. Duzan, Technical Assistant 
Rosert O. Evuis, Assistant for Operations 
Larry D. Gross, Maintenance Supervisor 
Luoyp E. HUFFMAN, Stockroom Manager 
J. Wiuu1am Lusk, Mailing and Distribution Services bE 
JERRY McNEAR, Maintenance Supervisor 
MEtvin E. ScHWARTZ, Financial Records 
JAMES E. SERGENT, Greenhouse Superintendent 


Publications and Public Relations 
Rospert M. ZEwapski, M.S., Technical Editor 
SHIRLEY MCCLELLAN, Assistant Technical Editor 
LAWRENCE S. Faruow, Technical Photographer 
Luoyp LEMERE, Technical Illustrator 


Technical Library 
Doris F. Dopps, M.S.L.S., Technical Librarian 
Doris L. SUBLETTE, MS.L.S., Assistant Technical 
Librarian 


Sratistics, Horace W.! 


CONTENTS 
SMR ERD CRTEIN TSE Aerie tals fats elec Os. rele Sisie: ee. stomioers eine <x ee Poemeeeees oe 317 
EPIL IES Be eteec fete ate olay niet cj oial eis! asics ate v c2) aualeedid win Gaatactt Sane s oe tke wee 318 
—ENTLIE 2 elds 6 eG ee seo Gio SOO ION NOR RENOID ORCS Cer ce eons ch ne cette iste ears 319 
Bhysical: Chemical Results) sticte tis. ai. veces acne oa Haeae Sees 319 
bifecino shine m@esulisMena- ce nus.as os lasalitsts viociatehuce Stsioaele CoM are 321 
© LE STOSSICOIN’ 3's vero leg! ee sh reap oR eg i er 332 
-_ Historical Changes in the Fish Populations of the Illinois River ...... 332 
Future Impacts on the Fish Populations of the Illinois River ........ 342 
MRO UAS MP RE Ne ete ei ccc «ys ue, shel evoye: oi oyaie. ©. 4% cle. Ghe ieioe Ae ihe qe ensmnege wuetors 344 
BRIA MUI OEEED katte. cieicv setts Piola rss aie) actialetsla: slnte ate ale So eins eretedina yest 377 
| THIBS . o.calh lly g.5 Oot or CRRETEES CE cen Oe ee ane Ae ae ea 378 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. 
It is a contribution from the Section of Aquatic Biology of the Illinois Natural History 
Survey. 


Richard E. Sparks is an Assistant Aquatic Biologist, and the late William C. Starrett 
was an Aquatic Biologist, at the Illinois Natural History Survey. 


(66938—4M—8-75) 


EZ 


i) 


An Electrofishing Survey 


of the Illinois River, 1959-1974 


FROM AS FAR BACK as historical ac- 
counts are available, the Illinois River 
Valley has been described as unusually 
productive of fish and wildlife. The 
French explorer Marquette wrote in 
1673 (Mills, Starrett, & Bellrose 1966: 
3-4): 

“We have seen nothing like this 
river that we enter, as regards to 
its fertility of soil, its prairies and 
woods; its cattle, elk, deer, wildcats, 
bustards, swans, ducks, parroquets, 
and even beaver.” 

When Illinois was still a territory, 
the Illinois River Valley was considered 
one of the important sources of furs 
in the northwest part of the United 
States (Starrett 1972:139). There are 
older residents of the valley who recall 
the importance of fish and wildlife to 
some of the river towns in the early 
part of the century. Hugh Bell, Super- 
intendent of the Illinois Department 
of Conservation Fisheries Field Head- 
quarters at Havana, as a young man 
worked at filling specially constructed 
tank cars with fish to be shipped by 
rail from Havana to Chicago. At one 
time live fish also were shipped regu- 
larly to Boston and New York, and the 
Illinois River ranked as a major inland 
commercial fishery. There was a U.S. 
government fisheries station at Mere- 
dosia ( Forbes & Richardson 1920:XVI). 
During that same period a train called 
the Fisherman’s Special ran between 
Springfield and Havana, and there were 
many people in Havana who made their 
living outfitting and guiding fishermen 
and duck hunters. 

Because of their importance as 
unique resources, the Illinois River and 
its bottomland lakes were studied in- 
tensively by the Illinois State Labora- 


| tory of Natural History and its succes- 


317 


Richard E. Sparks 
William C. Starrett 


sor, the Natural History Survey, from 
1874 to 1927 (Forbes 1928:387). More 
recently, surveys of the fish populations 
of the river have been conducted regu- 
larly from the 1940's to the present. 
Various types of sampling gear have 
been employed in these surveys, for 
various purposes. For example, min- 
now seines were used regularly in mid- 
summer to collect small fish and there- 
by gauge the spawning success of 
species which spawn in the spring. 
Hoop nets were used to collect large 
fish in backwaters and bottomland 
lakes. The present report concerns pri- 
marily the electrofishing surveys, which 
have been conducted regularly in the 
Illinois River in the fall, from 1959 
through 1974. 


ACKNOWLEDGMENTS 


The electrofishing survey of the IlIli- 
nois River was conceived and carried 
out, for the most part, by Dr. William 
C. Starrett, Aquatic Biologist, Havana 
Field Laboratory, Illinois Natural His- 
tory Survey. Dennis L. Dooley worked 
on the electrofishing survey, and other 
Illinois River studies, for 9 years. Robert 
Crompton, Howard Crum, and Ron 
Barker also assisted in the project un- 
der Dr. Starrett’s direction. 

Following Dr. Starrett’s death in De- 
cember, 1971, the electrofishing survey 
was resumed in 1973 by the writer and 
Kenneth Walker, with assistance in 
locating stations and following previ- 
ously established methods from Mr. 
Dooley. Carl M. Thompson assisted 
with the 1974 electrofishing and helped 
compile and analyze data for this re- 
port. 

We thank Lloyd LeMere for drawing 
the figures, Dr. R. Weldon Larimore 


318 


for reviewing the manuscript, O. F. 
Glissendorf for final editing, and Judith 
L. Breckenridge who did the typing. 
We are grateful to all the students 
and other assistants who helped with 
the program from 1959 through 1974. 

Finally, the electrofishing survey 
could not have been continued in 1974 
without the support of the St. Louis 
District and the Waterways Experi- 
ment Station of the U.S. Army Corps 
of Engineers. 


PROCEDURE 

Twenty-four sampling sites were 
chosen in 1959 that provided good 
habitat for adult fish and that were 
fairly well distributed throughout the 
length of the river (Table 1). The 
same sites were usually sampled in 
succeeding years, except that one addi- 
tional station, Big Blue Island Chute, 
was sampled in 1974. Most of the sites 
are in chutes, that is, side channels of 
the river, and contain brush piles, un- 
dercut banks, and “holes” where vari- 
ous species of fish are apt to congregate. 
The four exceptions to this general de- 
scription are (1) the station above 
Pekin where both sides of the main 
channel were fished, (2) the station 
along the shore of Lower Peoria Lake, 
(3) the station in Middle Peoria Lake 
where docks and riprapping in various 
marinas were fished in the 1960’s and 
where riprapping at a state conserva- 
tion landing in Detweiller Park was 
fished in the 1970's, and (4) a station 
in the Des Plaines River where the 
wide mouth of the Du Page River and 
a boatyard were fished. The stations 
are located most accurately by river 
mile*—the exact number of miles up- 
stream from the mouth of the river 
at Grafton, based on the Corps of En- 
gineers’ chart book of the Illinois 
Waterway (U.S. Army Engineer Dis- 
trict, Chicago 1970). A river mile 
designation shows the approximate area 
that was fished. For example, at the 


* Stations are located by river miles rather 
than by kilometers because existing river 
charts and navigation aids along the river use 
mileages. 


ILturnois NATuRAL History SURVEY BULLETIN 


Vol. 31, Art. 8 


first station listed in Table 1, we fished 
that part of Mortland Island Chute 
which extended from mile 18.7 to mile 
19.4. 


Pools in Table 1 refer to the waters 
impounded behind the dams and locks” 
for navigation. Throughout this paper, 
references are made to these pools as 
convenient geographic locations of the 
various sections of the river. The lower 
part of the Illinois River is under the in- 
fluence of the Alton Dam on the Missis- 
sippi. The dams forming the other pools, 
in upstream order, are: La Grange (river 
mile 80.2), Peoria (mile 157.6), Starved 
Rock (mile 231.0), Marseilles (mile 
247.0), and Dresden (mile 271.5). Be- 
cause the upstream pools are shorter 
than the downstream ones, there are 
fewer stations in the upstream pools 

The Illinois River begins at the con- 
fluence of the Des Plaines and Kanka- 
kee Rivers, and a distance of only 14 
miles (2.25 km) separates the conflu 
ence and Dresden Dam. The Dresden 
Pool extends into the Des Plaines and 
Kankakee Rivers, and our one sampling 
station in the Dresden Pool is actually} 
located in the Des Plaines River. 
Kankakee is a relatively unpolluted 
stream, while the Des Plaines River 
receives municipal and industrial efflu 
ents from the Chicago metropolitan 
area, via the Chicago Sanitary and Ship 
Canal. The Des Plaines station is ex- 
cluded when results from the Illinois 
River stations are used to compute 
average yearly catches per unit effort 
for the whole Illinois River (Tables 
3-27). 

The navigation dams help to mai 
tain a 2.74-m deep navigation channel 
by impounding water during low-flow 
periods. When the water is thus i 
pounded the river behind the dam is 
said to be at pool stage. In order to 
sample under similar environmental 
conditions from year to year, electro- 
fishing was conducted at the same time 
every year, from late August to the 
middle of October, and only when the 
river was in pool behind each of the 
navigation dams. Not all stations coulé 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RivER 319 


be fished every year, because of high 
water levels, and no stations were fished 
in 1971 and 1972 due to high water. 
In addition, the Des Plaines River sta- 
tion was fished only in 1959, 1962, 
1973, and 1974, because it is not part 
of the Illinois River proper and was 
omitted whenever there was a limited 
amount of time available for sampling. 


Several physical-chemical measure- 
ments were made at each station be- 
fore sampling of the fish populations 
began. Dissolved oxygen concentra- 
tions at a depth of .91 m and at the 
bottom in the deepest part of the sta- 
tion were measured by the Winkler 
azide method and, in 1974, with a YSI 
Model 57 dissolved oxygen meter. Sur- 
face water and air temperatures were 
measured with a mercury thermometer. 
Wind direction and velocity and cloud 
cover were noted. Transparency was 
measured with a Secchi disk. In addi- 
tion, turbidity of the river was mea- 
sured with a Jackson turbidimeter dur- 
ing some surveys. 


Fish populations were sampled by 
means of electrofishing. Fish were 
stunned by an electric current pro- 
duced by a 230-volt, 180 cycles/sec, AC 
generator (Homelite QHY-1), and 
transmitted through the water via three 
cables suspended from booms in the 
front of a 5.49-m aluminum boat. The 
stunned fish were dipped from the 
water and placed in plastic garbage 
cans containing water. Electrofishing 
was conducted in 15-minute segments, 
and a total of 60 minutes was spent 
electrofishing at most stations. In small 
chutes, or where an abundance of fish 
was collected quickly, only 30 minutes 
were spent electrofishing. Fish were 
identified, counted, weighed, checked 
for disease, and returned to the river. 
The few fish that died were buried on 
shore. 


RESULTS 


PHYSICAL-CHEMICAL RESULTS 


Physical-chemical results for the fall 
of 1974 are shown in Table 2. Since 


the dissolved oxygen levels at both 
the .9l-m depth and on the bottom 
were approximately the same at every 
station, the water was presumably well 
mixed. The dissolved oxygen concen- 
tration was 77-97 percent of saturation 
in the Alton Pool, 65-122 percent of 
saturation in La Grange and Peoria 
Pools, and 47-104 percent of saturation 
in Starved Rock, Marseilles, and Dres- 
den Pools. At Ballard Island Chute 
(mile 247.8-248.2) and in Lower Peoria 
Lake (mile 163.0-163.4), the atypically 
high oxygen values (greater than satu- 
ration) were probably due to algal 
photosynthesis, since the waters had a 
greenish or brownish tinge. Ballard 
Island Chute is shallow, and has a 
large surface area, slow current, and 
a very dissected shoreline, with many 
marshy blind pockets. Thus, it should 
be a likely spot for phytoplankton to 
develop. The Secchi disk visibility here 
was much lower than in the river, al- 
though some of the turbidity on the 
sampling date can be attributed to 
wave action on the shallow bottom, 
as well as to phytoplankton. 


The upper river in 1974 was gen- 
erally more transparent, as measured 
by the Secchi disk, than the lower 
river. Starrett (1971:273) found that 
turbidity readings with a Jackson tur- 
bidimeter were higher in the lower 
three pools than in the upper three 
pools in the period 1963-1966. The 
Alton, La Grange, and Peoria Pools are 
generally more turbid than the upper 
pools, presumably because the lower 
pools have soft mud bottoms and re- 
ceive heavy silt loads from tributary 
streams that drain agricultural areas. 
The river above Hennepin (mile 207.5) 
generally has a rocky bottom, although 
the rock is overlaid with mud, sand, 
and/or gravel in some sections. 


Towboats (several barges pushed by 
a diesel-powered boat) have a marked 
effect on turbidity in the Illinois River. 
Fig. 1 shows that the turbidity in mid- 
channel at mile 25.9 was increased by 
about 100 Jackson turbidimeter units 
(JTU) as towboats passed on three 


320 Intinois NaturaAL History SurvEY BULLETIN Vol. 31, Art. 8 
180 
70 e) 
7 NOVEMBER 1963 170 


MILE 25.9 
MID-CHANNEL 


160 
a 
150 = 
~~ = 
S wo 
z fn 
= 30 8 
= = 
w 5 
Be 120i 
: 5 

= 

10 & 
PF 100 > 
a 
90 
> 

80 

70 

900 1000 1100 1200 1300 1400 1500 
CENTRAL STANDARD TIME 

OXYGEN = O——O SURFACE A—A MID-DEPTH [BOTTOM } BARGE PASSAGE @—@TURBIDITY 
Fig. 1.—Dissolved oxygen concentrations and turbidity in the middle of the navigation 


channel of the Illinois River at mile 25.9, during passages of towboats on 7 November, 1963. 
Symbols for dissolved oxygen are circles for at the surface, triangles for at mid-depth, and 
squares for at the bottom. Turbidity is indicated with black dots. The time at which each 
towboat passed mile 25.9 is marked by an arrow. 


occasions on 7 November, 1963. It took 
approximately 2% hours for the tur- 
bidity to return to background levels 
following passage of towboats. 

A Natural History Survey crew took 
a few dissolved oxygen readings in 
midchannel on 6 and 7 November, 
1963, before, during, and after tow- 
boats had passed (Fig. 1 and 2). One 
might expect turbulence from move- 
ment of the hulls and from the pro- 
pellers to aerate the water. Surpris- 
ingly, oxygen levels at the surface de- 
clined and then recovered following 
passage of a towboat on 6 November. 
On 7 November, oxygen levels at both 
the surface and bottom declined. The 
declines are significant; oxygen levels 
at the surface on 6 November and at 
the bottom on 7 November declined 
by 0.4 mg/l, and the standard deviation 


of the method used (azide modification 
of the Winkler method) is 0.1 mg/l, 
even in the presence of appreciable 
interference. The decline in dissolved 
oxygen and the increase in turbidity are 
both attributable to the resuspension 
of sediment caused by towboats moy- 
ing in the relatively shallow naviga- 


tion channel (2.74 m deep). Sediments 


in the Illinois River exert an appreciable 
oxygen demand, and the demand in- © 


creases 7-fold to 10-fold when the sedi- 


ments are disturbed. For example, — 
Butts (1974:12) reported an oxygen 

demand of 2.8 g/m?/day for sediment ~ 
at mile 198.8, under quiescent condi- 
tions, and 20.7 g/m’/day when the 
sediments were disturbed. The dis- — 
turbance was produced by water cur- 
rent within a special chamber which 


Butts had constructed to measure in ~ 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIs RIVER 321 


VATOCN \ 4 SALURATLUN 7 


1300 1400 1500 


CENTRAL STANDARD TIME 


1600 


6 NOVEMBER 1963 
MILE 25.9 
MID-CHANNEL 


OXYGEN 


O———O SURFACE 
A MID-DEPTH 
O—O ,BotTtom 


¥ BARGE PASSAGE 
@—® TuRBIDITY 


TURBIDITY (JACKSON TURBIDIMETER UNITS) 


1700 


Fig. 2.—The effects of towboat passage on dissolved oxygen concentrations and turbidity 
it the same location on 6 November, 1963. The symbols are the same as in Fig. 1. 


itu oxygen demand, and could logi- 
sally be equated to the effects of dis- 
urbances created by barges (Butts 
1974:6). 

It is not known why oxygen levels 
nereased slightly at the bottom on 6 
November, and at mid-depth on 7 
November, following passage of tow- 
oats. It may be that turbulence from 
owboats results in uneven mixing of 
yarcels of water aerated by turbulence 
with parcels of water deoxygenated by 
esuspended sediment. 


The water temperatures in Starved 
Rock, Marseilles, and Dresden Pools 
were generally higher than in the upper 
yart of Peoria Pool, even though the 
eadings in the upper pools were taken 
2 weeks later and the weather had 
urned colder. The upper river is evi- 
ently warmer because of warm in- 
ustrial and municipal discharges. Star- 
ett (1971:370-373) reported the same 


trend of warmer temperatures in the 
upper river in July and August, 1966. 


ELECTROFISHING RESULTS 


The electrofishing results for those 
species that were frequently taken are 
presented below in phylogenetic order. 


Shortnose Gar (Table 3) 

Table 3 shows that shortnose gar 
(Lepisosteus platostomus) were occa- 
sionally taken in the three downstream 
pools, but never taken in the three up- 
stream pools. Judging by the reports 
of commercial fishermen, shortnose gar 
are more abundant in the downstream 
pools than our records indicate, and 
these fish are probably less vulnerable 
to electric shock than other species. 
Although garfish are listed in the com- 
mercial catch from the Illinois River 
(Table 28) most fishermen consider 
them a nuisance because they easily be- 
come entangled in nets, with their 


322 


elongate snout and numerous sharp 
teeth, and there is little demand for 
them as a food fish. 


Bowfin (Table 4) 

Bowfin (Amia calva) is considered 
a commercial species, but was not com- 
mon in the Illinois River collections. 
Bowfin were taken as far upstream as 
Peoria Pool only in 1961, and other- 
wise were restricted to collections from 
La Grange and Alton Pools. Bowfin 
taken from Alton Pool in 1974 were 
in breeding color. 


Gizzard Shad (Table 5) 

Gizzard shad (Dorosoma cepedia- 
num) were most abundant in La 
Grange and Peoria Pools and were gen- 
erally abundant in our collections in all 
pools of the river. The numbers and 
pounds reported in Table 5 do not 
begin to reflect the actual abundance 
of the species, for two reasons. One 
is that small gizzard shad are stunned 
only momentarily by the electric shock, 
and usually get away before they can 
be netted. The second is that so many 
gizzard shad usually appear that it is 
futile to try to net them all, and our 
netting efforts are concentrated on the 
other species. 


Gizzard shad are neither a commer- 
cial nor a game species, but small shad 
are valuable forage for largemouth bass, 
crappies, and even species such as 
drum, which ordinarily prefer molluscs 
when they are available. 


Shad are sensitive to low oxygen and 
probably sensitive to cold temperatures, 
and die-offs of gizzard shad sometimes 
occur in the bottomland lakes and back- 
waters in midsummer and usually oc- 
cur in winter. Nevertheless, because 
of their high reproductive capacity, 
gizzard shad populations do not seem 
to be much affected by these die-offs. 


Goldeye, Mooneye (Tables 6 and 7 


— Discussed under ‘‘Species 
Infrequently Taken’”’) 


ILLinois NATuRAL History SuRVEY BULLETIN 


q 
Vol. 31, Art. 8: 


q 
* 


Goldfish, Carp x Goldfish Hybrids 
(Tables 8 and 9) 


Goldfish (Carassius auratus) wet 
probably introduced into the Illinois 
River between 1908 and 1935; Forbes: 
& Richardson do not mention them in 
The Fishes of Illinois (1920) and; 
O'Donnell (1935) mentions that they 
occur frequently in the Illinois River. 
O'Donnell (1935) also mentions that. 
two carp x goldfish hybrids were taken 
at Peoria. t 

Goldfish and carp x goldfish hybrids 
were generally abundant in the Des’ 
Plaines and upper Illinois electrofishing ; 
collections from 1959 through 1974° 
(Fig. 3 and 4). Goldfish were usually 
most abundant in Dresden, Marseilles, \ 
and Starved Rock Pools, and carp xX! 
goldfish were most abundant in Peoria « 
and Starved Rock Pools from 1964: 
through 1974. 

The catch of goldfish generally de- 
clined in the downstream direction, ) 
From 1959 through 1974 no goldfish! 
were taken in the Alton Pool, although! 


| 


1974 declined dramatically. The carp 
x goldfish hybrids did not exhibit such 
a dramatic decline. Hybrids may occu 
in the polluted upper river becaus€ 
“hybrid vigor” confers some resistance | 
to pollution, or simply because both’ 
carp and goldfish occur there. 2 


Carp (Table 10) | 
Carp (Cyprinus carpio) were intro- 


duced into the Illinois River in 1885. 
By 1898, carp brought more money t¢ 


commercial fishermen along the Illinoi 


(2,720,000-3,630,000 kg) per year and: 
was worth more than $200,000 ( Forbes 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 323 


Aug., 1975 Sparks & STARRETT 


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Vol. 31, Art. 8° 


Ituino1s NaturAL History SURVEY BULLETIN 


324 


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(1928:285). At present, carp and giz- 
zard shad are the only species that 
occur abundantly in our electrofishing 
collections in all pools of the river 
(Fig. 5). Carp and bigmouth buffalo 
comprise the bulk of the commercial 
catch in the Illinois River. The carp 
catch from the Illinois River was 213, 
000 pounds (104,000 kg) in 1973. 


River Carpsucker, Quillback 
Carpsucker (Tables 11 and 12) 

The greatest number of quillback 
carpsuckers (Carpiodes cyprinus) was 
usually taken in three pools of the Illi- 
nois River: Marseilles, Starved Rock, 
and Peoria. 

In contrast to the quillback, the most 
river carpsuckers (Carpiodes carpio) 
were generally taken in the three lower 
pools, Alton, La Grange, and Peoria, 
prior to 1973. In 1973 and 1974 most 
were taken in Starved Rock Pool, so 
their distribution in the river may have 
changed after the high-water period 
1971-1973. The quillback and river 
carpsuckers are both commercial spe- 
cies. 

Smallmouth Buffalo (Table 13) 

The largest numbers of smallmouth 
buffalo (Ictiobus bubalus) were taken 
from Peoria and La Grange Pools. An 
unusually large number of smallmouth 
buffalo were taken from Starved Rock 
Pool in 1974. The smallmouth buffalo 
is a commercial species. 

Bigmouth Buffalo (Table 14) 

Like the smallmouth buffalo, the big- 
mouth buffalo (Ictiobus cyprinellus), 
was most commonly taken in Peoria 
and La Grange Pools. Prior to 1974 
no bigmouth buffalo had been taken 
from Dresden and Marseilles Pools, and 
bigmouth buffalo had been taken in 
Starved Rock Pool in only one year, 
1966. In 1974 they were taken in both 
Starved Rock and Marseilles Pools. It 
is surprising that few buffalo were 
ever taken in Alton Pool, and that no 
buffalo were taken there in 1974. Sev- 
eral commercial fishermen at Kamps- 
ville Landing and Godar Landing on 


Inyinois NATuRAL History SURVEY BULLETIN 


Vol. 31, Art. 8 


the Alton Pool said that they also were 
catching very few bigmouth buffalo in 
1974. Bigmouth buffalo rank «ea 
to carp in the commercial catch from 
the Illinois River. % 


Black Buffalo (Table 15) ; 

The black buffalo (Ictiobus niger) is 
a commercial species. It was not abun- 
dant in the Illinois River electrofishing 
collections, and was taken only in th 
lower three pools prior to 1974. It was 
most commonly taken in Peoria rd 
La Grange Pools. In 1974, the few 
black buffalo taken all came from 
Starved Rock Pool. 


Shorthead Redhorse (Table 16 — 

Discussed under “Species : 

Infrequently Taken’) 
§ 


Black Bullhead (Table 17) 

The black bullhead (Ictalurus melas 
is considered a commercial specie’ 
but most of the bullheads in our elec 
trofishing collections were quite small, 

Most of the black bullheads we 
taken from one station, Ballard Islan 
Chute (river mile 247,8-248.2) 
Marseilles Pool (Fig. 6), which wat 
described earlier as being an uno 
shallow, broad, marsh-fringed area, with 
very little current. The black bullhea 
probably prefers this type of habitat. 

Black bullheads were collected oce 
sionally in the main navigation channel 
by means of an otter trawl. For e 
ample, on 26 August, 1964, 51 bla 
bullheads averaging 18 cm in tot 
length were taken in 49 minutes 
trawling at mile 193. 


Yellow Bullhead (Table 18 — 
Discussed under ‘“‘Species 
Infrequently Taken’) 


Channel Catfish (Table 19) 

Channel catfish (Ictalurus punct 
tus) were taken in Marseilles Pool fot 
the first time in 1974. Also, the sec 
ond largest number and weight of fi 
were taken in the river in 197 
(Fig. 7). Most channel catfish wer 
taken below Beardstown (river mil 


327 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 


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Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIvER 329 


88.5). They were taken occasionally 
from the main navigation channel by 
trawling. On 13 November, 1964, 68 
young channel catfish averaging 9 cm 
in total length were taken in 53 minutes 
of trawling in the channel at mile 156. 
Prior to 1973, the numbers and weights 
of channel catfish taken appear to be 
unrelated to water levels. Channel cat- 
fish have declined in the Illinois River 
since 1899 as evidenced by the follow- 
ing commercial fishing statistics: 241, 
000 pounds (109,316 kg) in 1899, 105, 
304 pounds (47,878 kg) in 1950, about 
98,000 pounds (44,452 kg) in 1964 
(Mills, Starrett, & Bellrose 1966:17), 
and 45,000 pounds (20,412 kg) in 1973. 
(Larry Dunham, Fishery Biologist, Illi- 
nois State Department of Conservation, 
personal communication. ) 


Flathead Catfish (Table 20) 

Flathead catfish (Pylodictis olivaris ) 
are a desirable commercial species and 
often reach weights of 9-18 kg. Flat- 
head catfish were never abundant in 
the electrofishing collections, and were 
confined to the lower two pools. An 
§.16-kg individual was taken in La 
Grange Pool and several 1- or 2-year- 
old flatheads were taken at several 
stations in both Alton and La Grange 
Pools in 1974. 


White Bass (Table 21) 

The white bass (Morone chrysops) 
is a game species. The largest number 
of white bass was taken from the river 
in 1974, but the greatest catch by 
weight was in 1968. White bass popu- 
ations generally increased in the down- 
stream direction, with the largest num- 

er and greatest weights usually taken 
in Alton Pool. 


reen Sunfish (Table 22) 

Green sunfish (Lepomis cyanellus) 
re considered game fish by some 
eople, although they do not grow as 

rge as their relative, the bluegill. 
he green sunfish was taken in the Des 
laines River in two of the four years 
this station was sampled, whereas the 
bluegill was never taken from this sta- 


| 


tion. The largest numbers of green 
sunfish were generally taken in Peoria 
Pool. The number of green sunfish 
taken did not increase dramatically 
after the high-water period 1971-1973, 
as did the number of bluegills. 


Bluegill (Table 23) 

The largest number and greatest 
weight of bluegills (Lepomis macro- 
chirus) per 30 minutes of electrofishing 
were taken in 1974. Bluegill popula- 
tions generally increase in the down- 
stream direction, with either Alton or 
La Grange Pools having the greatest 
number and weight. However, in only 
one year, 1969, were more bluegills 
obtained in Starved Rock Pool than in 
the next pool upstream. 


Largemouth Bass (Table 24) 

The largemouth bass (Micropterus 
salmoides) is a game species. Large- 
mouth populations generally increase 
in the river in a downstream direc- 
tion (Fig. 8), with the greatest num- 
bers taken from La Grange and Peoria 
Pools. However, fewer bass were taken 
at the two stations in Starved Rock Pool 
than at the three stations in the next 
pool upstream, Marseilles. Bass popu- 
lations in the river as a whole reached 
their peak in 1960 and 1961, then 
showed a drastic decline during and 
following the drought years 1962-1964. 
The recent increase in largemouth pop- 
ulations follows the high-water years 
1971-1973. 


Crappies (Tables 25 and 26) 

The largest catch of both black 
crappie (Pomoxis nigromaculatus ) and 
white crappie (Pomoxis annularis), in 
weight and numbers, was taken in the 
river in 1974, following the high-water 
years 1971-1973. Populations of both 
species showed a steady decline in the 
years 1962-1965, during a drought pe- 
riod. Prior to 1973, few crappies were 
taken in the upper three navigation 
pools, but increased numbers of both 
species were taken in the Starved Rock 
and Marseilles Pools in 1974. In 1962, 


Vol. 31, Art. 8: 


Intino1is NATuRAL History SURVEY BULLETIN 


330 


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\ug., 1975 Sparks & STARRETT: ELEcTROFISHING SuRveY OF ILLinois River 331 


964, 1966-1969, and 1974, more black 
rappie were taken in La Grange Pool 
han in Alton Pool, perhaps because 
nore backwater and side channel areas 
vith brush piles (a favorite habitat of 
rappie) were usually available in La 
Srange Pool. In 1974 a larger number 
yf small white crappie was taken in 
La Grange Pool than in Alton Pool but 
1 greater weight of large white crappie 
was taken in Alton Pool. Both species 
we popular game fish. 


Freshwater Drum (Table 27) 
Freshwater drum (Aplodinotus grun- 
niens) is a commercial species. Most 
were taken in La Grange Pool. The 
largest number of individuals and the 
second greatest weight were taken in 
1974, following a high-water period. 


Species Infrequently Taken 

The yellow bullhead (Ictalurus na- 
talis) (Table 18) was uncommon in 
our collections, and has been taken 
only from the three lower pools, Alton, 
La Grange, and Peoria. 

The shorthead redhorse (Moxostoma 
macrolepidotum) (Table 16) occurred 
sporadically in our collections through- 
out the river. 

A female spotted gar (Lepisosteus 

oculatus) was taken by a commercial 
fisherman at Havana on 26 February, 
1973. Spotted gar are uncommon in the 
Illinois River. This specimen was the 
largest that had been taken in Illinois 
(3.41 kg, 83.8 cm in total length) and 
was full of ripe eggs. 
Mooneye (Hiodon tergisus) (Table 
7) were taken rarely, and only from 
e Alton Pool until 1974, when one 
as taken from upper Peoria Pool at 
mile 215. Goldeye (Hiodon alosoides ) 
(Table 6) were taken rarely, but 
ranged farther upstream than their 
relative, the mooneye. In 1974 only two 
‘goldeye were taken, both from one sta- 
tion at mile 261 in Marseilles Pool. 


The American eel (Anguilla rostrata ) 
as rarely taken. One was taken from 


—_— 


Alton Pool at mile 19 and two from 
Peoria Lake in 1974. 

The white catfish (Ictalurus catus) 
is a native of brackish to fresh waters 
along the East Coast from Pennsyl- 
vania to Florida. It has been intro- 
duced widely in the Midwest, and sev- 
eral have been taken from the Illinois 
River by commercial fishermen at Ha- 
vana, including one on 13 May, 1974. 
White catfish have never been taken 
in our electrofishing surveys. 

The few smallmouth bass (Microp- 
terus dolomieui) that were taken were 
probably introduced from tributary 
streams that are smaller and colder 
than the Illinois River. 

Skipjack herring (Alosa chrysochlor- 
is) were taken sporadically throughout 
the Illinois River. Large numbers ap- 
parently moved up the river during 
the spring flood of 1973, and sport fish- 
ermen were catching them on minnows 
at Havana. 

One sauger (Stizostedion canadense ) 
was taken at Big Blue Island Chute 
(river mile 57.5-58.9) in 1974. This 
species was common in the river before 
1908 (Forbes & Richardson 1920:275). 

Orange-spotted sunfish (Lepomis 
humilis) and pumpkinseeds (Lepomis 
gibbosus) were taken sporadically. 

One species, the longear sunfish 
(Lepomis megalotis), listed as being 
extirpated from the Illinois River and 
its bottomland lakes between 1908 and 
1970, by W. C. Starrett and P. W. 
Smith (Starrett 1972:163), was taken 
from La Grange Pool, Turkey Island 
Chute (mile 147.3-148.2) on 5 Sep- 
tember, 1973. Three adults, ranging in 
total length from 10.7 to 15.5 cm were 
taken. 

Northern pike (Esox lucius) were 
taken by sport fishermen in the river 
below Marseilles Dam in 1973, and 
were netted in Lake Chautauqua in 
1973 (river mile 126.0), but were not 
taken by electrofishing. Northern pike 
were common in the river before 1908 
(Forbes & Richardson 1920:209 ). 


332 Iuyinois NaturAL History SuRvEY BULLETIN Vol. 31, Art. 8 


Catfishes may be more abundant in 
the river than our collections indicate. 
They are bottom-dwelling species and 
when shocked they do not always come 
to the surface where they can be seen 
to be netted. Under nearly ideal con- 
ditions for electrofishing, Larimore 
(1961) reported taking only 10 percent 
of the total population of catfishes in a 
reach of Jordan Creek, whereas 52 per- 
cent of the sunfishes were taken. In 
the generally turbid waters of the lower 
Illinois River, a fish must be within 
10-15 cm of the surface to be seen. 
So our collecting efficiency for catfishes 
must have been lower than the 10 per- 
cent obtained by Larimore in clear 
water. 

Since we used a shocker, and 6.35 
mm mesh dip nets, minnows and other 
small fishes were generally not taken. 
We did obtain emerald shiners (No- 
tropis atherinoides) throughout the 
river in 1974 and in previous years 
(Mills, Starrett, & Bellrose 1966:15). 


DISCUSSION 
HISTORICAL CHANGES 
IN THE FISH POPULATIONS 
OF THE ILLINOIS RIVER 

The Illinois-Michigan Canal along the 
upper Illinois River was completed in 
1848, before any biological data were 
being collected on the Illinois River. 
Prior to 1871, it is unlikely that this 
canal had much of an impact on the 
middle and lower sections of the river, 
below Hennepin (river mile 208), 
which are the sections most productive 
of fish and wildlife. These are the 
most productive because the Illinois 
River below Hennepin follows a large 
valley developed in the late Pleistocene 
epoch, and the Illinois has developed 
lateral levee lakes, side channels, back- 
waters, and marshes which fill this an- 
cient valley and provide excellent habi- 
tat for fish and wildlife. 

In 1871, the flow of the Chicago 
River was reversed in order to conduct 
sanitary wastes from the city of Chi- 
cago away from Lake Michigan, which 


a 


served as the drinking water supply\ 
for the city. The polluted waters of; 
the Chicago River were directed 
through the Illinois-Michigan Canal) 
into the Des Plaines River and thence; 
into the Illinois River. Some of the 
polluted water apparently backed up; 
into the lower reaches of the Kankakee, » 
The effect of the polluted water on the: 
fishes of the Kankakee and _ Illinois) 
rivers was dramatic, according to a re-’ 
port by Nelson (1878:798): 
“Previously to the opening of the) 
Chicago River into the canal in 1871.) 
rock-bass, (Ambloplites rupestris);) 
black-bass, (Micropterus pallidus) 
[largemouth bass, Micropterus sal-| 
moides]; silver bass, (Roccus chry-) 
sops) [white bass, Morone chrysops];) 
wall-eyed pike, (Stizostethium vit-t 
reum) [walleye, Stizostedion vitreum) 
vitreum]; mud-pike, (?); pickerel,! 
(Esoxlucius) [northern pike, Esox) 
lucius]; mud-eel, (?) [lamprey?];) 


ican eel]; buffalo fish, (Bubalichthys 
bubalus) [buffalo, Ictiobus 
red horse, (Myxostoma macrolepi- 
dota) [shorthead redhorse, Moxosto-) 
ma macrolepidotum]; suckers, Catos-) 
tomus ?); bull-heads, ( Amiurus! 
catus) [bullhead, Ictalurus Py; 
spoon-fish, or shovel-bill, (Polyodon) 
folium) [paddlefish, Polyodon spath-\ 
ula]; sun-fish, (Pomotis ?) [sun- 
fishes, Lepomis 
(Amiurus 
____?]; dog-fish, (Amia calva)) 
[bowfin]; gar pike, (Lepidosteus; 


flows 6 miles east of Joliet, and emp 
ties into the Desplaines 8 miles south! 
of that town; also in Hickory Creek 
which rises about 14 miles east 0 
Joliet, and empties into the Des~ 
plaines just south of the town, and) 
in any of the streams of sufficien 
size in this vicinity. 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIVER 333 


“When the current of Chicago 
River was first turned through the 
canal and the rivers, it caused the 
fish in them to bloat to a large size, 
and rising to the surface they floated 
down the stream in large numbers. 
It was estimated at the time that 
several tons of dead fish passed 
through one of the canal locks just 
after the foul water commenced run- 
ning through the canal. 

“When these bloated fish chanced 
to float into the clear water at the 
mouth of some tributary of the river 
they would revive and swim up the 
clear stream. Such large numbers 
of the fish revived in this manner that 
all the small streams flowing into 
the Desplaines and Kankaku [sic] 
rivers were filled with fish in such 
numbers that many were taken with 
hook and line, one man taking over 
300 in a day in this manner at that 
time. 

“When the spring freshets occur 
the current is so rapid and the 
amount of pure water in the river 
is so great, that the foul water does 
not have much effect upon the fishes, 
and large numbers of the species 
mentioned ascend the rivers and are 
caught with hook and line. Later in 
the season as the water subsides, and 
the water from Chicago River pre- 
dominates, the fish which came up 
in the spring die and are floated 
down the river. In July and August 
when the water is the worst even the 
mud turtles leave the river in dis- 
gust and seek less odorous homes.” 


Water from the [Illinois-Michigan 
Canal also entered the Illinois River 
at La Salle (mile 223), but the wastes 
were sufficiently decomposed at that 
point that there was only a slight im- 
pact on the ecosystem of the Illinois 
River below La Salle (Starrett 1972: 
145). 


The carp was introduced into the IIli- 
nois River in 1885, out of a_ stock 
brought to the United States a few 


years earlier from Europe (Forbes & 
Richardson 1920: 105). By 1898, the carp 
catch exceeded the value of all other 
commercial fishes from the Illinois River 
(Thompson 1928:285). Forbes & Rich- 
ardson (1920:108-109) reported fishery 
statistics which showed that increasing 
carp populations did not adversely af- 
fect the populations of other species, 
although they did predict that carp 
might displace the native buffalo fishes, 
which have the same food preferences 
as carp. Forbes & Richardson (1920: 
108-110) did not feel that carp had in- 
creased the turbidity of the water in the 
Illinois River by their rooting habit of 
bottom feeding. In contrast, Jackson 
& Starrett (1959:163-165) observed lo- 
cal areas of heavy turbidity in Lake 
Chautauqua, a bottomland lake along 
the middle section of the river, pro- 
duced by schools of carp. They felt that 
some instances of carp activity may 
have been stimulated by low oxygen 
levels. The activities of carp may have 
had a greater effect on turbidity in 
more recent times because of the pres- 
ence of flocculent bottom muds that 
have been carried into the bottomland 
lakes by the river (Starrett & Fritz 
1965:88 ). 


Forbes (1928) does not mention any 
changes in fish fauna associated with 
the construction, prior to 1900, of the 
low navigation dams on the Illinois 
River at Marseilles, Henry, Copperas 
Creek, La Grange, and Kampsville. 
Nelson (1878:798) was of the opinion 
that a dam at Seneca (mile 252.5) 
hindered the upstream movement of 
fishes. On 1 January, 1900, the Sani- 
tary and Ship Canal was opened at 
Chicago, connecting the Des Plaines 
and Illinois Rivers with Lake Michi- 
gan. The canal was used to flush mu- 
nicipal and industrial wastes into the 
Illinois River system, and away from 
Chicago’s municipal water intakes in 
Lake Michigan. The quantity and 
quality of this diverted water had a 
tremendous impact on the Illinois 


334 


River. There was an average rise in 
water levels at Havana of 2.8 feet 
(.85 m), and during the normal low- 
flow period between June and Sep- 
tember the rise was 3.6 feet (1.10 
m) (Forbes & Richardson 1919:140- 
141), The tree line along the river re- 
treated as a result, and the loss of 
mature pin oak (Quercus palustris ) and 
pecan (Carya illinoensis) trees meant 
a loss of food for mallard ducks (Anus 
platyrhynchos) and wood ducks (Aix 
sponsa) (Mills, Starrett, & Bellrose 
1966:5). Populations of cavity-nesting 
tree swallows (Iridoprocne bicolor) 
and prothonotary warblers (Protono- 
taria citrea) increased, as a result of 
the increased supply of nest sites in 
zones of dead trees bordering the river 
and lakes. Populations of these species 
declined markedly during the 1940’s, 
as the last of the dead trees finally 
collapsed (Dr. Frank C. Bellrose, 
Waterfowl Biologist, Illinois Natural 
History Survey, personal communica- 
tion). 


One beneficial effect of the diver- 
sion was to increase the surface area 
of water in lakes and backwaters, 
which apparently improved the fishery 
(Forbes & Richardson 1919). It is also 
likely that the stumps and snags left 
after the trees had died temporarily 
provided cover for certain species such 
as bass, crappie, and other sunfishes. 
The increased shallow water areas and 
nutrient loading of the Illinois River 
and its bottomland lakes initially may 
have increased the plankton popula- 
tions and the biomass of bottom fauna 
in the middle and lower river (Forbes 
& Richardson 1913:494495). In the 
river proper, populations of molluscs, 
especially fingernail clams, probably 
increased the most, with a beneficial 
effect on mollusc-consuming species of 
adult fish such as carp, catfish, buffalo, 
and drum. 


After approximately 1910, however, 
as the pollution load increased, criti- 
cally low dissolved oxygen levels oc- 


Ittivois NaTuRAL History SURVEY BULLETIN 


Vol. 31, Art. 8 — 


curred farther and farther downstream 
with detrimental effects on food or- 
ganisms and fish (Richardson 1921b;— 
33). Populations of molluscs, including — 
fingernail clams (Sphaeriidae), in the — 
middle section of the Illinois River and — 
in several bottomland lakes were quite — 
high in the early 1950’s (Paloumpis & — 
Starrett 1960). 


In 1938, by order of the Supreme 
Court of the United States, the amount 
of water that could be diverted from 
Lake Michigan at Chicago was limited” 
to a yearly average of 42.48 m*/sec 
and minimum gage readings in the — 
middle section of the river at Havana — 
dropped about .61 m as a result (Star- 
rett 1972:146). In spite of an increasing 
human population in the Illinois basin, 
the population equivalent of the total 
combined domestic and _ industrial 
waste emptied into the river declined 
from 6,211,471 in 1922 to 2,417,000 in 
1960 (Mills, Starrett, & Bellrose 1966: 
9), because more waste was receiving” 
primary and secondary waste treatment. 
Population equivalents are based om 
the average amount of carbonaceous” 
oxygen demand in the waste produced 
per person, and do not take into ac 
count the oxygen demand of the nit 
trogenous fraction of human waste: 
The demand placed on the oxygen re= 
sources of the river by nitrogenous 
wastes has actually increased in recent 
years (Butts 1975). 


Minimum dissolved oxygen levels 
near the surface in the channel of 
the Illinois River during midsummer 
in the period 1911-1966 are reported 
in tables in Mills, Starrett, & Bellrose 
(1966:9) and Starrett (1971:370-373) 
In 1966, oxygen levels generally were 
below saturation throughout the whole 
length of the river. Levels below 1.0 
mg/1 occurred in Dresden, Peoria, and 
La Grange Pools. The reduction im 
dissolved oxygen concentration so far 
downstream of the Chicago and Peoria 
metropolitan areas results from the 
oxygen demand of sediment (Butts 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RiveR 335 


1974) and from the oxygen demand 
as ammonia in municipal waste is con- 
verted to nitrate (Butts 1975). During 
the winter, bacterial nitrification is 
slowed, oxygen demand is thereby re- 
duced, and higher ammonia concentra- 
tions extend farther downstream from 
Chicago (Butts 1975), 

Ammonia places aquatic organisms 
in double jeopardy; it not only removes 
oxygen from water, but is also toxic. 
Only the un-ionized fraction of the total 
ammonia concentration (approximately 
5 percent of the total ammonia in the 
Illinois River) is toxic, and the un-ion- 
ized ammonia concentrations were gen- 
erally well below lethal levels for fish 
in 1972 and 1973, although concentra- 
tions may have been high enough on 
occasion in the upper river to stress 
fish (Lubinski et al. 1974). 

It is not known to what extent the 
low dissolved oxygen concentrations, 
perhaps acting in combination with 
other stresses such as silt and toxic 
materials, contributed to the die-off of 
fingernail clams and snails in the mid- 
dle section of the river in the mid- 
1950's ( Mills, Starrett, & Bellrose 1966: 
12). As late as 1973, fingernail clams 
had not reappeared in areas of the river 
where dead shells indicated that they 
were formerly abundant. The loss of 
these important food organisms, ac- 
cording to the Mills, Starrett, and Bell- 
rose report, has resulted in a reduction 
of the number of diving ducks migrat- 
ing along the Illinois River and a de- 
cline in the condition factor of the com- 
mercially valuable carp. 


In addition to affecting the food sup- 
ply of fish, low oxygen levels have di- 
rect effects on fish. Carlson & Siefert 
(1974) have shown that oxygen levels 
at 35 percent saturation reduced the 

survival of larval largemouth bass by 
13.7 percent, and oxygen levels at 70 
percent saturation and below retarded 
the growth of larval bass. In two areas 
that provide good physical conditions 
for largemouth bass, Lower Bath Chute, 


La Grange Pool (Fig. 9) and Chilli- 
cothe Island Chute, Peoria Pool (Fig. 
10), midsummer oxygen levels were at 
35 percent saturation or below for 
4-5 years out of the 8-year period 
1963-1970. 

The discharge and water levels were 
generally high preceding the resurgence 
in bass populations at Lower Bath Chute 
(Fig. 9). Therefore, it is difficult to 
separate the beneficial effects of high 
water levels from the beneficial effects 
of increased discharge. During high 
water, flooded areas provide good 
breeding habitat for many adult fish 
and good nursery areas for juvenile 
fish. High discharge results in in- 
creased dilution of toxic wastes and 
oxygen-demanding wastes. At Chilli- 
cothe Island Chute (Fig. 10) the rela- 
tive importance of the two effects can 
be separated, because the water levels 
in Peoria Pool were maintained within 
fairly narrow limits by flow regulation 
at the Peoria Lock and Dam, while the 
discharge varied considerably. 

The resurgence in bass populations 
at Chillicothe Island Chute was as- 
sociated with increased discharge. Al- 
though we took no oxygen readings in 
the chute during midsummer 1973 or 
1974, oxygen readings in other parts of 
the river were generally 80 percent of 
saturation, and indicate that oxygen- 
demanding wastes were being diluted. 
Toxic wastes probably were diluted 
during this period also. 


Lubinski et al. (1974) indicated that 
the combined toxicity of the chemicals 
routinely monitored by the Illinois En- 
vironmental Protection Agency was 
generally well below levels lethal to 
fish at 17 locations on the Illinois 
River during an 18-month period in 
1972 and 1973, when discharge was 
high. Extensive monitoring of toxic 
materials in the Illinois River has been 
undertaken only recently, so Lubinski 
et al. (1974) were not able to estimate 
the combined toxicity of chemicals to 
fish during low discharge. The real 


336 


test of whether pollution abatement 
programs in the Illinois Valley have re- 
sulted in improvement of water quality 
for fish will occur during low discharge 
periods in the years to come. 

One of the major impacts on the 


NO. OF LARGEMOUTH BASS TAKEN 
PER 30 MIN OF ELECTROFISHING 


T9 096T 


Die (Oe 69s el BO 29 90R MSC ee ed! Gu 


Gl 


HZ6T &/ 


£ 
oO 


WGN 
uw 


= 


JULY-AUG. DISSOLVED OXYGEN LEVELS 
(% SATURATION) 


oe #8 


Iuuinois NaTuRAL History SuRVEY BULLETIN 


( ZOT/FIIW ) JLNHD HIVE YSKOT 


Vol, 31, Art. 8 


Illinois River below Hennepin was 
the leveeing and draining of bottom- 
land areas, primarily in the period 
1903-1926. Of 400,000 bottomland 
acres (161,874 ha) subject to overflow 
by the river, approximately 200,000 


JULY-AUGUST DISCHARGE 
AT KINGSTON MINES (m3/sEc) 


NW ££ uu 
ouowmodwvw 
(a ee = es ee =) 


009 


SPN Venn aS 

ODN LEMNWOAON LM 
JULY-AUGUST WATER LEVELS 

AT HAVANA (M) 


>_> 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIVER 337 


acres (80,937 ha) are now behind 
levees (Mills, Starrett, & Bellrose 1966: 
5), with a consequent reduction in 
wildlife and fish habitat. The back- 
waters and bottomland lakes of the 
Illinois River were, and are, critically 
important to fish and wildlife produc- 
tion. 


Richardson (1921a:464) reported 
that the largest weights of fish per 
acre were taken in reaches of the river 
‘with the largest connecting lake area: 

“Taking the year 1908 as an illus- 
tration, and using the figures for 
separate shipping points obtained 
by the Illinois Fish Commission in 
that year, we find for the 59.3 miles 
of river and lakes between Copperas 

Creek dam (river mile 136.9) and 

La Grange dam (river mile 77.6), 

with about 90% of its acreage con- 

sisting of lakes and ponds, an average 
fish-yield per acre for water levels 

prevailing half the year, of 178.4 

pounds; for the 87 miles from La 

Salle (river mile 223.9) to Copperas 

Creek dam, with about 83% lakes, 

130.4 pounds; and for the lower 77 

miles, La Grange to Grafton, with 

around 63% lakes, only 69.8 pounds.” 


Richardson (1921a:463) indicates 
that well over 80 percent of the total 
fish yield in 1908 came from the lakes, 
with much less than 20 percent coming 
from the river itself. The bottomland 
lakes supported an abundant aquatic 
weed-inhabiting invertebrate fauna, 
which supplied food for young fishes 
of the sunfish, perch, and pike families. 


In the 1930’s high navigation dams 
were constructed at Dresden Heights 
(6.71 m high), Marseilles (7.32 m), 


Starved Rock (5.79 m), Peoria (3.35 
m), and La Grange (3.05 m). The 
navigation dam at Alton on the Missis- 
sippi raised water levels in the Illinois 
as far north as Hardin, at river mile 
21.0. Timber and brush were cleared 
from areas due to be inundated by the 
new dams. Clearing operations prob- 
ably did not markedly reduce the 
amount of mast available for water- 
fowl, according to Dr. Frank C. Bell- 
rose, Waterfowl Biologist, Illinois Natu- 
ral History Survey. The navigation 
dams temporarily increase dissolved 
oxygen levels as the water passes over 
and through the dams (Mills, Starrett, 
& Bellrose 1966:9-10; Forbes & Rich- 
ardson 1913:549). Starrett (1971:271- 
272) indicated that the reduction of 
diversion from Lake Michigan coupled 
with the higher dams on the river 
have resulted in a decrease of average 
current velocity from about 2.01-4.02 
km/hour prior to 1908 to 0.97 km/hour 
in 1966. Pools behind navigation dams 
on the upper river have filled with 
oxygen-demanding sediment which in 
places resembles sludge from secondary 
sewage treatment plants (Butts 1974). 

Richardson (1921a:457, 474-475) in- 
dicated that abundant populations of 
fingernail clams in the Illinois River 
were generally found in areas of re- 
duced current and favorable conditions 
for sedimentation. We (and _ others, 
such as Gale 1969) have found that 
abundant populations of fingernail 
clams occur in Pool 19 on the Missis- 
sippi River, over soft mud bottoms, 
and Gale (1971) reported that finger- 
nail clams will select mud substrates in 
preference to sandy mud and sand. 


Fig. 9.—The relationships among mean water levels (open triangles), mean discharge 
(black triangles), and mean dissolved oxygen levels (black dots) during the months of July 


and August and the number of largemouth bass taken per 30 minutes of electrofishing (circles) 
in the fall at Lower Bath Chute (mile 107) in the La Grange Pool. Oxygen levels below 35 
percent saturation (heavy line) reduce the survival of larval largemouth bass. Discharge was 
measured at Kingston Mines (mile 145), water levels at Havana (mile 120), and oxygen levels 
in the chute. The oxygen reading marked by an asterisk was taken on 12 September, rather 
than in midsummer. Discharge rates were obtained from Water Resources Data for Illinois, 
U.S. Dept. of the Interior, Geological Survey. Water levels were obtained from Missouri- 
Mississippi River Summary & Forecasts, U.S. Dept. of Commerce, National Oceanic and At- 
mospheric Administration, National Weather Service Central Region, Kansas City, Missouri. 
The other data were obtained by the Illinois Natural History Survey. 


She Che VEE O00 269%= “88h 79) M99. 299. ciS" eo =-296R 


HZ61 


338 


If the high navigation dams constructed 
in the 1930’s did reduce the current 
and increase sedimentation in parts of 
the Illinois River, then the habitat suit- 
able for fingernail clams may have 
increased, with a benefit to the mollusc- 


NO. OF LARGEMOUTH BASS TAKEN 
PER 30 MIN OF ELECTROFISHING 


ho 
S = 


Oa 
— 
= 
= 
I 
= 
ton) 
=) 
= 
=x 
m 
= 
= 
= 
=) 
(om) 
Se 
= 
= 
m 
-_ 
= 
= 
= 
m 
he 
(ee) 
=) 
~— 


nN 
oO 


JULY-AUG. DISSOLVED OXYGEN LEVELS 
(% SATURATION) 


nae 


WNW 
ul © 


In.tino1s NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 8 


eating fish. It is puzzling that condi- 
tions have been so dramatically dif- 
ferent since 1955, when a die-off of 
fingernail clams occurred in the middle 
section of the Illinois River (Mills, Star- 
rett, & Bellrose 1966:12). As late as 


JULY-AUGUST DISCHARGE 
AT MARSEILLES (M3/sEC) 


= = Pou oe 
= f—) ui So uw Oo 
JULY-AUGUST WATER LEVELS 

AT PEORIA \M 


—— Sea 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SuRVEY OF ILLINOIs River 339 


1973, the fingernail clams had not re- 
turned to areas of the river where dead 
shells indicated they had formerly been 
abundant. 


Starrett (1971:272) felt that the in- 
crease in sluggishness of the river and 
the increased planting of row crops in 
the Illinois basin have made siltation 
in the last 30 years an important factor 
adversely affecting the survival of mus- 
sels and other organisms in the Illinois 
River and its bottomland lakes. Silt 
physically removes habitat by filling 
in areas such as Lake Chautauqua, near 
Havana (river mile 124-130), which 
has lost 18.3 percent of its storage 
capacity in a period of 23.8 years (Stall 
& Melsted 1951:1). Areas in Quiver 
Lake near Havana where boats could 
formerly be launched are now only a 
few centimeters deep in low water 
stages, and willows are encroaching on 
the lake. Jackson & Starrett ( 1959:160) 
stated: 

“The sediments in Lake Chautau- 
qua are mostly of a fine texture and 
form a loose, flocculent ‘false bottom’ 
(not similar to the type found in 
bog lakes) over the original lake 
bottom. A slight disturbance of the 
‘false bottom’ causes particles to be- 
come resuspended and so increases 
the turbidity of the water.” 

The same authors found that an in- 
crease in wind velocity from light to 
strong increased the turbidity from 162 
to 700 Jackson turbidimeter units 
(JTU) and that it took a calm period 
of 7-12 days for much of this sediment 
to settle from Lake Chautauqua. As a 
consequence, this lake and other bot- 
tomland lakes are highly turbid most of 
the time. 


The turbidity levels in bottomland 


lakes and backwaters along the Illinois 
River are within the range that reduces 
fish production. Buck (1956) studied 
fish production in farm ponds, hatchery 
ponds, and reservoirs in Oklahoma 
which had a wide range of turbidities. 
The farm ponds were treated with 
rotenone, then restocked with large- 
mouth bass and bluegills or largemouth 
bass and redear sunfish (Lepomis 
microlophus ). Twelve farm ponds were 
divided into three turbidity classes. 
After two growing seasons, the average 
total weights of fish were: 
Clear ponds 
(less than 25 JTU)—161.5 lb/acre 
(181.0 kg/ha ) 
Intermediate ponds 
(25-100 JTU)—94.0 Ib/acre 
(105.4 kg/ha) 
Muddy ponds 
(>100 JTU )—29.3 Ib/acre 
(32.8 kg/ha ) 


The decline in production in turbid 
ponds resulted from a decline in both 
reproduction and growth (Buck 1956). 

The results from hatchery ponds, 
where turbidities were artificially con- 
trolled, and from the reservoirs which 
harbored a variety of fishes, generally 
paralleled the results from the farm 
ponds, except for two species, channel 
catfish and flathead catfish. 

Channel catfish spawn in dark cavi- 
ties, such as hollow logs or in holes 
in banks. Turbid waters are likely to 
have more suitably dark cavities per 
surface area or length of shoreline than 
do clear waters, and thus reproduction 
of channel catfish was probably greater 
in the turbid waters. Flathead catfish 
grow well in turbid waters and appear 
to be well adapted to turbid conditions. 


Fig. 10.—The relationships among mean water levels (symbols are the same as in Fig. 9), 
mean discharge, and mean dissolved oxygen levels during the months of July and August and 
the number of largemouth bass taken per 30 minutes of electrofishing in the fall at Chillicothe 


Island Chute (mile 180) on the Illinois River. 


Oxygen levels below 35 percent saturation 


(heavy line) reduce the survival of larval largemouth bass. Chillicothe Island Chute is in the 
Peoria Pool. Discharge was measured at Marseilles (mile 247), water levels at Peoria (mile 
163), and oxygen levels in the chute. The oxygen reading marked by an asterisk was taken 
on 30 September, rather than in midsummer, Data were obtained from the same sources as 


given for Fig. 9. 


340 


Buck (1956:257) concludes that in 
newly formed reservoirs bass, crappies, 
and other scaled fish out-produce cat- 
fish and then limit them by predation 
on the young. Turbid waters offer cat- 
fish protection from these predators. 
In addition, sunfishes prefer to con- 
struct nests on firm substrates, rather 
than mud. Their eggs and fry are prob- 
ably more susceptible to smothering 
by sediment than those of catfish and 
rough fish. 


The disappearance of the yellow 
perch (Perca flavescens) from the Illi- 
nois River and its bottomland lakes is 
probably also associated with the dis- 
appearance of the plant beds and clean 
sandy or pebbly bottoms the perch uses 
for spawning. 


Catfish feed on the types of food 
organisms which can grow in turbid 
waters with mud bottoms, such as 
midges, worms, fingernail clams, and 
snails. Catfish can use their highly 
developed sense of smell to locate food, 
whereas other game fish rely more 
heavily on sight. Food habits studies 
have shown that young game fish 
feed first on zooplankton, then on in- 
sects such as dragonfly and damselfly 
nymphs, then on larger organisms such 
as fishes and crayfishes. These types 
of food organisms are associated with 
weed beds and moderately clear water. 
The bottomland lakes along the Illinois 
River have been transformed from the 
latter type of ecosystem to a turbid type 
of system, by the influx of sediment 
from the river. 


Recently, even the fish and duck food 
organisms which are adapted to mud 
bottoms have died out in the channel 
and lateral areas of the middle section 
of the Illinois. Fingernail clams in this 
section died out in 1955, and have not 
since recolonized the area. It is pos- 
sible that some of the heavier benthic 
animals such as the molluscs find it 
difficult to remain near the top of the 
flocculent bottoms or that the sus- 
pended material interferes with their 


Intiwois Naturau History SuRVEY BULLETIN 


Vol. 31, Art. 8 


feeding activities. The senior author 
suspects that the sediments exert an 
oxygen demand in the lakes, just as 
they do in the river. In August, 1974 
dissolved oxygen levels in Meredosia 
Lake (river mile 72-77) were approxi- 
mately 3 mg/l when a strong wind was 
blowing that stirred bottom sediments 
in the shallow lake. A die-off of gizzard 
shad was occurring, and almost all the 
fingernail clams maintained in plastic 
cages on the bottom of the lake had 
died since they were last checked in 
mid-July. Oxygen levels may have been 
lower than 3 mg/1 on previous occa- 
sions. Oxygen levels in the river on 
the same date were approximately 6 
mg/l. It is also possible that toxic 
materials, such as pesticides, that are 
bound to soil particles, were taken up 
by aquatic organisms such as clams 
that ingested the soil particles or passed 
them over their respiratory membranes. 
In addition, toxicants such as hydrogen 
sulfide may have been formed and 
released from bottom muds under 
anaerobic conditions. 

The increased barge traffic (Starrett 
1972:153) associated with the improved 
navigation channel increases the tur- 
bidity of the river. The turbulence 
produced in midchannel, as well as the 
washing action along shore, resuspends 
sediment, thereby increasing the tur- 
bidity (Fig. 1 and 2). W. C. Starrett 
made numerous observations of the 
effect of barges on turbidity of the 
river, for example (Starrett 1971:273): 

“A towboat underway causes a 
strong current and washing action on 
the silt bottom (“false bottom”) in- 
shore, which resuspends the silt par- 
ticles, thereby increasing the tur- 
bidity. The increase in turbidity is 
more noticeable in the lower three 
pools, particularly in the Alton Pool, 
than it is upstream because of dif- 
ferences in bottom types. ... The 
outrush of water from shore toward 
the channel caused by a towboat 
also temporarily ae the shallow 
areas. On November 18, 1964, in 

the Alton Pool at river mile 65.1, 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS River 341 


the turbidity just prior to the passing 

of two towboats was 108 units (Jack- 

son turbidity units), and within 6 

minutes after the tows had passed, 

the turbidity was 320 units. Sixteen 
minutes later the turbidity had 
dropped to 240 units.” 

Some personal observations were 
made on the effects of towboats during 
the 1974 electrofishing investigations. 
On several occasions, flow reversals 
in chutes were observed as tows passed 
first one end, then the other, of a 
chute. In a narrow part of the river 
channel above Pekin on 19 September, 
1974, in the midst of electrofishing, our 
boat was stranded on the mud when 
the water rushed out from shore as a 
tow of nine fully loaded coal barges 
passed upstream. Mussel shells were 
clearly visible on the bottom for several 
seconds before the water rushed back 
again. We had been in approximately 
0.5 m of water. 

Such washing along the shore and 
flow reversals in side channels may 
have a detrimental effect on benthic 
organisms and fishes that make nests 
in shallow water, such as sunfishes. 

Low flows from 1962 to 1964, and 
consequent low oxygen levels and re- 
duced dilution of toxic wastes, ap- 
parently are responsible for the decline 
during the same period of game species 
such as largemouth bass, crappies, and 
bluegill. Catches of these species 
showed dramatic recoveries following 
the high-water period 1971-1973. In 
14 years of electrofishing, covering the 
period 1959-1974, the largest numbers 
of the following species were obtained 
in 1974, following the high water pe- 
riod: black crappie, white crappie, 
flathead catfish, white bass, bluegill, 
bigmouth buffalo, and black buffalo. 
The maximum weights of the following 
species were obtained in 1974: white 
crappie, channel catfish, and bluegill. 
Fig. 8, 9, and 10 show that bass popula- 
tions still had not recovered to the peak 
levels observed in Peoria and La 
Grange Pools in the years 1959-1962. 


High water levels stimulate certain 
species, such as white bass, to run up 
tributary streams and spawn. White 
bass were obtained in the upstream 
pools, Starved Rock and Marseilles, in 
fairly substantial numbers in 1973 and 
1974, whereas none were obtained in 
these pools in 1959, 1961, 1963, 1964, 
1968, and 1969. High water also in- 
creases the space available for spawn- 
ing activities of fishes that build nests 
in shallow water, such as sunfishes, and 
the amount of protected habitat avail- 
able for juvenile fish, in shallow, flooded 
areas and around brush and _ tree 
stumps. As mentioned above, higher 
oxygen levels have occurred in the 
Illinois River in association with the 
high flows, with beneficial effects on 
fish and fish food organisms. 

In spite of the improvement in the 
electrofishing catch in 1973 and 1974, 
apparently due to high water levels in 
1971-1973, the commercial catch of fish 
in the Illinois River continued its his- 
torical decline in the 1970's (Table 28). 
Depending on whether the Illinois De- 
partment of Conservation figures or the 
National Marine Fisheries Service sta- 
tistics are used, the catch dipped under 
1 million pounds (454,000 kg) in 1971 
or 1972. The decline is not explained 
by a reduction in the number of com- 
mercial fishermen—there were 13 full 
time and 56 part time Illinois River 
commercial fishermen in 1973, and 9 
full time and 47 part time in 1971. Nor 
is it explained by a decline in economic 
value of the catch. The catch from the 
Mississippi River bordering Illinois has 
been relatively constant from 1950 
through 1973 (Table 28). A general 
decline in profits would be reflected 
in a general decline in fishing effort 
in both the Illinois and Mississippi 
Rivers and a corresponding decline in 
catch. It is possible that because fish- 
ermen generally take large adult fish, 
an increase in the catch of commer- 
cially important sizes of fish will not be 
seen until the fish spawned in 1973 and 
1974 reach marketable size. 


342 


FUTURE IMPACTS 
ON THE FISH POPULATIONS 
OF THE ILLINOIS RIVER 


In 1971 the Chicago Metropolitan 
Sanitary District began a large-scale 
sludge recycling project near the IlIli- 
nois River at St. David. In 1974, the 
District began aerating a section of 
the Chicago Sanitary and Ship Canal, 
and more of the canal will be aerated 
in succeeding years. In the future, all 
Chicago storm water probably will be 
captured and stored in a deep tunnel 
under Chicago, instead of being dis- 
charged into the canal, and will be 
treated before it is released to the 
canal, Advanced waste treatment plants 
should be capable of removing the 
ammonia that now exerts an oxygen 
demand so far down river. All of these 
improvements in waste treatment will 
have a beneficial impact on the aquatic 
life in the river, by reducing the oxygen 
demand on the river and improving 
oxygen levels during critical low-flow 
periods. Waste treatment probably will 
also be improved in the Pekin-Peoria 
metropolitan area. 


A proposed increase in the depth of 
the navigation channel of the Illinois 
River (from 2.7 to 3.7 m), would be 
accomplished by a combination of rais- 
ing low-flow water levels and dredging. 
Depending on local topography, the 
water surface area might be increased. 
Judging by the increased fishery in the 
Illinois River following a rise in water 
levels in 1900, as a result of water di- 
version from Lake Michigan, one might 
expect a beneficial effect. However, 
bottomland lakes that now have a 
chance to clear during periods when 
they are cut off from the river might 
then become permanently connected to 
the river and receive a continuous, 
rather than intermittent, input of oxy- 
gen-demanding sediment. In 1921, 
Richardson (1921a:418) reported that 
Quiver Lake (mile 121.0-mile 124.0) 
and Matanzas Lake (mile 114.5-117.0) 
received spring water from the sandy 


InLino1is NaTurAL History SurvEY BULLETIN 


Vol. 31, Art. 8 


bluffs on the east side, and that the 
waters in these lakes were somewhat 
clearer than in other bottomland lakes. 
According to an Illinois Water Survey 
report (Singh & Stall 1973:19), the 
influx of ground water to the river from 
Kingston Mines (mile 145.3) to Mere- 
dosia (mile 71.1) amounts to 8.75 m°3/ 
sec, or about one-twelfth of the total 
input to this section of the river, during 
the lowest flow expected for a 7-day 
period at a recurrence interval of 10 
years. According to Matanzas Beach 
residents, the water and shoreline of 
Lake Matanzas still are cleared of silt 
deposited by the river, due to the flush- 
ing action of ground water coming 
through the sandy bottom along the 
bluff. In contrast, Quiver Lake is now 
filled with silt. 


The Illinois Department of Conserva- 
tion has been able to restore aquatic 
vegetation to Rice Lake (mile 133-137) 
and Stump Lake (approximately mile 
5) by pumping water out of the lakes 
or allowing them to dry out naturally 
(personal communication, Robert L. 
Glesenkamp, Area Wildlife Manager, 
Illinois Department of Conservation). 
Midsummer drying was a natural oc- 
currence in this type of shallow lake, 
during low-flow years, prior to Lake 
Michigan diversion and construction of 
navigation dams (Richardson 1921a: 
419). On drying, the bottom muds were 
compacted, and when the lakes were 
reflooded, the turbid water generally 
cleared, and the plants gained root- 
hold in the firm bottom. Restoration 
efforts would be more difficult if sum- 
mer water levels were higher. In addi- 
tion, private duck clubs and state and 
federal wildlife refuges along the river 
would find it difficult to reduce water 
levels. They attempt to reduce water 
levels to expose mud flats and encour- 
age the growth of moist-soil food plants 
for waterfowl. Once again, a natural 
drying cycle has had to be replaced 
or supplemented by pumping, because 
water levels do not attain the low 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIVER 343 


levels they once did. Such management 
techniques require energy, equipment, 
and manpower. 


Larger towboats using the improved 
navigation channel and an increased 
number of towboats would keep more 
silt in suspension and increase the 
washing action along the shore and 
flow reversals in chutes. Fig. 1 shows 
that if towboats pass a point in the 
river more frequently than once every 
2% hours, the resuspended sediment will 
not have a chance to settle out and the 
average amount of sediment suspended 
in the water will increase with a con- 
sequent increase in oxygen demand 
and turbidity. The more silt there is 
in suspension in the river, the faster 
bottomland lakes such as Lake Chau- 
tauqua (mile 124-130) will fill with 
oxygen-demanding sediment, as they 
are periodically overflowed by the river. 


The effect of various future channel 
improvement schemes and_ various 
levels of boat traffic on the siltation rate 
in the critical backwater areas and 
lakes needs to be predicted. In addi- 
tion, the joint effects of man’s activities 
in the river and drainage basin needs 
to be assessed. For example, it is pos- 
sible that the proposed increase in di- 
version of Lake Michigan water at Chi- 
cago (discussed in more detail below) 
may make it possible for the present 
channel to accommodate deeper-draft 
barges in certain areas, without addi- 
tional dredging or higher dams. 


It would be counter-productive for 
one arm of government to spend re- 
sources in improving and restoring 
refuge areas if another arm of govern- 
ment engages in practices which de- 
grade such areas. There will be little 
benefit to the fisheries of the Illinois 
River by having the Chicago Metro- 
politan Sanitary District and other mu- 
nicipalities and industries expend bil- 
lions of dollars in improved waste 
treatment if the river and its bottom- 
land lakes are increasingly degraded 
by silt. Refuges, unpolluted lakes, and 


unpolluted tributary streams must be 
maintained if the river is to show the 
recovery pattern in the future that it 
exhibited in 1973-1974, following the 
high-water period and improved oxy- 
gen levels from 1971-1973. When for- 
merly degraded areas are restored, they 
can be recolonized rapidly by species 
that are desirable to man, if reservoirs 
of such species, and reservoirs of food 
organisms for desirable species, are 
available in undegraded pockets in the 
ecosystem. In a properly functioning 
system, the refuges maintained by man 
have precisely this function. 


The most practicable solution to the 
silt problem may be to reduce the 
amount entering the river in the first 
place, if predictive studies indicate that 
a reduction of silt input would actually 
reduce siltation in the lakes and back- 
waters. Once the silt is in the river 
and lakes, it may be recycled and re- 
suspended there, and it is possible that 
no reduction in turbidity or oxygen 
demand would be achieved by reduc- 
tion of silt input without the use of 
restoration techniques, such as drying 
out of lakes. On the other hand, it is 
possible that reduced silt input may 
cause the river to flush out backwater 
areas and lakes during periods of high 
flow, thus bringing about a natural 
restoration of these areas. Once the 
turbidity was reduced, fringing marshes 
and beds of aquatic plants might ap- 
pear again, further accelerating restora- 
tion by acting as silt filters and nutrient 
traps. 


The silt entering the river could be 
reduced by wide adoption of soil con- 
servation practices in the Illinois basin, 
including such new practices as no-till 
farming, where row crops are planted 
without greatly disturbing the soil. Be- 
fore the latter practice is adopted on a 
wide scale, the total energy require- 
ments (including the energy for the 
manufacture of agricultural chemicals ) 
of various alternative farming methods 
need to be determined, and the en- 


344 


vironmental impact of the herbicides 
that must be used with present no-till 
farming methods needs to be assessed. 


The City of Chicago and lakefront 
residents whose property has been dam- 
aged as a result of current high water 
levels in Lake Michigan have requested 
an increased diversion of Lake Michi- 
gan water into the Illinois River. An 
increased diversion would probably 
raise water levels, with some of the 
detrimental effects discussed above. 
However, Lake Michigan water is good 
quality water and probably would im- 
prove the quality of the upper river 
by a simple dilution, if diversion oc- 
curred during the summer months. On 
the other hand, if ammonia removal is 
not achieved by the Chicago Metro- 
politan Sanitary District, the effect of 
increased diversion might be to push 
this oxygen-demanding waste farther 
downstream before its oxygen demand 
could be satisfied. 


Two introduced species have entered 
the Illinois River recently and will 
probably become more abundant, just 
as the introduced carp, goldfish, and 
white catfish have. It is difficult to pre- 
dict whether the latest arrivals will in- 
crease explosively, as carp and goldfish 
did, or whether they will barely main- 
tain themselves, as white catfish have. 
White catfish are only occasionally taken 
from the Illinois River and do not seem 
to reproduce abundantly in the river. 
The white amur (Ctenopharyngodon 
idella), a plant-eating fish introduced 
from Asia, is now being taken regularly 
by commercial fishermen from the Mis- 
sissippi River at Crystal City, Missouri 
and from the Missouri River (Personal 
communications, William L. Pflieger, 
Fishery Biologist, Missouri Department 
of Conservation, and Peter Paladino, 
District Fishery Biologist, Illinois De- 
partment of Conservation), and has 
probably entered the lower Illinois 
River. If rooted aquatic vegetation 
could be restored to the Illinois River 
and its bottomland lakes by the lake 


Ixutinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 8 


restoration techniques discussed above, 
or by a reduction of silt loads in the 
river as a result of improved soil con- 
servation practices in the basin, the 
white amur might have a detrimental 
impact. On the other hand, white amur 
from the Mississippi are being mar- 
keted in small quantities commercially 
and their flavor is reported to be ex- 
cellent. White amur in the Missis- 
sippi grow to a large size (4.56.4 
kg) in 2 years (Personal communica- 
tions, Pflieger and Paladino). They 
might become a useful commercial 
species in the Illinois River. 

Another exotic species, the Asiatic 
clam (Corbicula manilensis) was found 
at three locations on the Illinois in 
the course of the 1974 electrofishing 
survey: at Kampsville (river mile 
32.0), Bath Chute (mile 106.7), and 
Turkey Island Chute (mile 148.4) 
(Thompson & Sparks, in press). The 
Asiatic clam is a serious nuisance, be- 
cause it has blocked condenser tubes 
of power plants in Illinois and else- 
where. In addition, it may displace the 
native fingernail clams. 

The future of the Illinois River will 
largely be determined by man’s activi- 
ties in the river and adjacent flood- 
plain and by his use of the land in the 
drainage basin. Predictions of the im- 
pacts of various activities must be de- 
veloped, so a rational management 
scheme for the Illinois River can be 
designed and the river can continue to 
serve a variety of purposes in the fu- 
ture. 


SUMMARY 


1. The upper Illinois River is 
warmer than the lower River, as a re- 
sult of warm municipal and industrial 
effluents. 

2. The upper river is less turbid, be- 
cause the bottom is generally rocky, 
whereas Peoria, La Grange, and Alton 
Pools contain flocculent muds that have 
entered the river and are kept in sus- 
pension by the river current and by 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIvER 345 


wave action resulting from wind, tow- 
boats, and pleasurecraft. 

3. Dissolved oxygen levels at the sur- 
face and the bottom of the river were 
virtually the same in the fall of 1974, and 
dissolved oxygen levels were 77-97 per- 
cent of saturation in Alton Pool, 65-122 
percent of saturation in La Grange and 
Peoria Pools, and 47-104 percent of 
saturation in the upper Pools of Starved 
Rock, Marseilles, and Dresden. Local 
areas of super-saturation occurred 
where plankton blooms appeared to 
be in progress. In two areas that 
provided good physical habitat for 
largemouth bass, Lower Bath Chute, 
La Grange Pool (mile 107) and Chilli- 
cothe Island Chute, Peoria Pool (mile 
180), midsummer oxygen levels were 
at 35 percent saturation or below for 
4-5 years out of the 8-year period 
1963-1970. Laboratory experiments 
have shown that oxygen levels below 
35 percent saturation reduce the sur- 
vival of larval largemouth bass and 
levels below 70 percent retard their 
growth. 


4, The number of fish species taken 
by electrofishing in the Dresden Pool, 
Des Plaines River portion of the Illinois 
Waterway during the period 1959-1974 
was consistently low (Tables 29 and 
30). Only carp and goldfish and hy- 
brids of these two pollution-tolerant 
species were commonly taken. 

5. The following species showed a 
trend of increasing abundance in the 
downstream direction, away from Chi- 
cago, with the largest number occur- 
ring in Alton Pool: shortnose gar, 
bowfin, goldeye, mooneye, channel cat- 
fish, flathead catfish, and white bass. 

6. Goldfish showed a trend of in- 
creasing abundance in the upstream 
direction, toward Chicago. 


7. The following species were most 
abundant in one or both of the two 
middle pools of the river, La Grange 
and Peoria Pools, which have the most 
connecting lake area: gizzard shad, 
carp, river carpsucker, smallmouth buf- 


falo, bigmouth buffalo, black buffalo, 
yellow bullhead, green sunfish, bluegill, 
largemouth bass, white crappie, black 
crappie, and freshwater drum. 

8. Gizzard shad and carp were gen- 
erally abundant throughout the river. 

9. Black bullheads were abundant 
at one atypical station, Ballard Island 
Chute, Marseilles Pool (mile 247.8- 
248.2), which apparently provides pre- 
ferred habitat for this species. 

10. Gamefish populations declined 
during the low water years 1962-1964, 
and recovered following the high water 
years 1971-1973. Largemouth bass 
populations did not recover to 1959- 
1962 levels. The recovery appears at- 
tributable to improved oxygen levels 
in the river, and perhaps to increased 
dilution of toxic materials, and demon- 
strates how rapidly fish populations re- 
spond to improved conditions in the 
river. 

11. The commercial and sport fish- 
eries in the Illinois River have gen- 
erally declined from levels around the 
turn of the century. The decline is 
attributable to a loss of habitat and in- 
creasing pollution. Habitat was lost 
due to leveeing and draining of bottom- 
land areas in the period 1903-1926 and 
due to sedimentation in the remaining 
areas. Sedimentation has resulted in 
undesirable habitat modification, as 
well as habitat reduction. 

12. Northern pike, yellow perch, and 
walleye (Stizostedion vitreum vitreum ) 
were once abundant in the river but 
are now rare or limited in their distribu- 
tion. Yellow perch populations have 
declined probably as the result of the 
disappearance of beds of aquatic plants 
and disappearance of clean sand or 
pebble substrates perch use for spawn- 
ing. 

13. In the past the bottomland lakes 
and backwater areas offered havens for 
fish and fish food organisms, as the 
river became increasingly polluted. 
Now dissolved oxygen levels in the 
river seem to have improved, while 


346 


the lakes have filled with sediment that 
apparently exerts an oxygen demand, 
keeps aquatic plants from growing, and 
does not support an abundance of food 
organisms. 

14. More and better waste treat- 
ment facilities are being constructed 
by industries and municipalities in the 
drainage basin of the Illinois River. 
However, the production of fish and 
wildlife in the Illinois River and its 
bottomland lakes is not likely to im- 
prove unless sediment pollution is also 
brought under control. 


15. The consequences of future uses 
of land in the drainage basin and the 
consequences of future uses of the river 
must be predicted, so that a wise se- 
lection of alternatives can be made. 
If the river is to be managed in the 
future for a variety of beneficial uses, 
then the various state, federal, and pri- 
vate agencies charged with managing 
land and water within the drainage 
basin must work in a coordinated 
fashion, rather than at cross purposes. 


GUIDE FOR USE OF TABLES 
OF ELECTROFISHING RESULTS 
(Tables 3-27) 


SYMBOL EXPLANATION 
1 Dresden Pool, Des Plaines 
River—not included in tabu- 
lated value for the [Illinois 
River at bottom of each table. 


2 Values represent the total 
number of fish or total weight 


Iuuiwois Natural History SuRvEY BULLETIN 


Vol, 31, Art. 8 


of fish taken during the desig- 
nated year in the Illinois River 
divided by the number of half- 
hour intervals fished. Illinois 
River pools are Alton, La 
Grange, Peoria, Starved Rock 
and Marseilles. The Dresden 
Pool, Des Plaines River, is ex- 
cluded from this tabulation. 


# Denotes less than 0.01 kilo- 
grams or fish per 30 minutes 
fished. 


Note: Fish species are listed in phylo- 
genetic order. All common and 
scientific names are taken from 
A List of Common and Scientific 
Names of Fishes from the United 
States and Canada, 3rd edition, 
1970, American Fisheries Society 
Special Publication No. 6. Spe- 
cies that were rarely taken by 
electrofishing are not shown in 
the tables, but are discussed in 
the text. The values in the body 
of each table are determined by 
summing the number of fish or 
weight of fish obtained at all 
stations in the navigation pool 
and dividing the sum by the 
total number of half-hour inter- 
vals fished in that pool. Thus 
the values are average catches 
per unit effort for each pool. 
The number of electrofishing sta- 
tions in each pool are as follows: 
Alton Pool (4-5), La Grange 
Pool (6), Peoria Pool (8) 
Starved Rock Pool (2), Mar- 
seilles Pool (3), and Dresden 
Pool (1). 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIvER 347 


Table 1.— Illinois Natural History Survey 
electrofishing sites on the IIlinois Waterway, 
1959-1974. 


Pool Station River Mile* 


Alton Mortland Island 
Chute 


Below Hardin 


Diamond Island 
Chute 
Above Hardin 24.0-25.5 


Hurricane Island Chute 
Above Hardin 26.0-27.2 
Crater Island and 
Willow Island Chutes 
Below Kampsville 
Big Blue Island 
Chute? 

Above Florence 57.5-58.9 


LaGrange Bar Island and 
Grape Island Chutes 
Below Beardstown 
Sugar Creek Island 
Chute 
Below Browning 
Lower Bath Chute 
Above Browning 
Upper Bath Chute 
Above Bath 
Turkey Island Chute 
Above Kingston 
Mines 
Illinois River 
Above Pekin 154.5-155.3 


Peoria Lower Peoria Lake 
Near East Peoria 
Middle Peoria Lake 
Near Peoria Heights 
Conservation Landing 
at Detweiller Park 169.2-171.0 


18.7-19.4 


29.3-30.7 


86.2-87.1 


94.3-95.2 


106.8-107.5 


112.8-113.3 


147.3-148.2 


163.0-163.4 


Table 1.—Continued 


Pool Station River Mile* 


Peoria Chillicothe Island 
Chute 


Above Chillicothe 


Henry Island Chute 
Below Henry 


Lower Twin Sisters 
Island Chute 
Above Henry 


Upper Twin Sisters 
Island Chute 
Above Henry 


Hennepin Island 
Chute 
At Hennepin 


Clark Island Chute 
Below Spring 
Valley 214.9-215.6 


Starved Bulls Island 
Rock Chute 
Above Ottawa 


Bulls Island Bend 
Section 
Above Ottawa 241.4-241.9 


Marseilles Ballard Island Chute 
Above Marseilles 247.8—-248.2 


Johnson Island Chute 
Above Marseilles 249.4-249.9 


Sugar Island Chute 
Below Morris 260.2—261.0 


Dresden, Rapp’s Boat Yard 
Des Plaines and Du Page River 
River Mouth 
Above Channahon 276.8—277.8 


180.1—-181.0 


193.5-194.1 


202.2-203.1 


203.1-203.5 


207.0—208.0 


240.5-241.1 


8 Stations are located by river miles rather 
than by kilometers because existing river 
charts and navigation aids along the river use 
mileages, 

> Fished in 1974 but not in previous years. 


Vol. 31, Art. 8 


Intino1is NATURAL History SURVEY BULLETIN 


348 


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6T CS) oLL &T9 VLE ggg0-3ny §Z L0&-€°6¢ sojnyO PUBIS] MOTITM Pues 19}e1) 
&@ 819 v8 819 616 Ges0-sny 7 @ L2-0°96 9yngOD pues] sueolliny mo}vIV 
8T 60°L $16 Teh G66 O¢ST-2ny 17 G'G2-0'°F6 aynygO pue[s] puowmeid 
8T org S08 819 61Z 0s60-Sny TZ ¥6T-L'8T 9ynyOD pues] puelj}IOW 
wa mad Ws % wmdd Do (LS0) awuL IW LO0teT 4017018 100d 
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"pL6| “ACmiaieny 


sioul|j] 243 40 Aeauns Burysijosj29;9 ue Bulunp pauieygo sanjea AyjigisiA (°G°S) >SIP !y229eg pue uaBAxo panjossip ‘ainyesaduua}y Jayep~,—'Z FIFE) 


ELECTROFISHING SURVEY OF ILLINOIS RiveR 349 


Aug., 1975 Sparks & STARRETT 


b0°0 00°0 
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000 
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# T0'0 400 00°0 400 
00°0 00°0 600 00°0 00°0 
00°0 #00 60°0 00°0 €T'0 
200 00°0 00°0 00°0 00°0 
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10°0 00'0 00°0 00°0 00°0 
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SONU, OF 4ad 42QUWNN 
0°22 az 0°92 "8% a'8Z 
L96T 996T G96T $961 £961 


00°0 


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paysig sino {0 42QUNN pud 1DaX 


zu Tl 

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zu Tl 
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“bL61-6S61 ‘Aemuazeny slourj|; ayy ul Burysijosjoaj9 Aq uayxe} (snwojysojeyd snoysosiday) 1e6 asouwroys—e 3jqe) 


Vol. 31, Art. 8 


Inuino1s NATuRAL History SURVEY BULLETIN 


350 


T0°0 00°0 or 0 00°0 10°0 00°0 00°0 00°0 00°0 00°0 T0°0 80°0 00°0 00°0 zu il 
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00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 YWooY Peareys 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So[[TesreW 
00°0 00°0 00°0 00°0 depseiqg 


Sanur OF 4ad 42QUnN 


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paysig sino [0 4aQUNN pUD 1DaX 


SS esse sees 8 ees ee ee 
————————— 


"PLO L-6S61 ‘Aemuaye ny Siourjj] ayy Ul Busysiyos329]9 Aq Uayey (BAJED ErMIY) ULYMOg—"p 2I/qe 1 


ELECTROFISHING SURVEY OF ILLINOIS RIvER 351 


Aug., 1975 Sparks & STARRETT 


290 £r'0 970 030 8F'0 
Gz°0 # oro 
690 6F'0 0+'0 oz £0'T 
080 670 &r'0 6z'0 9z°0 
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760 810 910 970 620 
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0z9t L8TZ OFTT O0FL 69°8F 
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Saynurpl OF 4aq Swuvs60)1y 


8H'OF 
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02°69 
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Bulysijosjoaja Aq uaxe} (winueipeda> ewosoi0g) peys PseZZIQ—"G a/4P 1 


: 


Vol. 31, Art. 8 


ILLino1is NaturRAL History SuRVEY BULLETIN 


352 


# T0'0 # # # T0'0 0070 200 T0'0 # # 00°0 # 00°0 vu Il 
00°0 T0'0 00°0 10°0 00°0 10°0 60'0 # 00°0 m01Ly 
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S00 00°0 00'0 00°0 000 00°0 00°0 00°0 00°0 00°0 £0'0 00°0 00°0 00°0 So] [10812] 
00°0 00°0 00°0 \wepseiq 
Sanur, OF 42d swmvs6071y 
&0'0 £0°0 # ¢0'0 200 120 00°0 110 610 200 Z0'0 00°0 400 00°0 rar a 10 
00'0 620 00'0 0ST 000 $90 &TT 810 00°0 u071V 
00°0 80°0 010 00°0 00°0 00°0 00°0 00°0 00°0 00°0 400 00°0 00°0 esueiy eT 
00°0 00°0 00°0 00°0 10°0 00°0 00°0 810 00°0 00°0 00°0 000 010 00°0 BI10ed 
00°0 00°0 00°0 00°0 00°0 000 0070 00°0 00°0 00°0 00°0 00°0 00°0 00°0 yooy peareis 
0+'0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 10°0 00°0 00°0 00°0 SoT[Tes1e 
00°0 00°0 00°0 wepseiq 


SaINWI, OF 4ad 42QUNN 


81s S'6T gt 0°32 0°06 0°82 GTé 0°93 G'&s G&S GP 0°0T Gor OCT 100q 
PLET €L6T OL6T 696T 896T L96T 996T G96T v96T €96T 6961 T96T 096T 6S6T 


paysig sino fo 4aqunn pun 02x 


“pL61-6S6| ‘Aemuayepy siourj|| 243 ut Buyysizosjoaja Aq uaye} (S@prosoje uopolpyy) aAepjog—'9 2/qe 1 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 353 


Aug., 1975 Sparks & STARRETT 


# 00°0 00°0 00°0 00°0 00°0 00°0 # 00°0 100 00°0 00°0 00°0 00°0 zu lll 
00°0 00°0 00°0 00°0 00°0 £0°0 00°0 S00 00°0 UovIV 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 aduBiy eT 
# 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 Bl10ed 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 HooY P9areys 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So] [os1e AL 
00°0 00°0 00°0 00°0 Wepseaiqd 
saynurpy OF 4ad Swvsb0]1y 
60°0 00°0 00°0 00°0 00°0 00°0 00°0 80°0 00°0 60°0 00°0 00°0 00°0 00°0 cu TL 
00°0 00°0 00°0 00°0 00°0 98°0 00°0 610 00°0 moyy 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 asuBlyH BT 
0oT'0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 BIL09d 
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81S G6. Get 002 03% 0°33 ia 4 0°93 ie 4 SEs GPP 00T Ger as 100d 
FL6T €L6T OL6T 696T 896T L96T 996T G96T F96T €96T 96T T96T 096T 6S6T 


paysig sinoy fo 4aqunn pup 1vax 


eee nny rt 


“pL61-6S61 ‘Aemsayeny siouy|| 24) U! 


Busysijosjoaja Aq uaye} (snsi61ey uoporpyy) 2BA2U00\Y—"/ 9/921 


Vol, 31, Art. 8 


Intinois NaturAL History SuRVEY BULLETIN 


354 


$00 oro 89°0 120 8T0 620 92°0 09°0 Le0 TL'0 660 C6 T ce T £10 vu Tl 
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00°0 tat OF ST 00° L9°0T €e°9 €&°9 GL'té SLOT GLLT 0g°0€ 0s"r9 OF FZ 0g3 Wo0Y Peasejys 
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09°8T LOVE Gc TOT 0g'9T yaepseiqg 


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STs G6T S&T 0°36 0°32 0°66 ia 4 0°96 ges g'&s GOP 00T Ger OCT 100d 
vL6T €L6T OL6T 696T 896T L96T 996T G96T v96T €96T C96T T96T 096T 6961 


peysig sinog fo 4aqunn pup 02x 


“pL61-6S6| ‘Aemuareny stour|| ayy ul Burysizouj2a]9 Aq uaye} (Sngeane SmIsseAeD) YS1xP][ON—'B 2I9eL 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 355 


Aug., 1975 Sparks & STARRETT 


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“pL6l-6S6| “Aemsayeny siourj| 24} ul Gurysijosyoaj9 Aq uaxe} (Snyeane snisseses x o1dued snundAy) ysiypjop x duey—'6 ajqe) 


Vol. 31, Art. 8 


Ittivo1s NaturAL History SuRvEY BULLETIN 


356 


8 TT 18°8 &¢'&T be TT 68°0T 86C1 SL FT £26 LV TC 99°81 Ciara § 96°0T 6L0T 619 vu TL 
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"bL61-6S61 ‘AeMIaIeAA sloulj|| 24} Ul Bulysijosj99}a Aq uayxe} (oldaed snundAD) died—'O| 9/921 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 357 


_ Aug., 1975 Sparks & STARRETT 


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Ons 00'T 00°0 00°0 £0 00°0 £80 00°0 00°0 00°0 oro 00°0 09°0 00°0 yooy pealeys 
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ee OO oOoq“qzgq?zepeaeoe0eOS=$S= SSS sa 
“bL61-6S6L ‘AemsayeAQ Sioun|| ay} u! Buiysijosj>a;9 Aq uaxe} (o:daed sapoidsed) dayonsdied 49ANY— "| | FIG2L 


_ a = os = — — — I 


Vol. 31, Art. 8 


In~inois NaTuRAL History SURVEY BULLETIN 


358 


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oro 880 &T0 00°0 00°0 60°0 00°0 G20 00°0 movLVy 
00°0 ST 0 00°0 0 £0 00°0 G20 610 £20 *5°0 80°0 00°0 00°0 aZUBIt) BT 
00°0 &T0 oT 0 TT eet 09°0 610 &&°0 88°0 TT 620 9£°0 0¢°0 00°0 Bl109d 
OFT 19°0 OFT 00T L9@ LOT 00% 00°§ GLc SLT 09°0 00°0 0F'0 00°0 HOY Psaleys 
01°70 00 T os'T LTT eT 00% 0s°0 00°0 ee T LT0 00°0 G20 00°0 00°0 So] [Tos rey 
00°0 00°0 00°0 00°0 uepseiqd 


SaInurpl OF 42d 42QUNN 
EEE ES ot ie a Se, Ne Oa Se eee ee 
81S Go 6r gst ad 02 0°03 bia 4 0°92 gto Gs GF 00T Gor OCT 100d 
LET €L6T OL6T 6961 896T L96T 996T G96T F96T €96T 961 T96T 096T 6S6T 


paysig Sino {0 4aQuUNnN PUD 12K 
os — OOOH 


"bL61-6S61 ‘Aemiayepy soul) a4} ul Gurysiyosj99]9 Aq uae} (smujadAd sapoidaey) sax2nsdse> xIeEqIJINO—'Z1 FIGEL 


ELECTROFISHING SURVEY OF ILLINOIS RivER 359 


Aug., 1975 Sparks & STARRETT 


S70 &T0 
00°0 

$20 600 
€&°0 120 
OTT cr 0 
00°0 600 
00°0 00°0 
Sr'0 920 
00°0 

0f°0 80°0 
0L°0 0F'0 
00% ££°0 
00°0 02°0 
00°0 00°0 
81s G'6T 
PL6T &L6T 


9T 0 


8£°0 
0T 0 
*T 0 
00°0 


0&0 


09°0 
02°0 
0f°0 
0T0 


G'éT 
OL6T 


oT 0 
00°0 
60°0 
GeO 
00°0 
00°0 


oT 0 
00°0 
8T0 
+10 
00°0 
00°0 


St°0 
00°0 
8£°0 
£0'T 
00°0 
00°0 


saynurpy OF 4ad swo0sbo]Ly 


820 LT0 92°0 
oT 0 00°0 $00 
Lg°0 g0°0 12'0 
Te°0 S¥'0 +90 
00°0 00°0 60°0 
00°0 00°0 600 
69°0 Sr'0 08°0 
6T 0 00°0 6T 0 
PET LT0 €8°0 
09°0 02 T LYT 
00°0 00°0 €¢°0 
00°0 00°0 LT 0 
0°62 0°32 0°32 
696T 896T L96T 


£80 
00°0 
G20 
610 
00°0 
00°0 


GTZ 


9961 


eT 0 
00°0 
s20 
&T0 
00°0 
00°0 


0°92 


G96T 


T3°0 
00°0 
690 
L3'T 
00°0 
00°0 


SaINUL OF 4ad 4aQuUnN 


GES 


P96T 


6L°0 
£0°0 
02% 
690 
00°0 
00°0 


160 
&T0 
ead 
glo 
00°0 
00°0 


G&% 


€96T 


£0 
&T0 
€L°0 
£0 
00°0 
00°0 
00°0 


89°0 
09°0 
PST 
9¢°0 
00°0 
00°0 
00°0 


GP 
o96T 


paysiq sinoy fo 4aqunyn pup wax 


02°0 


62°0 
00°0 
00°0 


G20 


9€°0 
00°0 
00°0 


0°0T 
T96T 


st 0 


00°0 
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490 
00°0 


02°0 


00°0 
02°0 
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00°0 


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096T 


Le°0 


99'T 
or 0 
00°0 
00°0 
00°0 


OTT 


Gog 
&T0 
00°0 
00°0 
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OCT 
6961 


zu Tl 

mov 
aZueIy eT 
B1100d 

YyooY Pealeig 
So] [1esIey 
,Wepseig 


vu ll 
uoyLV 
asueiy ey 
el100g 


WooYy peaiej}s 
Se[[1os1e 


,Uepseiq 


100d 


"PLO 1-656 1 ‘Aemuaieny Slour|| 43 UI Burysiyosyoaja Aq uaxey (SMJEGnG SNgoN>}) oJe}4Nq YINoWjeUIs—"_E| >qeL 


Vol, 31, Art. 


Iutino1is NaturAL History SURVEY BULLETIN 


360 


60'S LET 490 Lass LOE 99°9 LvY es 194 evy ¥8G 19° oT £Vs ra: 0g 
00°0 ££°0 rr0 8L°0 TOT 00°0 98°0 TL'0 Lv0 > mouV 
pL T PIT 89°0 LTT ePT 869 89°€ 18 T 198 068 LOY LLY aa s esueIp eT 
ST8 08% 8IT 118 9¢°8 IT &t 166 819 80°9 96° 867 L9¢ €L0 COT BI109d 
610 00°0 00°0 00°0 00°0 00°0 610 00°0 00°0 00°0 000 00°0 00°0 00°0 HooY Pearejs 
10°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So] [Tesre yl 
00°0 00°0 00°0 00°0 1uepseiqd 
SaINUIAL OF 4aq Swvsb0]1y 
8r9 oss 0s°0 96% eL% 01g £07 88 T Ce F 8OF 81 08'T 0c T G8°s vu Ul 
00°0 610 &10 £9°0 1&0 00°0 690 0s°0 09°0 movV 
oss C6 T 0r0 £80 eT Sos Pat rT 668 oS'6 Tes 60¢ SL6 esueIp eT 
Oc LT eeP 06°0 02°9 L8°9 kar § 666 00°S ors ooh 68'S Lg 0L'0 Ses e1109d 
0&0 00°0 00°0 00°0 00°0 00°0 €€0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 WooY Peareys 
020 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So[[Tosreyl 
00°0 00°0 00°0 00°0 1depseid 
SaInurp OF 4ad 42QUnN 
81s S6T gst 0°66 036 0°66 GTs 0°96 Go Gs GHP 00T Ger OCT 100g 


PL6T €L6T OL6T 696T 896T L96T 996T G96T P96T €96T G96T T96T 0961 6S6T 


paysig sinoy fo 4aquUnn pud w0ax 


"PLO 1-656 1 ‘Aemuayeny Siouijj| 243 Us Burysijos399}9 Aq uaxey (SMyjauIadAd snqon>}) Oje4¥3nq YINoWBIg—"p| 1921 


ELECTROFISHING SURVEY OF ILLINOIS River 361 


Aug., 1975 Sparks & STARRETT 


# 90°0 Lv 0 STO L110 It0 1) Si) # G20 1¢°0 Te°0 0F'0 00°0 90°0 
00°0 00°0 IT0 00°0 120 00°0 6o'0 00°0 00°0 

00°0 9T 0 00°0 00°0 #10 Tr'0 90°0 60°0 29°0 6TT cé0 00°0 Te°0 
00°0 00°0 9F°0 3r0 e'0 00°0 ot'0 00°0 IT 0 st 0 89°0 89°0 # 00°0 
620 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 

saynuipL OF 4ad swps6o0)1y 

T0°0 90°0 02°0 +10 90 60°0 Z10 Z0°0 110 Ge'0 92°0 ge0 400 oro 
00°0 00°0 €T0 00°0 oT0 00°0 ot 0 00°0 00°0 

00°0 ST 0 00°0 00°0 80°0 £80 80°0 90°0 9F°0 80T €¢°0 00°0 0g°0 
00°0 00°0 0*'0 0F'0 ££°0 00°0 620 00°0 10°0 90°0 6g°0 0s°0 1) att) 00°0 
0¢'0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 

SaInurpl 0G 42d 42QUNN 

81S G6t Gg ét 0°32 02 0°36 gTs 09d G&s G&s GPP 0-0T a & OCT 
PL6T €L6Tt OL6T 696T 896T L96T 996T S96T F96T €96T 96L T96T 096T 6S6T 


peysig sinoy {0 4aqunn pud vax 


vu Ul 

m0vVy 
esuely BT 
eBll08g 

YOOY PIaArIVys 
So] [1esIey 
juepseiqg 


7a TI 
uovLy 
esuely eT 
e1100q 


yooYy Peareys 
So][TeSsIe 


jWepseiq 


100d 


SS 


“pL61-6S61 ‘Aemuayeny siourjj| 24} ut 


Burysijosj2aja Aq uaxe} (4961 snqoir}) Oje43nq 42e}G—S1 FI9eL 


Vol. 31, Art. 8 


I_ttinois NaturAL History SuRvEY BULLETIN 


362 


80°0 90°0 200 # # # T0°0 200 T0'0 # Too 200 700 40'0 
£20 00°0 T0'0 00°0 00°0 00°0 00°0 00°0 00°0 

60'0 90°0 200 To'0 # 00°0 £0°0 400 # 00°0 00°0 100 90°0 
£0°0 $0°0 # # 00°0 00°0 00°0 To'0 20°0 # £0°0 £0°0 200 010 
00°0 £2°0 010 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 910 00°0 
00°0 00°0 00°0 00°0 00°0 400 # 400 00°0 T0'0 T0'0 00°0 00°0 00°0 
sT'0 0°0 00°0 40°0 


Sanur OF 42d swv160]1y 

92°0 g2°0 020 ¢0°0 400 60°0 10°0 LT0 10°0 400 80°0 g0°0 82°0 020 
09°0 00°0 &T0 00°0 00°0 00°0 00°0 00°0 00°0 

0&0 SsT0 0r0 80°0 80°0 00°0 LT0 tF0 80°0 00°0 400 62°0 G10 
oro $80 oT 0 L0°0 00°0 00°0 00°0 10°0 §T0 90°0 IT0 L0°0 1) 1) &T0 
00°0 L9°0 0&0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 020 00°0 08°0 00°0 
00°0 00°0 00°0 00°0 00°0 LT0 LT 0 9T 0 00°0 LT0 L0°0 00°0 00°0 00°0 
0L°0 £20 00°0 ny 

SOINUL OF 49d 42QUNN 


STs S'6T séT 0°66 0°3¢ 006 GTS 0°9¢ GES G&S i 2 0°0T Ger 0°CT 
PL6T €L6T OL6T 696T 896T L96T 9961 S96T P96T €96T G96T T96T O96T 6S6T 


paysig sinoy [0 4aQuUNN PUD 10aX 


vu TI 

uoily 

esueIy) eT 

BI 100g 

YoY peaie}s 
Sal [19s1e 
,uepseiqg 


zu Ud 
movLy 
esueIy) BT 
Bl100g 


WooY poaieys 
Sol[Iesiey 


,wepsaiq 


100d 


*pL61-6561 ‘Aemuazyeny sioujj] 243 ul Buiysijosjoaja Aq uaxe} (unyopidajossew ewoysoxoW) asi0upes peaywous—'9| 2/qe1 


ELECTROFISHING SURVEY OF ILLINOIS RIVER 363 


Aug., 1975 Sparks & STARRETT 


90°0 90°0 60°0 £0'0 600 400 
00°0 00°0 00°0 00'0 
00°0 00°0 00°0 # # 400 
T0'0 T0'0 20°0 T00 00°0 # 

00°0 00°0 00°0 00°0 00°0 60'0 
£50 S80 180 020 Ge'0 910 


400 
00°0 
a) 
£0°0 
00°0 
8T0 


300 
00°0 
T0'0 
# 

220 
420 


100 
00°0 
00°0 
# 

00°0 
90°0 


Sanur OF 4ad swmps6011y 


4 i ¥6°0 03°0 6£°0 T6°0 99°0 
00°0 00°0 00°0 00°0 
00°0 00°0 00°0 80°0 80°0 6r0 
0T0 10°0 0T0 90°0 00°0 &10 
00°0 00°0 00°0 00°0 00°0 £&°0 
09°6T 09°9 g's oss 099 og’ 


88°0 
00°0 
G10 
a) 
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ety 


T8°0 
00°0 
6T0 
&T0 
Soe 
Loy 


812 G61 get 066 036 0°32 
vL6T €L6T OL6T 696T 896T L96T 


GTe 


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6&0 
00°0 
00°0 
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00°0 
L1G 


SaINUL, OF 4ad 12QUNN 


GES 


p96T 


pays. sinoy [0 4aQuUNnN pup wax 


620 


1c°0 
+T 0 
62°0 


SPT 


zu TIT 

uoyLV 

esueIy eT 
BI100q 

yooy peaieig 
So] [los Ie yAy 
juepseiqg 


vu UL 
uovly 
esuBIy BT 


ell0eg 


WooY peaiej}s 
So[[los1e yy 


juepseig 


100d 


“PLO 1-656 ‘Aemuayen siourjj| ayy ul Busysiyosjoaja Aq uaye} (Sejou snanjeyr]) Peayljng 422|g— Z| F921 


Vol. 31, Art. 8 


Intivois NaturAL History SuRVEY BULLETIN 


364 


20'0 T0'0 00°0 T0°0 # # 200 # T0'0 # To'0 00°0 200 00°0 Paes a 10 
00°0 00°0 00°0 00°0 00°0 00°0 20°0 000 00°0 u0jTy 
£0°0 00°0 00°0 To'0 00°0 20°0 # 00°0 £00 T0'0 To'0 60°0 00°0 asuBiy ey 
£0'0 20'0 00°0 T0'0 10°0 # 10°0 200 To'0 00 20'0 00°0 0070 00°0 B1100g 
00°0 00°0 00°0 00°0 00°0 000 000 00°0 00°0 00°0 00°0 00°0 00°0 00°0 H00Y Peareis 
00°0 00°0 00°0 00°0 00'0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So][1es1e 
00°0 00°0 00'0 00°0 \Wepseiq 
Sanu, OF “aq swupsb6oNy 
10°0 £0°0 00°0 10°0 Z0°0 60'0 600 20°0 80°0 400 90°0 00°0 020 00°0 vu ‘Ill 
00°0 00°0 00°0 00°0 00°0 00°0 210 00°0 00°0 u0z1V 
010 00°0 00°0 80°0 00'0 80°0 80°0 00°0 cT'0 80'0 40°0 ZL'0 00°0 asuBiy eT 
010 10°0 00°0 810 10°0 10°0 Tz'0 10°0 90°0 90°0 sT'0 00°0 00°0 00°0 el100g 
00°0 00°0 00°0 00°0 00°0 00'0 00°0 00°0 00°0 0070 00°0 00°0 00°0 00°0 400 peareis 
00°0 00°0 00°0 00°0 000 00°0 00°0 000 00°0 00°0 00°0 000 00°0 0070 So] [Tes1ey 
00°0 00°0 00°0 00°0 \wepseiq 


SaInurp OF 49d 42QUNnN 
oo EE i ON rt oS de les 6 ee a i ee ee Eee 
812 G6T Get 032 03 032 big 4 0°92 Gets Gs AZ 00T Gor OCT 100d 
PLET €L6. OL6T 696T 896T L96T 996T G96T $96T 961 96T T96T 096T 6S6T 


peysig sinoy [0 4aQUNN PUD 1D2ax 
a ——— ————————————————_ 


"bL61-6S61 ‘Aemuayeny S}our|] 243 U! Burysiyoajoaja Aq aye} (syeseU Smanjeyr]) PEAYyING MOj}2A—'BL F921 


ELECTROFISHING SURVEY OF ILLINOIS RIvER 365 


Aug., 1975 Sparks & STARRETT 


g9°0 T8°0 &T0 E00 62°0 $20 st0 60°0 020 89°0 92°0 100 8T0 00°0 ras i 10 6 
T6'T 06°0 80 89°0 LYT g92°0 TOT ta ad 88°0 u0vTV 
vF'0 9F°0 rr0 G20 SPO C¥'0 820 10°0 oT 0 610 Gt0 Lo°0 00°0 eduBID BT 
610 120 00°0 T'0 s0°0 400 100 100 00°0 00°0 L0°0 600 £0°0 00°0 BIL08d 
00°0 99°0 60°0 00°0 00°0 00°0 00°0 LT0 00°0 00°0 rE0 00°0 s0°0 00°0 HOY PI9aAsezs 
6T'0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So] [los1e Ay 
00°0 00°0 00°0 00°0 duepseid 
Sanu, OF 4ad Swvs60)14 
PLT 99°0 0r0 SOT o9'T 00°T Set +90 Ts'0 61% 96°0 oro 2 T 00°0 vu Ud 
oes Gls GLY €9'T 00°¢ 00% Les 0s'8 0st moV 
Oe T 690 Ort ort 19's ors 19's G20 180 69'S 9FT 00°F 00°0 esueiy BT 
0r0 0+0 00°0 Lv0 020 10 40°0 &T0 00°0 00°0 6T 0 +10 02°0 00°0 Bl1l09d 
00°0 00% 0t°0 00°0 00°0 00°0 $80 Get 00°0 00°0 010 00°0 02°0 00°0 HOY P9areys 
02°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 00°0 So] [Tos1e 
00°0 00°0 00°0 00°0 uepseid 


Sanur OF 4d 42QUNN 


812 9'6T GT 0°32 0°32 0°33 G12 0°92 GES Gs GPP 0'0T Ger OCT 100g 
PL6T €L6T OL6T 6961 896T L96T 996T S96T v96T €96T 696T T96T 096T 696T 


pays sinoy fo 4aqunn pup vax 


*pL61-6S61 ‘Aemuayeny stout] 243 ul Gusysiyosj9aj9 Aq uaye} (Smgegound snanjeyry) Ysiy3eD jauUeYD—"6| 2/42) 


Vol. 31, Art. 8 


Inuino1is NATURAL History SURVEY BULLETIN 


366 


180 200 2r0 # 100 # 00°0 00°0 00°0 90°0 # 00°0 00°0 00°0 cu UI 
00 200 90°0 T0°0 0070 00°0 00°0 100 400 mow 
80°T g0°0 SPT 00°0 0070 00°0 00°0 00°0 0070 120 00°0 00°0 00°0 asUBIN VT 
00°0 00°0 0070 00°0 00°0 00°0 00°0 0070 00°0 000 00°0 00°0 00°0 0070 B1100d 
00'0 00°0 00°0 00°0 00°0 000 00°0 00°0 00°0 00°0 00°0 0070 00°0 00°0 qooy peains 
00°0 000 00°0 00°0 00'0 00°0 000 00°0 0070 00°0 00°0 00°0 0070 00°0 so] [}esre 
00°0 00°0 00°0 0070 \depseiq 
SaynupL OF 4aq swmp160)14 
62°0 £0°0 0T'0 300 ¢0°0 30°0 00°0 00°0 00°0 90'0 400 00°0 00°0 0070 vu Ul 
03°0 92'0 20 92'0 00°0 0070 00°0 rm) 08°0 tory 
09°0 80°0 0F'0 00°0 0070 00°0 00°0 00°0 00°0 gT'0 00°0 00°0 00°0 aguvID eT 
00°0 00°0 0070 00°0 00'0 00°0 00°0 0070 00°0 00°0 00°0 00°0 00°0 00°0 e1100q 
00°0 00°0 0070 00'0 00°0 00°0 00°0 00°0 00°0 0070 00°0 00°0 00°0 00°0 yoy peareis 
00°0 000 00°0 00°0 00°0 0070 00°0 0070 00°0 00°0 00°0 00°0 00°0 00°0 so]]1os1eW 
00°0 00°0 00°0 00°0 \depseiq 


SaInurp. OF 42d 42QUNN 


81s g'6T g'éT ad 0°06 ad GTZ 0°92 Gs i 4 GHP 00T Ger OCr 100d 
PL6T €L6T OL6T 696T 8961 L96T 996T G96T v96T £961 696T T96T 096T 6S6T 


poysig sinoy {0 4aqQunn pup LDax 


a 
—EEEoEEoEoESESSSSSSESEEEEEooooooooooooeeeEeSESESEESSESeSeEeeoeeEeESESaEaSaSaSaESaSaESEESSESESESESESSaaaSSSaSESESESaSQSQooana SESS ")§/:0—0°00—5——5—™— 


"bL6 1-656 1 ‘Aemuajeny Siour|] 243 ul Burysijo1yo99 Aq ures (SHEANO SHD>POJAg) YsI1¥Je> Peayie|j— OZ P1921 


ELECTROFISHING SURVEY OF ILLINOIS RIvER 367 


Aug., 1975 Sparxs & STARRETT 


€2°0 $00 ST'0 S20 of 0 120 810 ST 0 10°0 STO 80 00°0 s0°0 00°0 
0s°0 tT T 8eT 290 40 G20 92°0 08°0 €0T 00°0 
oT 0 # 12°0 00°0 oT'0 02°0 02°0 st 0 00°0 90°0 60°0 &1T0 00°0 
120 S00 9T'0 oO 610 120 92°0 ST 0 10°0 00°0 oro 00°0 60°0 00°0 
80°0 90°0 10°0 00°0 00°0 orto 120 10°0 00°0 00°0 90°0 00°0 # 00°0 
60°0 100 # 00°0 00°0 00°0 00°0 # 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 
Sanur OF 4aq swo.borLy 
IPT 9¢°0 09°0 02'T +80 LL'0 LL0 00°T 0*'0 890 8L'0 00°0 910 00°0 
OL'T GLs 6F'E 00% 00'T LOT ooT 69% Orr 00°0 
03°0 80°0 08°0 00°0 G30 890 gL'0 +60 ST 0 90 £e°0 62°0 00°0 
02% €6°0 0st L¥0 0*'0 99°0 £60 90°T G20 00°0 rr0 00°0 oro 00°0 
09°0 ee 1 OL'T 00°0 00°0 €¢°0 00°T GL0 00°0 00°0 02°0 00°0 02°0 00°0 
0F'0 02°0 02°0 00°0 00°0 00'0 00°0 LT 0 00°0 00°0 00°0 00°0 00°0 00°0 
00°0 00°0 00°0 00°0 
SaINuUL OF 42d 49QUNN 
81s S6T Gét 0°03 0°66 086 gTs 0°93 Gs Gs GHP 00T Ger OCT 
PLE €L61 OL6T 6961 8961 L961 9961 S961 F961 £961 C96T T96T 0961 6S6T 


paysigq sino fo 4aqunyn pup vax 


zu dl 

u0j;TV 
esuely eT 
Bl100q 

YooYy poarejs 
So][Ios1eyy 
,Wepseiq 


aes 00 E 
movLy 
esuely eT 
Bl100q 


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so] [1es1e 


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100d 


0.30—00O“—oowowr"“"AaOWwqSASSSeSsSSeeeeee 


‘pL61-6S61 ‘Aemsiayeny sour) a4} ul BusysijosjDa}9 Aq uae} (sdosAay> @uosow) sseq aHUA\—' |Z FIGeL 


Vol. 31, Art. 8 


Intinois NaturAL History SURVEY BULLETIN 


368 


ee ee SS 


90°0 £10 40'0 110 90°0 £0'0 400 # # T0'0 20°0 £0°0 800 T0'0 zu Il 
40°0 Z0°0 00°0 # T0°0 000 ag # # u071V 
TS'0 010 0'0 020 10°0 T0'0 To'0 # To'0 T0'0 £0'0 00°0 asuBiy BT 
ya) 810 ¢0'0 +10 010 90°0 600 # # £0'0 £0°0 £0'0 00°0 400 el10ed 
600 19 a1) 00°0 ¢0'0 800 T0'0 To'0 00°0 00°0 # # 00°0 00°0 00°0 YoY peaivis 
60°0 S00 G0'0 # To'0 200 # 000 00°0 000 # £0°0 # # So] [losreW 
00°0 # 00'0 # \wepseiqg 
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Vol. 31, Art. 8 


Inuiwois NaTuRAL History SURVEY BULLETIN 


370 


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Vol. 31, Art. 8 


Intiwois NatTurAL History SURVEY BULLETIN 


372 


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Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RIVER 375 


' Table 29.—Average number of kilograms of fish taken per 30 minutes of electrofishing in 
2ach navigation pool of the Illinois Waterway during the period 1959-1974. 


Pools 

Ref. Downstream Upstream 
Table La Starved Mar- Dres- 
No. Species Alton Grange Peoria Rock seilles den 
3 Shortnose gar 0.01 0.04 0.04° 0.00 0.00 0.00 
4 Bowfin 0.05% 0.05 0.01 0.00 0.00 0.00 
5 Gizzard shad 0.36 0.88 0.902 0.40 0.47 0.03 
6 Goldeye 0.028 # # 0.00 # 0.00 
7 Mooneye 0.018 0.00 # 0.00 0.00 0.00 
8 Goldfish 0.00 0.04 0.05 2.38 2.10 3.05 
9 Carp x goldfish # 0.05 0.38 Hesiy(e 0.53 0.35 
10 Carp 19.01° 17.40 8.02 9.24 10.85 5.82 
11 River carpsucker 0.10 0.12 0.10 0.148 0.02 0.00 
12 Quillback carpsucker 0.03 0.03 0.14 0.508 0.20 0.00 
13 Smallmouth buffalo 0.03 0.52" 0.34 0.17 # 0.00 
14 Bigmouth buffalo 0.48 4.21 5.70" 0.02 # 0.00 
15 Black buffalo 0.06 0.25* 0.20 0.01 0.00 0.00 
16 Shorthead redhorse 0.03 0.02 0.02 0.04 0.01 0.06" 
17 Black bullhead 0.00 # 0.05 0.04 0.248 0.00 
18 Yellow bullhead # 0.01 0.01* 0.00. 0.00 0.00 
19 Channel catfish 1.122 0.36 0.07 0.09 0.01 0.00 
20 Flathead catfish 0.03 0.21° 0.00 0.00 0.00 0.00 
21 White bass 0.672 0.10 0.11 0.05 0.01 0.00 
22 Green sunfish 0.01 0.09% 0.06 0.02 0.02 # 
23 Bluegill 0.15 0.20" 0.06 # 0.01 0.00 
24 Largemouth bass 0.44 Table 0.78 0.04 0.08 0.00 
25 White crappie 0.19 0.20 0.23 0.01 0.03 0.00 
26 Black crappie 0.44 0.65% 0.43 0.03 # 0.00 
27 Freshwater drum 0.12 0.29% 0.05 0.01 # 0.00 


8 Indicates the pool or pools where the maximum number of kilograms of each species was taken 
in the period 1959-1974. 


# Less than 0.01 kilogram taken, 


376 Inuinois NaturAL History SURVEY BULLETIN Vol. 31, Art. 8 


Table 30.—Average number of fish taken per 30 minutes of electrofishing in each naviga- | 
tion pool of the Illinois Waterway during the period 1959-1974. 


Pools 

Ref. Downstream Upstream 

Table La Starved Mar- Dres- 
No. Species Alton Grange Peoria Rock seilles den 
3 Shortnose gar 0.072 0.078 0.05 0.00 0.00 0.00 — 
4 Bowfin 0.07% 0.02 0.01 0.00 0.00 0.00 
5 Gizzard shad 18.09 43.55 63.20% 9.22 12.52 2.66 
6 Goldeye 0.41" 0.02 0.02 0.00 0.03 0.00 
7 Mooneye 0.052 0.00 0.01 0.00 0.00 0.00 
8 Goldfish 0.00 0.37 0.73 17.02 12.02 42.76" 
9 Carp x goldfish 0.01 0.18 1.39 2.028 0.80 1.05 
10 Carp 19.81 34.692 18.67 18.92 14.29 12.06 
11 River carpsucker 0.27 0.39 0.449 0.34 0.04 0.00 
12 Quillback carpsucker 0.11 0.16 0.52 1.38° 0.71 0.00 
13 Smallmouth buffalo 0.11 1.04 0.67 0.25 0.03 0.00 
14 Bigmouth buffalo 0.33 4.21 5.798 0.05 0.01 0.00 
15 Black buffalo 0.04 0.242 0.19 0.02 0.00 0.00 
16 Shorthead redhorse 0.08 0.21 0.09 0.02 0.00 0.00 
17 Black bullhead 0.00 0.12 0.35 0.38 4.398 0.00 
18 Yellow bullhead 0.01 0.10" 0.07 0.00 0.00 0.00 
19 Channel catfish 3.76% 1.60 0.17 0.34 0.01 0.00 
20 Flathead catfish 0.192 0.09 0.00 0.00 0.00 0.00 
21 White bass 2.65" 0.42 0.64 0.44 0.07 0.00 
22 Green sunfish 0.52 1.77 2.918 0.65 1.01 0.23 
23 Bluegill 4.90 7.12 3.10 0.06 0.33 0.00 
24 Largemouth bass 1.70 4.238 2.82 0.13 0.47 0.00 
25 White crappie 1.07 1.752 1.47 0.05 0.32 0.00, 
26 Black crappie 2.99 5.55% 2.44 0.18 0.04 0.00 
27 Freshwater drum 1.49 2.498 0.34 0.03 0.14 0.00: 


"Indicates the pool or pools where the maximum number of individuals of each species was 
taken in the period 1959-1974, 


LITERATURE CITED 


Buck, D. H. 1956. Effects of turbidity on 
fish and fishing. Twenty-First North 
American Wildlife Conference Transac- 
tions: 249-261. 


Burts, T. A. 1974. Measurements of sedi- 
ment oxygen demand characteristics of 
the upper Illinois Waterway. Report of 
Investigation 76. Illinois State Water 
Survey. 32 p. 


1975. Nitrification effects on the 
dissolved oxygen resources of the Illinois 
Waterway. In: Water—1974: II. Mu- 
nicipal Wastewater Treatment. American 
Institute of Chemical Engineers Sympo- 
sium Series 71(145) :38-43. 


Cartson, A. R., and R. E. Sierert. 1974. 
Effects of reduced oxygen on the embryos 
and larvae of lake trout (Salvelinus 
namaycush) and largemouth bass (Mi- 
cropterus salmoides). Journal of the 
Fisheries Research Board of Canada 
31(8) :1393-1396. 


Forses, S. A. 1928. Foreword, p. 387-388. 
In: R. E. Richardson. The bottom fauna 
of the Middle Illinois River, 1913-1915. 
Illinois Natural History Survey Bulletin 
17(12) :387—475. 


, and R. E. RicHarpson. 1913. Stud- 
ies on the biology of the upper Illinois 
River. Illinois State Laboratory of Nat- 
ural History Bulletin 9(10):481-574, 21 
plates. 


, and 1919. Some recent 
changes in Illinois River biology. Illinois 
Natural History Survey Bulletin 13(6): 
139-156. 


, and . 1920. The fishes of Illi- 
nois. Second ed. Illinois Natural History 
Survey. cxxxvi + 357 p. 


GaLe, W. F. 1969. Bottom fauna of Pool 19, 
Mississippi River, with emphasis on the 
life history of the fingernail clam, Sphaer- 
ium transversum, PhD dissertation. Iowa 
State University. Ames, Iowa. 234 p. 


1971. An experiment to determine 
substrate preference of the fingernail 
clain, Sphaerium transversum (Say). 
Ecology 52 (2) :367-370. 


Jackson, H. O., and W. C. Srarrerr. 1959. 
Turbidity and sedimentation at Lake 
Chautauqua, Illinois. Journal of Wild- 
life Management 23(2) :157-168. 


Larimorg, R. W. 1961. Fish population and 
electrofishing success in a warm-water 
stream. Journal of Wildlife Management 
25(1) :1-12, 


LusinskI, K. S., R. E. Sparks, and L. A. 
JAHN. 1974. The development of tox- 
icity indices for assessing the quality of 
the Illinois River. Research Report No. 
96. Water Resources Center, University 
of Illinois at Urbana-Champaign. 46 p. 


Mitts, H. B., W. C. Starrett, and F. C. 
BELLROSE. 1966. Man’s effect on the fish 
and wildlife of the Illinois River. Illinois 
Natural History Survey Biological Notes 
No. 57. 24 p. 


NeEtson, E. W. 1878. Fisheries of Chicago 
and vicinity. In: Report of the U.S. 
Commissioner of Fish and Fisheries for 
1875-1876, Part 4, Appendix B, p. 783-800. 


O’DonneELL, J. D. 1935. Annotated list of 
the fishes of Illinois. Illinois Natural 
History Survey Bulletin 20(5) :473-500. 


Patoumpis, A. A., and W. C. STARRETT. 
1960. An ecological study of benthic or- 
ganisms in three Illinois River flood plain 
lakes. American Midland Naturalist 64 
(2) :406—435. 


RicHarpson, R. EB. 1921a. The small bottom 
and shore fauna of the Middle and Lower 
Illinois River and its connecting lakes, 
Chillicothe to Grafton: its valuation; 
its sources of food supply; and its rela- 
tion to the fishery. Illinois Natural His- 
tory Survey Bulletin 13(15) :363-522. 


1921b. Changes in the bottom and 
shore fauna of the middle Illinois River 
and its connecting lakes since 1913-1915 
as a result of the increase, southward, of 
sewage pollution. Illinois Natural His- 
tory Survey Bulletin 14(4) :33-75. 


. 1928. The bottom fauna of the mid- 
dle Illinois River, 1913-1925, its distribu- 
tion, abundance, valuation, and index 
value in the study of stream pollution. 
Illinois Natural History Survey Bulletin 
17(12) :387-475. 


SincH, K. P., and J. B. Stati. 1973. The 
7-day, 10-year low flows of Illinois 
Streams. Illinois State Water Survey 
Bulletin 57. 


Sra, J. B., and S. W. Metstep. 1951. The 
silting of Lake Chautauqua, Havana, IIli- 
nois. Illinois State Water Survey, in co- 
operation with Illinois Agricultural Ex- 
periment Station, Report of Investigation 
8. 15 p. 


Srarrett, W. C. 1971. A survey of the 
mussels (Unionacea) of the Illinois Riv- 
er: a polluted stream. Illinois Natural 
History Survey Bulletin 30(5): 267-403. 


377 


. 
378 ILLinois NaTurAL History SuRvEY BuLLETIN Vol. 31, Art. 8° 


———. 1972. Man and the Illinois River, tauqua, Illinois. Illinois Natural History 
p. 131-169. In: R. T. Oglesby, C. A. Carl- Survey Bulletin 29(1):1-104. 


son, and J. A. McCann (eds.). River THompson, D. H. 1928. The “Knotheaa” 
ecology and man. Proceedings of an In- carp of the Illinois River. Illinois Nat- 


ternational Symposium on River Ecology —yral History Survey Bulletin 17(8):285- 
and the Impact of Man, held at the Uni- 320. 


versity of Massachusetts, Amherst, Mas- 

sachusetts, June 20-23, 1971. Academic U. S. Army ENGINEER District, CuHrcaco, 

Press. New York. 465 p. 1970. Charts of the Illinois Waterway 
from Mississippi River at Grafton, IIli- 

, and A. W. Frirz. 1965. A biological nois to Lake Michigan at Chicago and 


investigation of the fishes of Lake Chau- Calumet Harbors. 77 p. 
INDEX 
A Chicago River, 332, 333 
Alosa chrysochloris, 331 arya Sanitary and Ship Canal, 318, 333, 
rene oe Geese Island Chute, 236 345 
mre Had S19; 822, S26) S20 iba Ks SORE, Gimnere lat nsufente 321, 322, 326, 329, 331, 


333, 341, 344, 345, 374 


ee aU Ronde Bet MR SRT Ae Commercial fishermen, 321, 322, 326, 331, 


American eel (see Anguilla rostrata) 


5 341, 344 
=o) PAI ECNERE SE Commercial fishery, 317, 326, 329, 345 
ammonia, 334, d8b, 342, 344 Commercial river traffic (see navigation) 
Anguilla rostrata, 331, 332 C a Greek as vie = 
Aplodinotus grunniens, 322, 331, 334, 345, Opperas , 
373 Copperas Creek Dam, 337 


Corbicula manilensis, 344 
Crappie (see Pomozis) 
Otenopharyngodon idella, 344 


Asiatic clam (see Corbicula manilensis) 


B Current, 337, 338, 344 
Ballard Island Chute, 319, 326, 345 Cyprinus carpio, 322, 325, 326, 333-335, 344, 
Barges (see navigation) 345, 355, 356 
Bath Chute, 335, 344 
Big Blue Island Chute, 318, 331 D 


Bigmouth buffalo (see Ictiobus cyprinellus) 3 A 

Black buffalo (see Ictiobus niger) Des Plaines on 318, 322, 329, 345 

Black bullhead (see Ictaluwrus melas) Detweiller Park, 318 

Black crappie (see Pomozis Discharge (river flow), 335 
nigromaculatus) Dorosoma cepedianum, 322, 326, 345, 351 


Bluegill (see Lepomis macrochirus) Dresden Dam, 318 


Boat traffic (see navigation) Dresden Heights, 337 


Dresden Pool, 318, 319, 321, 326, 345 
Bottoml kes, , , , , , 2 7 : : , 
eee lakes, 317, 336, 337, 339, 340 Drought effects, 329, 341 


Ducks, 334, 335, 342 


Bowfin (see Amia calva) Du Page River, 318, 332 


Brown bullhead (see Ictalurus nebulosus) 


Bullheads (see Ictalurus) i 
E ; 
Cc Electrofishing, 317, 321, 322, 332, 347, 375, 
Carassius auratus, 322, 323, 324, 344, 345, 376 
354, 355 Eel (sée Anguilla rostrata) 
Carp (see Cyprinus carpio) Emerald shiner (see Notropis atherinoides) 
Carp x goldfish hybrids (see Carassius Esox lucius, 331, 332, 345 
auratus) Exotic species (see introduced species) 
Carpiodes 
carpio, 322, 326, 345, 357 
cyprinus, 326, 358 F 
Catfishes (see also Ictalurus), 332, 339, 340 Fingernail clams (see Sphaertidae) 
Catostomus, 332 Fish (see names of species, commercial 
Channel catfish (see Ictalurus punctatus) fish, sport fish, etc.) 
Chicago, 317, 332, 335, 342 Fisherman’s Special (train between 
Chicago Metropolitan Sanitary District, 342, Springfield and Havana), 317 


343 Flathead catfish (see Pylodictis olivaris) 


Aug., 1975 Sparks & STARRETT: ELECTROFISHING SURVEY OF ILLINOIS RiveR 379 


Food organisms (see also Sphaeriidae), 
340, 341, 345, 346 

Freshwater drum (see Aplodinotus 
grunniens ) 


G 


Game fish (see sport fish) 

Gar (see Lepisosteus platostomus) 

Gizzard shad (see Dorosoma cepedianum) 

Godar Landing, 326 

Goldeye (see Hiodon alosoides) 

Goldfish (see Carassius auratus) 

Goldfish x carp hybrids (see Carassius 
auratus ) 

Grafton, 318, 337 

Green sunfish (see Lepomis cyanellus) 


H 


Habitat, 318, 332, 335, 344, 345 
brush piles, 318, 331 
degradation by pollution, 335, 339, 
340-345 
increase due to high water, 334, 335 
loss by leveeing, 335, 337, 345 
sampling, 318 
Hardin, 337 
Havana, 317, 331, 339 
Hennepin, 319, 332 
Henry, 333 
Herbicides, 344 
Hickory Creek, 332 
Hiodon alosoides, 331, 345, 352 
Hiodon tergisus, 331, 345, 353 
Historical background of Illinois River, 
317, 332-341 
Hoop nets, 317 
Hybrids (see Carassius duratus) 
Hybrid vigor, 322 


Ictalurus, 334 
catus, 331, 344 
melas, 326, 327, 345, 363 
natalis, 331, 345, 364 
punctatus, 326, 328, 329, 339, 341, 345, 
365 
Ictiobus, 332, 334, 345 
bubalus, 326, 345, 359 
cyprinellus, 326, 341, 345, 360 
niger, 326, 341, 345, 361 
Illinois-Michigan Canal, 332, 333 
Illinois River 
description, 318 
historical background, 317, 332-341 
lower section, 318, 319, 329, 331, 334, 344 
middle section, 319, 334, 338 
navigation pools, 318 
upper section, 319, 321, 322, 332, 344 
valley, 317 
Introduced species (see Carassius auratus, 
Corbicula manilensis, Ctenopharyngodon 
idella, Cyprinus carpio, Ictalurus catus) 


K 


Kampsville, 333, 344 
Kampsville Landing, 326 
Kankakee River, 318 
Kingston Mines, 342 


L 


LaGrange, 333, 337 
LaGrange Dam, 318, 337 
LaGrange Pool, 318, 319, 322, 326, 329, 331, 
335, 344, 345 
Lake Chautauqua, 331, 339 
Lake Michigan, 333, 334, 337, 342, 343 
Lake restoration, 342, 344 
Lamprey, 332 
Largemouth bass (see Micropterus 
salmoides ) 
LaSalle, 333 
Lepisosteus 
oculatus, 331 
osseus, 332 
platostomus, 321, 322, 345, 349 
Lepomis, 334 
cyanellus, 329, 345, 368 
gibbosus, 331 
humilis, 331 
macrochirus, 329, 339, 341, 345, 369 
megalotis, 331 
microlophus, 339 
Longnose gar (see Lepisosteus osseus) 
Longear sunfish (see Lepomis megalotis) 


Marquette, 317 
Marseilles, 333, 337 
Marseilles Dam, 318, 331 
Marseilles Pool, 318, 319, 321, 326, 329, 331, 
341, 345 

Matanzas Beach, 342 
Matanzas Lake, 342 
Meredosia, 317, 342 
Micropterus, 334 

dolomieui, 331 

salmoides, 322, 329, 330, 332, 335, 339, 

344, 345, 370 
Minnows (see also Notropis), 331, 332 
Minnow seines, 317 
Mississippi River, 318, 341, 344 
Molluscs (see also Sphaeriidae), 322, 334, 
340 

Mooneye (see Hiodon tergisus) 
Morone chrysops, 329, 332, 341, 367 
Mortland Island Chute, 318 
Mozostoma macrolepidotum, 331, 332, 362 


Navigation, 342-344 
effects on aquatic life, 341 
effects on dissolved oxygen, 320, 321, 343 
effects on turbidity, 319, 320, 340-344 
channel, 318, 342 
channel dredging, 342, 343 


380 


dams, 318, 333, 337, 342, 343 
pools, 318, 337 
Northern pike (see Esox lucius) 
Notropis atherinoides, 332 


Oo 


Orangespotted sunfish (see Lepomis 
humilis) 
Oxygen 
dissolved, 319-321, 333-339, 341-345, 348 
demand, 320, 321, 334, 337, 342-345 


P 
Pekin (see also Peoria-Pekin), 318, 341 
Peoria, 337 
Peoria Dam, 318 
Peoria Lake, 318, 319, 331 
Peoria-Pekin metropolitan area, 342 
Peoria Pool, 318, 319, 321, 322, 326, 329, 331, 
335, 344, 345 
Perca flavescens, 340, 345 
Perches (see also Perca, Stizostedion), 337 
Pesticides, 340 
Pikes (see also Hsoz), 337 
Plankton, 319, 334, 345 
Plants (aquatic), 340, 342, 343, 344, 345, 346 
Pollution (see also ammonia, oxygen, 
pesticides, sediment, toxic 
chemicals, turbidity), 345 
agricultural, 339, 340, 343, 344 
industrial, 333, 335, 344 
municipal (sewage), 332-335, 337-341, 
344 
Pomozis, 322, 334, 340, 341 
annularis, 329, 331, 341, 345, 371 
nigromaculatus, 329, 331, 341, 345, 372 
Pumpkinseed (see Lepomis gibbosus) 
Pylodictis olivaris, 329, 339, 341, 366 


Q 


Quillback (see Carpiodes cyprinus) 
Quiver Lake, 339, 342 


Recovery from pollution (see also lake 
restoration), 3438, 345 

Redear sunfish (see Lepomis microlophus) 

Refuges, 343 

Restoration, 342, 343, 345 

Rice Lake, 342 

River (see Illinois River, Kankakee River, 
etc.) 

River carpsucker (see Carpiodes carpio) 

River redhorse (see Moxostoma carinatum ) 

Rock bass (see Ambloplites rupestris) 


Sampling method, 318, 319 
Sauger (see Stizostedion canadense) 


Intinois Natura History SuRVEY BULLETIN 


Vol. 31, Art. 8 


Sediment, 319-321, 333, 337, 340, 342, 343, 
345 

Seneca, 333 

Shortnose gar (see Lepisostus 
platostomus) 

Shorthead redhorse (see Mozostoma 
macrolepidotum ) 

Siltation (see sediment) 

Skipjack herring (see Alosa chrysochloris) 

Smallmouth bass (see Micropterus 
dolomieui) : 

Smallmouth buffalo (see Ictiobus bubalus) 

Soil conservation, 343 j 

Sphaeriidae, 334, 335, 340 

Sport fish (es), 317, 329, 331, 339-341, 345 

Sport fishermen, 331 

Spotted gar (see Lepisosteus oculatus) 

Starved Rock, 337 

Starved Rock Dam, 318 

Starved Rock Pool, 318, 319, 321, 326, 329, 
341, 345 

St. David, 342 

Stizostedion 

canadense, 331 
vitreum vitrewm, 332, 345 

Stump Lake, 342 

Suckers (see Catostomus, Moxostoma) 

Sunfishes (see also Lepomis, Micropterus, — 
Pomozcis), 332, 334, 337 


T 


Temperature, 321, 348 

Towboats (see navigation) 

Toxic chemicals, 335, 340, 341 
Trawling, 326, 329 

Turbidity, 319, 320, 333, 339, 340, 342 
Turkey Island Chute, 331, 344 


U 


U.S. Government 
Corps of Engineers, 318 
fisheries station, 317 q 


Ww 


Walleye (see Stizostedion vitrewm vitreum) 
Water levels 

effects on fishes and other organisms, 

329, 331, 334, 335-339, 341, 343 

effects on sampling, 319 q 
White amur (see Ctenopharyngodon idella) 
White bass (see Morone chrysops) 
White catfish (see Ictalurus catus) 
White crappie (see Pomozis annularis) 


Y 


Yellow bullhead (see Ictalurus natalis) 
Yellow perch (see Perca flavescens) 


Some Publications of the ILLINOIS NATURAL HISTORY SUR 


BULLETIN 


Volume 31, Article 2—The Reproductive 
Cycle of the Raccoon in Illinois. By Glen 
C. Sanderson and A. V. Nalbandoy. July, 
1973. 57 p., index. 


Volume 31, Article 3—Nutritional Respon- 
ses of Pheasants to Corn, with Special 
Reference to High-Lysine Corn. By Ron- 
ald F. Labisky and William L. Anderson. 
July, 1973. 26 p., index. 


Volume 31, Article 4—An Urban Epiphy- 
totic of Phloem Necrosis and Dutch Elm 
Disease, 1944-1972. By J. Gedric Carter 
and Lucile Rogers Carter. May, 1974. 31 
p., index. 


Volume 31, Article 5.—Larvae of the Seri- 
cothripini (Thysanoptera: Thripidae), 
with Reference to Other Larvae of the 
Terebrantia, of Illinois. By Thomas C. 
Vance. August, 1974. 64 p., index. 


Volume 31, Article 6——Root Infection of 
Woody Hosts with Verticillium albo- 
atrum. By Gerald L. Born. August, 1974. 
41 p., index. 


Volume 31, Article 7—-The Mecoptera, or 
Scorpionflies, of Illinois. By Donald W. 
Webb, Norman D. Penny, and John C. 
Marlin. August, 1975. 66 p., index. 


BIOLOGICAL NOTES 


83.—Illinois Birds: Laniidae. By Richard 
R. Graber, Jean W. Graber, and Ethelyn 
L. Kirk, June, 1973. 18 p. 


84— Interactions of Intensive Cultures of 
Channel Catfish with Largemouth Bass in 
1-Acre Ponds. By D. Homer Buck, Rich- 
ard J, Baur, and C. Russell Rose. Febru- 
ary, 1974. 8 p. 


85.—The Literature of Arthropods Associ- 
ated with Soybeans. III. A Bibliography 
of the Bean Leaf Beetle, Cerotoma trifur- 
cata (Forster) and C. ruficornis (Olivier) 
(Coleoptera: Chrysomelidae). By M. P. 
Nichols, M. Kogan, and G. P, Waldbauer. 
February, 1974. 16 p. 


86.—Illinois Birds: Tyrannidae. By Rich- 
ard R. Graber, Jean W. Graber, and 
Ethelyn L. Kirk. February, 1974. 56 p. 


87—The Literature of Arthropods Associ- 
ated with Alfalfa. I. A Bibliography of 
the Spotted Alfalfa Aphid, Therioaphis 
maculata (Buckton) (Homoptera: Aphi- 


List of available publications mailed on request 


No charge is made for publications of the ILL1no1is NATuRAL History SuRvVEY. 
copy of most publications will be sent free to anyone requesting it until the supply be 
low. Costly publications, more than one copy of a publication, and publications in | 
supply are subjects for special correspondence. Such correspondence should iden! 
writer and explain the use to be made of the publication or publications. 


Address orders and correspondence to the Chief 
illinois Natural History Survey 


dae). By D. W. Davis, M. P. N 8, 
E. J. Armbrust. February, 1974. 1: 


88.—The Literature of Arthropods 
ated with Alfalfa, II. A Bibliog 
the Sitona Species (Coleoptera: — 
lionidae). By W. P. Morrison, B, C 
M. P. Nichols, and E. J. Armbru 
ruary, 1974. 24 p. 


89.—The Life History of the Spot 
er, Etheostoma squamiceps, in 
Illinois, and Ferguson Creek, K er 
By Lawrence M. Page. May, 197: 

90.—A Bibliography of the Nort! 
Rootworm, Diabrotica longico 4 
and the Western Corn Rootwor 
brotica virgifera LeConte (Co 
Chrysomelidae). By W. H. Lue 
H. C. Chiang, E. E. Ortman, and 
P. Nichols. April, 1974. 15 p. 

91.—The Distribution of Period 
in Illinois. By Lewis J. Sta 
February, 1975. 12 p. 


92.—The Literature of Arthropod: 
ated with Soybeans. IV. A Biblio 
of the Velvetbean Caterpillar 4 
gemmatalis Hiibner (Lepidop' 
tuidae). By B. J. Ford, J. R. 
Reid, and G. L. Godfrey. Febr 
15 p. 

93.—The Life History of the 
Darter, Etheostoma kennicotti, 
Creek, Illinois. By Lawrence 
February, 1975. 15 p. 


94.—Illinois Pheasants: Their Distr! 
and Abundance, 1958-1973. By R 
Labisky. February, 1975. 11 p. 
95.—The Nest Biology of the Bee 
(Ptilandrena) erigeniae Robe 
menoptera: Andrenidae). By 
Davis, Jr. and Wallace E. LaBerg 
1975. 16 p. 


CIRCULAR th 

51.—Illinois Trees: Selection, Plant 
Care, By J. Cedric Carter. Aug 
123 p. 


52.—Fertilizing and Watering T 


54.—Corn Rootworm Pest Manager 
Canning Sweet Corn. By W. H 
mann, J. T. Shaw, D. E. Kuh 
Randell, and C. D. LeSar. Mar 
10 p. 


I ILLINOIS 
atural History Survey 
BULLETIN 


Pesticides and 
Environmental Quality 
in Illinois 


t L. Metcalf 


mel 
; R. Sanborn arunnt 08 ; 
4 \ 

oot 

yBRARY 

F ILLINOIS Hi LIBRARY OF THE 
TMENT OF REGISTRATION AND EDUCATION 
¥ DET 31375 


RAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 


Jr. at Y Ub ILLINOIS 
AT URBANA-CHAMPAIGN 


VOLUME 31, ARTICLE 9 
AUGUST, 1975 


ILLINOIS 


latural History Survey 
BULLETIN 


Pesticides and 
Environmental Quality 
in Illinois 


art L. Metcalf 
s R. Sanborn 


. 
. 
. 
: 
| 


F ILLINOIS 
TMENT OF REGISTRATION AND EDUCATION 


RAL HISTORY SURVEY DIVISION 
NA, ILLINOIS 


| VOLUME 31, ARTICLE 9 


STATE OF ILLINOIS DEPARTMENT OF REGISTRATION AND EDUCATION 

BOARD OF NATURAL RESOURCES AND CONSERVATION ' 
Ronaup E. STAcKieER, J.D., Chairman; THOMAS Park, Ph.D., Biology; L. L. Suoss, Ph.D., Geology; HWerBert S. 
Gutowsky, Ph.D., Chemistry; Ronert H, Anperson, B.S.O.E., Engineering; W. L. Everirt, E.E., Ph.D., Repre 
senting the President of the University of Illinois; Joun C. Guyon, Ph.D., Representing the President of Southern 
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Winus N. Bruce, Ph.D., Entomologist Hersert M. Dreier, M.S., Research Assistant 
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CONSULTANTS AND RESEARCH AFFILIATES: Sysrematic Exromonoay, Roperick R, Irwin, Chicago, Tl 
nois; WILDLIFE RESEARCH, WILLARD D. KuimstrRa, Ph.D., Professor of Zoology and Director of Cooperative Wil 
life Research, Southern Illinois University; Parasirouocy, Norman D. Levine, Ph.D., Professor of Vetert 
Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, University | 
Illinois; ENTomouoay, Ropert L. Mereanr, Ph.D., Professor of Zoology and of Entomology, University of Illinow 
and Ginpert P, WaAnpBauER, Ph.D., Professor of Entomology, University of Illinois; STATISTICS, Horace 
Norton, Ph.D., Professor of Statistical Design and Analysis, University of Illinois. 


CONTENTS 


SRE PEVVISEDCRATINGS & 2 tone wey EM ORO Os ts i ee SR 381 
BPEMBSAPLESTICHDES Ss eee ete TN AL MES A Ba 381 
BSeMISOH ES UR VEIMEICAN GE ot 8 fic tt 0S A ee eh 382 
BESRHISH ISK] OB WEESTIGIDEM USE: Best HAN a ee 383 
BML APNIN Gm LE CHNOLOG Var 222-272 kod aoe Bes Ne ed Lee Bim, 383 
Movet-Ecosystem MIECHNOLOG WER Pores < ents Tot iP ler ee 385 
See eID PEST aE SUTES Io ee. wien bee dE eon: Phe ee a ee. 386 
ORGANOPHOSPHORUS INSECTICIDE TEST RESULTS _......... 389 
BESEON CATE INSECTICIDE: LEST RESULTS (2) ceo 5 ens Se oo eon cec ceca dan cecesecdelineca bse 392 
SRISGerGANEOUS JINSEGTICIDE. LEST. (RESULTS 2.2.6.0. --2---0-6ccloceeecses see eee es 393 
SIRGANOGHLORINE INSECTIGIDE TEST RESULTS. ..........-.---.------cce-0c0ceceeenceeeeeeeeeeeneee 394 

BiGITOMEeM tl heTsistenCemaees CS. Sik 2 eg 395 
SECIONDE MNES TRE ESUINES pers ees Sr ee ee ee 399 
| CR ETUSSIOINT: - Lhe see Aion 0s el Oe seer SE cee Peet Sn OE ok 400 

Biological Biects s2..5. 7.02 Sac ne, ee 400 

Mp era cl atives DTOCUC Spe ee at ee BL 400 

LE walroyearea ll” IM Geyaa avn ey ato) apie teen eee eae ee ne ee 401 

Unextractable Radioactive Materials ~.......2.2.2-22-222222.--.2-eeeecceecee eee enn 402 
| TETRAGTCOT TD COP ODS Re ee oe et mE es 433 
GE sa ee ge es a Dt neces vas grist sh ne cnc needs enacteusscacttcssthartnee 436 


This report is printed by authority of the State of Illinois, IRS Ch, 127, Par. 58.12. 
It is a contribution from the Section of Economic Entomology of the Illinois Natural 
History Survey. 

Robert L. Metcalf is Professor of Biology and Research Professor of Entomology, 
University of Illinois. James R. Sanborn is an Assistant Entomologist, Illinois Natural 
History Survey. 


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Pesticides and Environmental Quality 


in Illinois 


ILLINOIS has 29,039,000 acres (1.18 
< 10’ ha) of farmland, amounting to 
34 percent of its land surface. This 
and is among the most fertile and pro- 
Juctive in the world, and Illinois ranks 
is the second state, after California, 
im producing farm crops, valued at 
33.167 billion in 1973. Illinois land 
oroduced 996,010,000 bushels (2.53 >< 
10" kg) of corn (17.6 percent of the 
U.S. total), 290,745,000 bushels (7.9 
10° kg) of soybeans (18.6 percent of 
the U.S. total), 37,800,000 bushels 
(1.03 < 10’ kg) of wheat (2.2 percent 
of the U.S. total), 19,780,000 bushels 
(2.88 < 10° kg) of oats (30 percent 
of the U.S. total), 3,251,000 tons (2.95 
< 10’ kg) of hay (2.4 percent of the 
U.S. total), and 4,225,000 pounds (1.92 
< 10° kg) of red clover seed (15 per- 
cent of the U.S. total). From these 
plant products Illinois produced an 
additional $1.906 billion worth of live- 
stock (4.2 percent of the U.S. total) 
(Illinois Cooperative Crop Reporting 
Service 1973). 

The value of Illinois farmland ex- 
ceeds $30 billion by current land value, 
and its corn crops alone have been 
valued at more than $30 billion over the 
past 100 years. However, in terms of 
its capability to help to feed a world 
which is growing ever hungrier, the 
value of Illinois soil can scarcely be 
overestimated. 


ACKNOWLEDGMENTS 


The research described in this report 
has been supported by a number of 
agencies, and portions of the data were 
obtained through the work of many 
individuals. Sponsors include the Her- 


Robert L. Metcalf 


James R. Sanborn 


man Frasch Foundation; American 
Chemical Society; Rockefeller Founda- 
tion; U.S. Environmental Protection 
Agency Grant EP-826 and Project R- 
800736; U.S. National Science Founda- 
tion Grant GI-39843; U.S. Department 
of the Interior through the Illinois 
Water Resources Center Grants B-050 
and B-070; World Health Organiza- 
tion; and Illinois Agricultural Experi- 
ment Station Regional Project NC-96. 
Individuals to whom particular thanks 
are due are Dr. Gary Booth, Dr. Dale 
Hansen, Dr. Asha S. Hirwe, Dr. Jorge 
Iwan, Dr. Inder Kapoor, Dr. Po-Yung 
Lu, Dr. Gurcharan Sangha, Dr. Ching- 
Chieh Yu, Margaret Anderson, Carter 
Schuth, and Patricia Sherman. 
Radiolabeled pesticides for evalua- 
tion were generously contributed by 
American Cyanamid, Badische Aniline 
Soda Fabrik Aktiengesellschaft, Chem- 
agro Corporation, Chevron Chemicals, 
CIBA-Geigy Corporation, Dow Chemi- 
cal, Eli Lilly and Company, FMC Cor- 
poration, Hercules Corporation, Mobile 
Chemical Company, Monsanto Com- 
pany, Morton Chemical Company, Na- 
tional Institutes of Environmental 
Health Sciences, Schering Corporation, 
Shell Chemical Company, Thompson- 
Hayward Company, Union Carbide 
Chemicals, Upjohn Company, Velsicol 
Corporation, and Zoecon Corporation. 


USE OF PESTICIDES 


Modern agricultural practices—in- 
volving superior plant varieties, im- 
proved cropping methods, heavy ap- 
plications of nitrogenous fertilizers, and 
extreme reliance on agricultural chemi- 
cals, especially herbicides and insecti- 


381 


382 


cides—have been responsible for the 
state's immense agricultural produc- 
tivity. These innovations have seen 
Illinois corn yields increase from 30 
bushels per acre (1,601 kg per ha) 
in 1920 to 105 bushels per acre (6,605 
kg per ha) in 1973. The use of pesti- 
cides in corn production has been de- 
scribed as being “as significant as the 
plow.” Their use has increased phe- 
nomenally, and in Illinois more total 
acreage, more than 14 million acres 
(5.67 10° ha), is treated with pesti- 
cides than is treated in any other state 
(Fowler & Mahan 1972). In 1972 herbi- 
cides were applied to 14,326,000 acres 
(5.79 < 10° ha) (49 percent of Illinois 
farmland ) and insecticides to 5,946,000 
acres (2.41 x 10° ha) (20 percent of 
Illinois farmland ) (Illinois Cooperative 
Crop Reporting Service 1973). On an 
acreage basis 14.7 percent of the herbi- 
cides and 14.1 percent of the insecti- 
cides used in U.S. agriculture were 
applied in Illinois although the state 
has only about 2.5 percent of the total 
cultivated land. We estimate (U.S. En- 
vironmental Protection Agency 1972a; 
Illinois Cooperative Crop Reporting 
Service 1973) that about 34 million 
pounds (1.54 x 10° kg) of the active 
ingredients of pesticides were applied 
to Illinois farm soil in 1971—equivalent 


to 1 pound for each acre (1.1 kg per - 


ha) in the state or 3 pounds (1.36 kg) 
for each of the state’s 11 million in- 
habitants. 

Much of the total amount of pesti- 
cides applied is dispersed throughout 
the environment (Frontispiece), enter- 
ing air, water, and food through vola- 
tilization and air currents, runoff and 
leaching, and uptake and concentration 
in food chains. 


NEED FOR SURVEILLANCE 


The heavy use of pesticides, chang- 
ing agricultural technology, and the 
rapid introduction of new pesticide 
products present a continuing demand 
for evaluation and surveillance of the 
effects of pesticides upon environmental 


It.ivois NaturaAL History SURVEY BULLETIN 


Vol. 31, Art. 9 


quality. The long-term effects of widely 
used pesticides are not well appreci- 
ated. Thus, von Riimker and Horay 
(1972), after a detailed survey of the 
most widely used pesticides, concluded 
that for 20 of the 35 compounds studied 
there was inadequate information about 
the nature of the environmental degra- 
dation products and their effects on en- 
vironmental quality. Considering that 
many of these pesticides, such as chlor- 
dane, toxaphene, dieldrin, propanil, 
captan, zineb, and maneb, were intro- 
duced 20 or more years ago, the mag- 
nitude of the problem is apparent. 
Furthermore, insect resistance to the 
organochlorine insecticides, together 
with increasingly severe effects of their 
use upon environmental quality, have 
resulted in their gradual replacement 
with organophosphorus and carbamate 
insecticides (Table 1). 

New pesticides are being introduced 
at a rate much faster than that of our 
scientific appreciation of their environ- 
mental effects. During the 30 years 
since World War II, the number of 
synthetic fungicides, herbicides, insecti- 
cides, nematocides, and rodenticides 
has increased from less than 100 to over 
900. The scene changes constantly with 
the development of new products and 
new technologies such as no-till farming. 
During 1974, for example, the following 
new pesticides were introduced under 
experimental permit into Illinois agri- 
culture: cyprazine (Prefox®), metri- 
buzin (Sencor®), bentazon (Basa- 
sran®), oryzalin (Surflan®), pro- 
fluralin (Tolban®), dinitramine (Co- 
bex®), bifenox (MODOWN®), gly- 
phosate (Round-up®), Rowtate®, and 
Counter®. Pesticides introduced under 
such experimental permits may be used 
on hundreds of thousands to millions _ 
of acres of Illinois soil in a few years. 
Thus, carbofuran, introduced in 1968, 
was used to treat 706,000 acres (287,- 
000 ha) in 1971, and trifluralin, intro- 
duced in 1964, was used to treat 1,226,- 
000 acres (496,000 ha) in 1971 (Petty 
& Kuhlman 1972). 


Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL Quatity 383 
Table 1.—Use of organochlorine insecticides on IIlinois farms. 
Year Insecticide Used and Acres Treated 
aldrin dieldrin DDT chlordane heptachlor toxaphene 
1968 3,438,000" aide ao 82,500 822,000 fin 
1969 3,512,000 11,000 9,000 160,000 1,131,000 24,000 
1970 2,690,000 is wae 63,800 822,000 
1971 1,690,000 0 0 233,000 232,000 
(2,240,000) (87,000) (654,000) 
1972 1,268,000 0 0 375,000 181,000 (35,000) 
(1,883,000) 
1973 afote 33e ioe aye 365 ee 
1974 1,400,000 0 0 200,000 400,000 (100,000) 


8 Data from Petty (1974) and data in parentheses from Illinois Cooperative Crop Reporting 


Service (1970 and 1973). 


In addition, farmers are increasing 
their use of combinations or mixtures 
of pesticides, either prepackaged or in- 
tank mixed. This proliferation of ma- 
terials and their persistence may pro- 
vide unintended soil mixtures. Pesti- 
cides are, by design, highly reactive 
biological compounds and may inter- 
act with one another in many ways 
to produce unintended effects, e.g., 
synergism in which the combined action 
is far greater than that of either of the 
components alone. Thus, the study of 
pesticide interactions in relation to 
environmental quality is much more 
complicated than the study of the indi- 
vidual components. As an example of 
this complexity, 29 combinations of 
herbicides were registered for use on 
corn and soybeans in Illinois in 1974 
(McGlamery et al. 1974). 


BENEFIT-RISK 
OF PESTICIDE USE 


The use of pesticides in such a prodi- 
gal way obviously poses benefit-risk 
questions which are very difficult to 
answer satisfactorily, especially in re- 
gard to the effects of pesticides on the 
total quality of the environment and on 
the long-term productivity of Ilinois 
soil. Two examples will illustrate this 
point. 

The use of certain preemergence 
herbicides allows no choice between 
planting corn or soybeans. The unusu- 


ally wet May and June of 1974 pre- 
vented corn production in many areas 
on land already treated with atrazine. 
This herbicide is highly toxic to soy- 
beans so that this crop was precluded 
as an alternative although it might have 
been the most profitable crop over a 
shortened growing season. 


The soil insecticide aldrin is con- 
verted by the action of air, bacteria, 
and enzymes in plants and animals to 
the epoxide dieldrin, one of the most 
persistent of all pesticides. More than 
60 million pounds (2.72 « 10’ kg) of 
aldrin have been applied in Ilinois 
since 1954, and the soil of this state 
has the highest average levels in the 
nation of aldrin (0.13 ppm) and diel- 
drin (0.11 ppm) ( Wiersma et al. 1972). 
The national averages are 0.02 ppm 
for aldrin and 0.03 ppm for dieldrin. 
Soybeans grown on soil long planted in 
corn average about 0.01 ppm of dieldrin 
although they have no federal toler- 
ance. Dieldrin residues in Illinois milk 
consistently exceed legal limits, and 
highly dieldrin-contaminated soybean 
sludges fed to poultry have resulted 
in the seizure and destruction of more 
than 25 million chickens in Mississippi 
(Anonymous 1974). 


EARLY-WARNING 
TECHNOLOGY 


The thoroughly unsatisfactory situ- 
ation in Illinois, resulting from the 


384 


widespread use of highly persistent 
organochlorine pesticides with little or 
no prior understanding of their fates in 
the total environment, has prompted 
both scientific and lay concern about 
a screening methodology which could 
serve as a simple early-warning system 
against potentially undesirable or haz- 
ardous effects of the large-scale use of 
new agricultural chemicals or com- 
binations of them. The wait-and-see 
system, followed in the use of aldrin, 
dieldrin, heptachlor, and chlordane and 
requiring a generation or more to dis- 
tinguish serious environmental pollu- 
tion, is demonstrably inadequate and 
has resulted in such disasters as the 
widespread contamination and seizure 
of milk supplies, the destruction of mil- 
lions of contaminated chickens, and 
the devastation of valuable fishing in- 
dustries. 

A recent comprehensive study, Pesti- 
cide Use on the Nonirrigated Crop- 
lands of the Midwest (U.S. EPA 1972a) 


Fig. 1.—The laboratory model ecosystem used to evaluate the fates and environmental 
effects of radiolabeled pesticides on terrestrial and aquatic organisms, including sorghum, 
salt-marsh caterpillar, plankton, alga, snail, mosquito larva, and mosquito fish. 


Iuyinors Natura History SURVEY BULLETIN 


ESTIGMENE 


OEDOGONIUM 


Vol. 31, Art. 9 


recommended that “a massive, interdis- 
ciplinary research effort be mounted to 
clarify the environmental behavior of 
major pesticides which are expected 
to continue in use for the forseeable 
future.” Information needed includes 
the fates of pesticides in the environ- 
ment after application; routes of me-— 
tabolism, degradation, and disappear- 
ance; natures of the ultimate break- 
down products; effects of long-term 
exposure of ecosystems to low-level 
residues; and interactions with othe 
chemicals in the environment. It wi 
be necessary to establish an order of 
priority among products to be investi 
gated in this fashion. { 

The investigations reported here rep- 
resent an effort by the State of Illinois 
through the Illinois Natural History 
Survey and the University of Illinois, 
to assume the responsibility for the 
comprehensive research so urgentl 
needed on the total environmental fate, 
of new pesticides. 


SORGHUM 


GAMBUSIA 


MODEL-ECOSYSTEM 
TECHNOLOGY 


The development of model-ecosys- 
tem or microcosm technology (Metcalf 
et al. 1971; Metcalf 1974) has provided 
a quick and sensitive laboratory tool 
for providing answers to these ques- 
tions about environmental pollution by 
pesticides: 

1. The nature of the biological ef- 

fects on non-target organisms 

2. The nature of degradative path- 

ways and the magnitudes of deg- 
radative products 

3. The bioconcentration and eco- 

logical magnification (EM) of 
parent compounds and degrada- 
tion products in living organisms 

4. The quantitative estimation of 

persistence and biodegradability 

Basically, model-ecosystem evalua- 
tion uses radiolabeled pesticides to fol- 
low qualitatively and quantitatively the 
movement and degradation of the com- 
pounds from a terrestrial (farm) en- 
vironment into an aquatic (lake) en- 
vironment and to demonstrate the pas- 
sage of the parent compound and 
its transformation products through 
aquatic food webs. The experimental 
model is shown in Fig. 1 and consists 
of a 20-gallon aquarium with a sloping 
shelf of washed quartz sand entering a 
lake of 7 liters of standard reference 
water (Freeman 1953), which provides 
mineral nutrition for plankton, alga, 
snail, mosquito larva, and fish and for 
sorghum plants growing on the ter- 
restrial farm area. The water phase 
of the system is aerated, and the entire 
system is kept in an environmental 
plant growth chamber at 80°F (26.5°C) 
with a 12-hour diurnal cycle of 5,000 
foot candles of fluorescent light. 

The radiolabeled pesticide to be 
tested is applied to sorghum plants, 
seeds, or to the soil of the system, 
using a realistic dosage of 1-5 mg per 
experiment, equivalent to 0.2-1.0 pound 
per acre (0.22-1.1 kg per ha). Ten 
last-instar salt-marsh caterpillars, Estig- 


— Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


385 


mene acrea, are introduced to consume 
the treated sorghum plants, and the 
caterpillars and their excretory prod- 
ucts, leaf frass, etc., contaminate the 
lake portion of the model system. The 
radiolabeled products enter the vari- 
ous aquatic food chains, e.g., plank- 
ton —> daphnia (Daphnia magna) > 
mosquito (Culex pipiens) —> fish 
(Gambusia affinis) or alga (Oedo- 
gonium cardiacum) —> snail (Physa 
spp. ). 

The movement of the radiolabeled 
products from plants to lake are mea- 
sured by counting the radioactivity of 
duplicate 1-ml water samples by liquid 
scintillation at intervals of 1, 2, 4, 7, 
14, 21, 28, and 33 days or whenever 
desired. After the system has been in 
operation for 26 days, 300 mosquito 
larvae are added, and after 4 more days 
50 are removed for analysis. The food 
chains are completed after 30 days by 
adding three mosquito fish, G. affinis, 
which are left for 3 days to eat the 
daphnia and mosquito larvae. 

The experiment is terminated after 
33 days, when weighed samples of the 
various organisms are homogenized in 
small volumes of acetonitrile. Aliquots 
are counted for total radioactivity by 
liquid scintillation. One liter of water 
from the system is extracted three times 
with diethyl ether to measure total 
radioactivity. The residual water is 
hydrolyzed with 1.0 N_ hydrochloric 
acid for 4 hours and reextracted with 
diethyl ether to determine the conju- 
gated materials, and the amount of un- 
extractable radioactive materials is de- 
termined by counting the radioactivity 
of the remainder. 

The acetonitrile extracts of the or- 
ganisms are concentrated to a few milli- 
liters and known volumes are applied 
to thin-layer chromatography (TLC) 
plates of fluorescent silica gel (E. 
Merck GF-254). TLC is carried out 
with appropriate solvents (identified in 
the tables) and with the incorporation 
of standard known metabolites of the 
pesticide under study. After the chro- 


386 


matograms are developed, they are 
placed against X-ray film and exposed 
for several weeks to several months to 
determine the areas containing radio- 
labeled products. These areas are 
scraped into scintillation vials, and scin- 
tillation counts are made to determine 
the amounts of individual degradation 
products present. The residues from 
the tissue extractions are combusted to 
determine the amount of unextractable 
radioactive materials, using either the 
Schoeniger oxygen flask technique 
(Kelly et al. 1961) or a tissue solubili- 
zation method. 


After the completion of these assays, 
the results of the experiment are as- 
sembled on balance sheets showing the 
amounts and natures of radiolabeled 
degradation products present. Wher- 
ever possible, the chemical identities 
of the degradation products are de- 
termined by cochromatography with 
known model compounds, by the use of 
specific microchemical reactions and 
by infrared and mass spectrometry. 
The results of such studies on 48 pesti- 
cides are shown in the tables. 


HERBICIDE TEST RESULTS 


The importance of examining the 
fates of herbicides in a_terrestrial- 
aquatic model ecosystem cannot be 
overestimated, especially in view of the 
exponential growth in the use of herbi- 
cides over the past 20 years in the 
United States. Pimental et al. (1973) 
estimated that in 1945 the use of herbi- 
cides for controlling weeds in corn was 
practically nonexistent. However, in the 
25-year period from 1945 to 1970 the 
use of herbicides increased significantly, 
and it was estimated that by 1970 
herbicide treatment averaged 1 pound 
of active ingredient per acre (1.1 kg 
per ha). Though figures were not avail- 
able for 1945, it is possible to examine 
figures for 1950-1970, which clearly 
demonstrate that herbicide use on corn 
increased at least twentyfold during 
that time. 


Inutinors NATuRAL History SURVEY BULLETIN 


0.2100, 0.05479, and 0.07356 ppm, re- 


Vol. 31, Art. 9 


Alachlor, or 2-chloro-2’, 6’-diethyl- 
N-(methoxymethyl)-acetanilide, is a 
member of a large class of chloro- 
acetanilide herbicides used to control 
annual grasses in cornfields and certain 
broadleaf weeds in corn or soybeans, 
The data clearly indicate the suscepti- 
bility of this herbicide to extensive 
degradation, as no residues of alachlor 
were isolated from any of the test 
organisms (Table 2). The high degree 
of degradation is further evidenced by 
the large number (10) of radiolabeled 
products of alachlor isolated from the 
water section of the ecosystem. Con- 
tinued use of this herbicide should not 
lead to its accumulation in aquatic 
food chains. 

Atrazine, or 2-chloro-4-(ethylamino)- 
6-(isopropylamino )-s-triazine, is one of | 
the most extensively used herbicides 
for controlling weeds in corn plantings. 
The alga, snail, and fish of the model 
ecosystem contained 2.4059, 0.2386, and 
0.3511 ppm, respectively, of atrazine 
(Table 3). The percentages of atrazine 
in the radioactive materials extractable 
from the alga, snail, and fish were 87.3, 
63.1, and 59.3, respectively. The EM 
values for atrazine for the alga, snail, 
and fish were 75.6, 7.5, and 11.0, re- 
spectively. In addition, the alga, snail, 
and fish contained smaller amounts, 


spectively, of N-dethylatrazine (com- 
pound A, Table 3). Another N-deal- 
kylated product, N-deisopropylatrazine 
(compound B, Table 3), was isolated 
from the alga (0.04934 ppm), snail 
(0.02796 ppm), and fish (0.05496 
ppm). The EM values of these two 
dealkylated metabolites were of the 
same order of magnitude as that ob- 
served for~atrazine. Continued use of 
atrazine would not appear to lead to 
major accumulations in aquatic food 
chains. 

Bentazon, or 3-isopropyl-1H-2,1,3- 
benzothiadiazin-4 - (3H ) -one-2, 2-diox- 
ide, is a new herbicide employed for 
the control of a selected number of 
broadleaf and sedge weeds. In the 


Aug.,1975 Metcatr & SANBORN: PEsTICIDES AND ENVIRONMENTAL QuaLity 387 


model ecosystem (Booth et al. 1973) 
it was susceptible to degradation, as 
indicated by the lack of residues in all 
organisms except the clam, which con- 
tained 0.622 ppm of N-isopropylanthra- 
nilamide, 1.266 ppm of anthranilic acid, 
and 0.510 ppm of unchanged bentazon 
(Table 4). The percentage of bentazon 
in the radioactive materials extractable 
from the clam was 18.7, and the EM 
value was about 10. Continued use of 
this herbicide should not lead to its ac- 
cumulation in aquatic food chains. 
Cyanazine, or 2-chloro-4-(1-cyano-l- 
methylethylamino )-6-ethylamino-s-tria- 
zine, is used for the control of annual 
grasses and broadleaf weeds in corn- 
fields. The behavior of this herbicide 
in the model ecosystem indicates that 
it is susceptible to degradation, as only 
the water plant, Elodea, contained resi- 
dues of this herbicide (Table 5). 
Neither the fish nor the snail contained 
residues of cyanazine or its degradation 
products. The high water solubility, 171 
ppm, of cyanazine and its apparent 
susceptibility to degradation clearly 
demonstrate that the continued use of 
cyanazine should not result in its ac- 
cumulation in aquatic food chains. 
Dicamba, or 3,6-dichloro-o-anisic 
acid, is an effective herbicide for the 
control of both annual broadleaf weeds 
and grasses in corn. The data indicate 
clearly that this herbicide is not ab- 
sorbed by the organisms of the model 
ecosystem (Yu et al. 1975a) (Table 6). 
This fact is probably related to the 
pH of the aqueous portion of the 
model ecosystem, which is higher than 
the pKa (dissociation constant) of this 
benzoic acid derivative; therefore, the 
herbicide exists in the ionic form. 
Dicamba in the ether-extracted water 
constitutes about 90 percent of the ex- 
tractable radioactive materials. Al- 
though the data do not indicate it, 
dicamba was recovered from the water 
only after acidification and heating for 
24 hours. It is impossible to state 
whether the dicamba was in the ionic 
form and that acidification facilitated 


the partition of dicamba into ether, or 
whether the dicamba was present as 
a conjugate and that the acid treatment 
broke down the conjugate and released 
the free acid. In any case, very little 
happened to dicamba in the water 
of the model ecosystem other than con- 
jugation through the carbonyl moiety. 

Phenmedipham, or methyl m-hy- 
droxycarbanilate m-methylcarbanilate, 
is a postemergence herbicide used in 
sugar beets to control a large variety 
of annual weeds. The fate of phen- 
medipham in this model ecosystem 
clearly indicates the susceptibility to 
degradation of this herbicide, as none 
of the organisms contained phenmedi- 
pham residues (Table 7). The radio- 
active material extractable from the 
fish remained at the origin of the TLC 
plate, indicating the polar nature of the 
radioactivity. The continued use of 
phenmedipham should not lead to its 
accumulation in aquatic food chains. 

2,4-D, or 2,4-dichlorophenoxyacetic 
acid, is one of the oldest synthetic 
herbicides in use today. After more 
than 30 years of its continued use, prob- 
lems relating to aquatic food-chain ac- 
cumulation of 2,4-D are nonexistent. 
The data from the experiment with 
“C-24-D corroborate the “outdoor” 
data that have accumulated for the past 
three decades, as no 2,4-D residues 
were found in any of the organisms 
of the model ecosystem (Table 8). As 
might be expected, the alga contained 
the greatest number of unidentifiable 
4C residues even though eight standard 
degradation products of 2,4-D were 
cochromatographed. Continued use of 
24-D does not appear to lead to en- 
vironmental problems relating to its 
accumulation in aquatic food chains. 
“Real-world” data and model ecosystem 
results are similar and clearly demon- 
strate the ability of this microcosm to 
predict potential environmental prob- 
lems. 

Propachlor, or 2-chloro-N-isopro- 
pylacetanilide, is one of a large number 
of a-chioroacetanilide herbicides, which 


388 


include alachlor, that are used to con- 
trol annual grasses and some broadleaf 
weeds in a number of crops including 
corn and soybeans. The structural simi- 
larity of propachlor to alachlor and its 
great susceptibility to degradation are 
evident, as none of the organisms con- 
tained residues of this herbicide (Table 
9). There was a very minute amount 
of propachlor (0.0564 ppb) in the 
water at the end of the experiment. 
Clearly the a-haloacetanilides are some 
of the most degradable herbicides ex- 
amined in this system, and continued 
use of these herbicides should not lead 
to their accumulation in aquatic food 
chains. 

Pyrazon, or 5-amino-4-chloro-2- 
phenyl-3-(2H)-pyridazinone, is used 
for the control of annual broadleaf 
weeds in sugar beets and beets. The 
model ecosystem data clearly demon- 
strate that pyrazon is susceptible to 
degradation, as only the crab contained 
residues (0.476 ppm) of this herbicide, 
which constituted 95.4 percent of the 
radioactive materials extractable from 
the crab (Table 10). The EM value for 
the pyrazon in the crab was 22.5 (Yu 
et al. 1975b). Continued use of this 
herbicide would not appear to lead to 
problems related to accumulations of 
it in aquatic food chains. 

Trifluralin, or a,a,a-trifluoro-2,6-di- 
nitro-N,N-dipropyl-p-toluidine, is used 
to control grasses and several broadleaf 
weeds in soybeans, cotton, and many 
other crops. Only the snail and fish 
contained 5.046 ppm and 0.261 ppm, 
respectively, of trifluralin as an ex- 
tractable residue (Table 11). The 
percentages of trifluralin in the ex- 
tractable radioactive materials in the 
snail and fish were 75.7 and 34.0, re- 
spectively. The EM values for the 
snail and fish were 17,872 and 926, re- 
spectively. In addition to trifluralin the 
snail contained lesser amounts of a,a, 
a-trifluoro-2,6-dinitro-N-propy]- p-tolui- 
dine (0.337 ppm), which had an EM 
value of 3,874. Trifluralin is the only 


Inuinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 9 


herbicide tested that showed a_pro- 
pensity to accumulate in either the fish 
or snail. Its tendency to accumulate is 
undoubtedly related to its low water 
solubility (0.58 ppm) and high lipid 
solubility (Probst & Tepe 1969). De- 
spite the accumulation in the snail 
and fish, trifluralin is unusual in that 
it is susceptible to degradation, form- 
ing at least 11 degradation products 
in water, yet demonstrates a tendency 
to be magnified to some extent through 
aquatic food chains. It is not, however, 
magnified at the level of chlorinated 
hydrocarbons, but at a level very similar 
to that of the insecticide methoxychlor, 
which has an EM value of about 1,500. 

Metrabuzin, or 4-amino-6-tert-butyl- 
3-(methylthio ) as-triazin-5- (4H )-one, 
is a new herbicide used for weed con- 
trol in soybeans. The data in Table 12 
clearly demonstrate the degradability 
of this herbicide in the model ecosys- 
tem, as no residues of this herbicide 
were isolated from the organisms. Fur- 
ther, the water contained numerous 
metabolites, which is indicative of the 
susceptibility of this herbicide to degra- 
dation under the conditions of this ex- 
periment. The major degradation prod- 
uct in the water is a mixture of DK and 
DADK, which were not resolvable by 
thin-layer chromatography. The data 
from this system clearly indicate that 
the continued use of this herbicide 
should not lead to its accumulation in 
aquatic food chains. 

Bifenox, or methyl-5-(2’,4’-dichloro- 
phenoxy )-2-nitrobenzoate, is a new pre- 
emergence herbicide somewhat related 
to 2,4-D. As shown in Table 13, bi- 
fenox is: degraded by hydrolysis of the 
methyl ester to form the parent benzoic 
acid (compound B, Table 13), and by 
reduction of the nitro group to the cor- 
responding amino compound (com- 
pound A, Table 13). There was no 
evidence of cleavage of the diphenyl 
ether moiety. Bifenox is of low water 
solubility (0.35 ppm) (Fig. 2) and was 
bioconcentrated about 200-fold by the 


Aug., 1975 Mercatr & SANBORN: PesTICIDES AND ENVIRONMENTAL QuaALITY 


fish. It falls in the borderline area of 
moderate biodegradability and should 
be used with care. 


ORGANOPHOSPHORUS 
INSECTICIDE TEST RESULTS 


The decline in the use of organo- 
chlorine insecticides to control pest 
species (Table 1) is the result of factors 
such as target-pest resistance, environ- 
mental hazards, and more recently, the 
ban imposed by the U.S. Environmental 
Protection Agency (EPA) on DDT and 
aldrin/dieldrin as general insecticides 
for home and agricultural use. Further, 
in view of the recent action of the EPA 
seeking to ban the use of chlordane, 
heptachlor, and heptachlor epoxide, it 
is certain that more phosphate and 
carbamate insecticides will be used to 
fill the void left by the elimination of 
the organochlorine insecticides. There- 
fore, it is essential to examine carba- 
mate and phosphate insecticides to 
insure that no problems of the enyiron- 
mental persistence and aquatic food- 
chain accumulations of these insecti- 
cides will occur. 

Chlorpyrifos, or O,O-diethyl-O- (3,5, 
6-trichloro-2-pyridyl) phosphorothio- 
nate, had EM values in the alga, snail, 
mosquito, and fish of 72, 691, 45, and 
320, respectively. Of the radioactive 
material extractable from each _or- 
ganism, the percentages of chlorpyrifos 
isolated from the alga, snail, mosquito, 
and fish were 30.3, 48.1, 7.9, and 49.5, 
respectively (Table 14). The position 
of the “C label in the pyridyl ring 
allows the investigation of the per- 
sistence of this moiety in the organisms 
of the system or its uptake by them or 
both. The ecological magnification and 
percentage of the extractable radio- 
active materials for the pyridinol in 
each organism were: alga, 44, 18.8 
percent; snail, 443, 32.3 percent; mos- 
quito, 191, 34.9 percent; and fish, 180, 
29.1 percent. The absence of the oxon 
of chlorpyrifos in any of the organisms 
is typical, as the oxons of the phosphate 


389 


insecticides were not found generally 
in any of the organisms. 

Chlorpyrifos-methyl is an insecticide 
similar to chlorpyrifos except for the 
substitution of O,O-dimethyl for O,O- 
diethyl groups to yield O,O-dimethy]-O- 
(3,5,6-trichloropyridinyl ) phosphorothi- 
onate. The chlorpyrifos-methy] ecologi- 
cal magnification values for the alga, 
snail, mosquito, and fish are 478, 544, 
1,875, and 95, respectively. The values 
for the snail and fish are substantially 
lower than those found in the organisms 
subjected to chlorpyrifos, the result of 
the greater susceptibility of the O- 
methyl groups to degradation as com- 
pared to that of the O-ethyl moieties in 
chlorpyrifos. The percentages of chlor- 
pyrifos-methyl in the radioactive ma- 
terials isolated from the alga, snail, 
mosquito, and fish were 49.0, 49.3, 68.2, 
20.7 percent, respectively (Table 15). 
Again, because the “C label is located 
in the pyridyl moiety, it is possible to 
investigate the fate of this group in 
the model ecosystem. The ecological 
magnification and percentage of the 
chlorinated pyridinol in the organisms 
were: snail, 41, 9.3 percent; fish, 54.5, 
29.7 percent. As was observed for 
chlorpyrifos, none of the organisms 
contained the activation product, chlor- 
pyrifosoxon-methyl. 

Counter® is one of the newer phos- 
phate insecticides under development 
for use as a soil insecticide, and it has 
the chemical name of O,O-diethyl S- 
(tert-butylthio )-methyl phosphorodi- 
thioate. This insecticide was therefore 
applied in the sand of the model eco- 
system to mirror its use in the field. 
The similarity in structure to phorate 
(Thimet®) and disulfoton (Di-Sys- 
ton®) is obvious, and the degradation 
in pathways of sulfur oxidation in the 
side chain of Counter® was similar to 
those of the other two pesticides. The 
percentages of Counter® in the radio- 
active materials extractable from the 
alga, snail, mosquito, and fish were 3.3, 
23.5, 4.7, and 25.0, respectively (Table 


390 


16). No other metabolites were iso- 
lated from the fish or mosquito although 
a small amount (0.0241 ppm) of 
Counter® oxon was observed in the 
snail. The Counter® ecological mag- 
nification values from the alga, snail, 
mosquito, and fish were 175, 1,830, 360, 
and 535, respectively. These values 
from the fish and snail are somewhat 
higher than those found for most other 
phosphate insecticides. Undoubtedly 
these higher values are related both 
to the initial stability of the phosphoro- 
dithionate and to the application of this 
chemical to the sand, which does not 
allow for the initial metabolism and 
degradation by the caterpillars. The 
water sector of the ecosystem contained 
only trace amounts of Counter® and 
of nearly all of the possible combina- 
tions of the oxidation products of phos- 
phorothioate and sulfide sulfur. 


Temephos (Abate®), or the bis-O,O- 
dimethylphosphorothioate ester of 4,4’ 
dihydroxydipheny] sulfide, is an excel- 
lent mosquito larvicide and appears to 
possess ideal environmental charac- 
teristics, as it is exceptionally degrad- 
able. No residues of temephos or any 
of its oxidative or hydrolytic metabo- 
lites occurred in the fish. Because of 
its high larvicidal activity, the mos- 
quitoes were killed throughout the 
usual duration of the experiment, and 
it was extended to 53 days. The alga 
and snail contained small amounts 
(0.00195 and 0.01876 ppm, respec- 
tively) of temephos (Table 17). The 
EM values of temephos from the alga 
and snail were 1,500 and 14,431, re- 
spectively. In addition, the alga con- 
tained small amounts (0.4-2.0 ppb) of 
all of the cochromatographed metabo- 
lites, and the snail contained substan- 
tially fewer of the metabolites though 
at somewhat higher concentrations 
(2-27 ppb). The higher concentrations 
in the snail again emphasize the low 
titer of enzymes in this organism ca- 
pable of degrading foreign compounds. 
The absence of data for the mosquito 


Iturnois NaTuRAL History SuRVEY BULLETIN 


Vol. 31, Art. 9 


emphasizes the outstanding larvicidal 
properties of this insecticide. 

Fonofos (Dyfonate®), or O-ethyl- 
S-phenyl ethylphosphonodithioate, is 
an effective soil insecticide which is 
finding increasing use as a replacement 
for the organochlorine insecticides, 
Although the organisms of the model 
ecosystem contained small amounts 
of the unchanged fonofos, none con- 
tained significant amounts of degrada- 
tion products (Table 18). The per- 
centages of fonofos in the radioactive 
materials extractable from the alga, 
snail, and fish were 32.1, 27.0, and 
80.5, respectively. Further, the fonofos 
in the alga, snail, and fish had EM 
values of 108, 86, and 77, respectively. 
The large number of degradation prod- 
ucts isolated from the water (14), 
coupled with the very low EM values, 
clearly indicates that fonofos does not 
accumulate significantly in aquatic food 
chains. 

Fenitrothion, or O,O-dimethyl]-O-(3- 
methyl-4-nitrophenyl) phosphorothio- 
nate, is one of the safest organophos- 
phorus insecticides, as the LD,, for 
the rat is 500 mg per kg and for the 
mouse is 1,200 mg per kg. The substi- 
tution of the methyl group in the meta 
position of the nitrophenyl ring of 
methyl] parathion is believed to be re- 
sponsible for the much reduced mam- 
malian toxicity as compared to that of 
methyl parathion, of which the LD,, 
for the rat is 13 mg per kg and for the 
mouse is 75 mg per kg. Fenitrothion 
EM values of 349, 2.2, and 9.8 were 
found for the alga, mosquito, and fish, 
respectively. The percentages of feni- 
trothion in the radioactive materials 
isolated from the alga, mosquito, and 
fish were 33.7, 6.6 and 44.4, respectively 
(Table 19). The only other degrada- 
tion product isolated from the orga- 
nisms was a small amount (5.7 ppb) 
of fenitroxon found in the fish. This 
degradation product of fenitrothion had 
an EM value of 6.5. The isolation of 
this phosphorus oxon from the fish is 


Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


unique, as none of the other oxons 
of the phosphate insecticides were 
found in the fish. 

Malathion, or O,O-dimethyl-S-(1,2- 
dicarboethoxyethy] )-phosphorodithio- 
ate, is widely used in the home and 
garden as an insecticide. It appears to 
be exceptionally degradable, as no 
traces were found in any of the model- 
ecosystem organisms (Table 20). The 
fish, snail, and mosquito contained sev- 
eral uncharacterized metabolites, which 
were also found in the water. It is 
apparent that malathion is one of the 
most degradable organophosphorus in- 
secticides examined in this system. This 
degradability, together with malathion’s 
low mammalian toxicity (rat oral LD,,, 
1,300 mg per kg), makes it a safe and 
useful product. 

Acephate (Orthene®), or O-methyl- 
S-methyl-N-acetylphosphoramidothio- 
ate, is a relatively new insecticide, 
which has found widespread use in the 
control of pests of vegetables. The 
parent insecticide was not isolated from 
any of the model-ecosystem organisms 
(Table 21), which is not unexpected in 
view of the high water solubility of 
acephate (650,000 ppm). However, an 
uncharacterized degradation product 
was isolated (R; 0.93) in all of the 
organisms except the clam and fish. 
In the crab this degradation product 
had an EM value of 4,273 times the 
concentration in the water. Further 
research is in progress to determine the 
structure of this degradation product. 

Leptophos (Phosvel®), or O-(4- 
bromo-2,5-dichloropheny] ) -O-methy] 
phenylphosphonothionate, is a new or- 
ganophosphate insecticide now under- 
going extensive development for use 
in controlling pests of cotton and veg- 
etable crops. The available environ- 
mental degradation information ( Holm- 
stead et al. 1973; Aharonson & Ben-Aziz 
1974) clearly indicates that this insecti- 
cide has a high degree of environmental 
stability. Other problems with this in- 
secticide have been found in its use in 


391 


Egypt on cotton, where it killed 1,300 
water buffaloes (Shea 1974). Labora- 
tory experiments with chickens have 
shown that leptophos has neurotoxic 
effects (Abou-Donia et al. 1974). 

The behavior of leptophos in our 
model ecosystem indicates that it is 
one of the most persistent phosphorus- 
derived pesticides examined (Table 
22). The experiment was extended to 
45 days, because each time the mos- 
quitoes were introduced, they immedi- 
ately died. Even though the mosquitoes 
died after their introduction on the 
45th day, the fish were then added to 
the ecosystem, and the experiment was 
terminated 3 days later. Every organism 
contained residues of leptophos, the 
alga having 13.221 ppm, the snail 52.27 
ppm, and the fish 1.559 ppm. These 
residues of leptophos in the radioactive 
materials extracted from the alga, snail, 
and fish constituted 41.8, 97.3, and 
83.5 percent, respectively, of the totals. 
The EM values for leptophos were 
12,243 for the alga, 48,398 for the 
snail, and 1,444 for the fish, respec- 
tively. Clearly, this is the most per- 
sistent organophosphorus insecticide ex- 
amined in the model ecosystem. 

Parathion, or O,O-diethyl O-4-nitro- 
phenyl phosphorothionate, and methyl 
parathion, its O,O-dimethyl analogue, 
were produced in the United States in 
1970 in the combined amount of about 
56 million pounds. The available in- 
formation on the behavior of parathion 
and methyl parathion in the environ- 
ment indicates that they have presented 
no problems of accumulation in aquatic 
food chains after more than 25 years 
of widespread use. The model-ecosys- 
tem data (Table 23) corroborate the 
outdoor data. The only organism con- 
taining a residue of parathion was the 
fish, and there the concentration was 
only 0.1006 ppm, which constituted 
about 52 percent of the radioactive ma- 
terials isolated from the fish. The ex- 
periment was lengthened to 38 days 
because of the toxicity of the water to 


392 


the mosquito. The use of 2,6-'C-la- 
beled 4-nitrophenol-labeled parathion 
allowed the examination of the fate of 
this moiety, and it was determined that 
the water (0.000136 ppm) and fish 
(0.0086 ppm) contained small amounts 
of this moiety. 


CARBAMATE 
INSECTICIDE 
TEST RESULTS 


The carbamate insecticides recently 
have assumed a large role in Illinois 
agriculture with the elimination of the 
organochlorine insecticides because of 
the resistance of target pests, the en- 
vironmental accumulative tendency of 
the organochlorine compounds, and 
their carcinogenic properties. The use 
of metalkamate, carbofuran, and car- 
baryl to control insect pests on corn 
and soybeans has proved to be effective 
and has eliminated the aquatic food 
chain accumulation problems of the 
formerly used chlorinated hydrocarbon 
insecticides. 

Metalkamate is a 3:1 mixture of 
m-(1-ethylpropyl)-phenyl and m-(1- 
methylbutyl)-phenyl N-methylcarba- 
mates introduced to control soil pests 
of corn. This insecticide does not have 
any tendency to accumulate in the 
higher members of the trophic web, 
though the alga (0.980 ppm); crab 
(0.0498 ppm), which died 7 days after 
the introduction of metalkamate; and 
Elodea (0.245 ppm) contained residues 
of the parent compound (Table 24). 
These residues of metalkamate in the 
alga, crab, and Elodea constituted 55.0, 
17.4, and 25.9 percent, respectively, of 
the extractable radioactive material 
from these organisms. The most inter- 
esting observation here is that these 
three organisms were the only orga- 
nisms that contained detectable amounts 
of “C, None of the other organisms had 
substantial amounts of “C residues. 
While this insecticide has not been as 
effective recently as it has been in the 
past in controlling pests of corn, its 
environmental behavior in the model 


Intinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 9 


ecosystem clearly indicates that should 
it become widely employed, no aquatic 
food chain accumulation problems are 
likely to arise. 

Carbaryl, or 1-naphthyl N-methyl- 
carbamate, was the first carbamate in- 
secticide to find widespread use in the 
home garden and in agriculture, and 
it is presently the most widely used 
insecticide in the United States. With 
the banning for general use of DDT 
in 1972, carbaryl is being used to con- 
trol the tussock moth in the Pacific 
Northwest; the gypsy moth, which is 
migrating westward from the eastern 
regions of the United States; and the 
spruce budworm. After more than 20 
years of widespread use, neither prob- 
lems of accumulations in food chains 
nor of ubiquitous food residues have 
been experienced. The data from the 
terrestrial-aquatic model ecosystem 
(Table 25) definitely corroborate the 
experience in the field, as no residues 
of carbaryl were found in any of the 
organisms. The water contained many 
degradation products of carbaryl, but 
no residues of carbaryl itself. Con- 
tinued widespread use of this insecti- 
cide will definitely not lead to prob- 
lems associated with accumulations in 
aquatic food chains. 

Carbofuran, or 2,2-dimethyl-2,3-di- 
hydrobenzofuranyl-7-N-methylearba- 
mate, is an excellent soil insecticide 
for the control of com and soybean 
pests. The behavior of this carbamate 
insecticide is similar to that of the other 
carbamates examined in that none of 
the organisms in the model ecosystem 
contained residues of the parent in- 
secticide (Table 26). The water con- 
tained a small amount of carbofuran 
(0.003889 ppm) as well as trace 
amounts of other metabolites and deg- 
radation products of carbofuran (Yu 
et al. 1974). It appears that the con- 
tinued use of this insecticide will not 
lead to environmental problems of ac- 
cumulations in aquatic food chains. 

Propoxur, or 2-isopropoxyphenyl N- 
methylcarbamate, is used for household 


Aug., 1975 Metrcatr & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


pest control and for residual spraying 
for adult mosquitoes. In the model 
system every organism contained resi- 
dues of propoxur at concentrations of 
0.0360, 0.0928, 0.4441, and 0.0468 ppm 
for the alga, snail, mosquito, and fish, 
respectively (Table 27). The percent- 
ages of propoxur in the radioactive ma- 
terials extracted from the alga, snail, 
mosquito, and fish were 7.8, 23.5, 19.4, 
and 39.9, respectively. The EM values 
for the alga, snail, mosquito, and fish 
are 112, 290, 1,388, and 146, re- 
spectively. In addition to the parent 
compound, the fish contained lesser 
amounts of 2-isopropoxyphenol (0.0252 
ppm) and 2-isopropoxyphenyl N-hy- 
droxymethyl carbamate (0.0180 ppm). 
Propoxur was the only carbamate ex- 
amined in this model ecosystem that 
was accumulated by the fish. This fact 
may be, in part, related to the high 
specific activity of the radiolabeled 
propoxur (10.4 mCi/mM ), which made 
it possible to determine the small resi- 
dues of this insecticide in the orga- 
nisms. 

Aldicarb is a systemic carbamate in- 
secticide, 2-methy]-2-methylthiopropi- 
onaldoximyl N-methylearbamate. Aldi- 
carb is readily oxidized in vivo to sulf- 
oxide and sulfone metabolites, both 
of which are insecticidal. These metab- 
olites and the parent compound form 
relatively persistent systemic toxicants 
in plant tissues (Metcalf et al. 1966). 
A single application to the roots of 
cotton plants kills boll weevil larvae 
during an entire growing season. There- 
fore, it was not unexpected to find 
these products persisting over the 33- 
day period of the model-ecosystem ex- 
periment (Table 28). However, the 
substantial water solubility of aldicarb, 
0.6 percent, clearly prevented high bio- 
magnification in the organisms, and the 
EM value in the fish was 42. Aldicarb 
was highly toxic to the snail, Physa, 
and all of these died early in the course 
of the experiment. 


Formetanate, or 3-dimethylamino- 
methyleneiminophenyl N-methylcarba- 


393 


mateehydrochloride, is a carbamate 
acaricide. As shown in Table 29, this 
compound is highly biodegradable, and 
no trace of the parent compound was 
found in the model ecosystem after 
33 days. The only identifiable degrada- 
tion product (compound A, Table 29) 
involved removal of the N-methylcar- 
bamoyl group and loss of the amidino 
moiety. We do not expect that this 
compound will cause problems in en- 
vironmental quality. 


MISCELLANEOUS 
INSECTICIDE 
TEST RESULTS 

Methoprene, or isopropyl-11-me- 
thoxy-3,7,11-trimethyldodeca-2,4-dieno- 
ate, is one of the “fourth-generation” 
insecticides believed to interfere with 
the normal metamorphic development 
of insects. This pesticide has shown 
some promise in the control of mos- 
quitoes developing in irrigated fields in 
California. The degradation of metho- 
prene has been examined in detail in 
several outdoor systems (Quistad et al. 
1974 and 1975; Schooley et al. 1975). 
In the model ecosystem every orga- 
nism contained residues of methoprene 
(Table 30), with the alga containing 
2,220 ppm, the snail 1.500 ppm, and the 
fish 0.0176 ppm. These methoprene 
residues in the alga, snail, and fish con- 
stituted 48.0, 30.7, and 25.1 percent, 
respectively, of the radioactive materials 
extracted from each organism. The EM 
values for methoprene in the alga, snail, 
and fish were 25,814, 17,442, and 205, 
respectively. Measurable amounts of the 
11-O-demethylated methoprene were 
isolated from the alga, 0.723 ppm; 
snail, 0.469 ppm; and fish, 0.0181 ppm 
though the water contained none of 
this degradation product. Finally, the 
water, snail, and fish contained small 
amounts of 11-hydroxy-3,7,11-trimethyl- 
dodeca-2,4-dienoic acid. 

Dimilin, or 1-(2,6-difluorobenzoy])- 
3-(4-chlorophenyl) urea, is a recently 
introduced insecticide which apparently 
interferes with the normal development 


394 


of the insect cuticle and leads to mor- 
tality at molting. The use of two dif- 
ferent “C-labeled sites in dimilin en- 
abled us to examine the fates of the 
two phenyl moieties. Every organism 
contained this insecticide (Table 31), 
from the high of 13.1369 ppm in the 
mosquito in the “C-chlorophenyl urea 
dimilin to the low of 0.1097 ppm in 
the fish in the “C-difluorobenzoy] dimi- 
lin. Despite the variation in the ab- 
solute quantity of dimilin in the fish of 
the two experiments, 0.1097 ppm for 
the “C-difluorobenzoyl and 0.3193 ppm 
for the “C-chlorophenyl urea, the EM 
values of 19.2 and 14.5 were very close. 
The percentage of dimilin in the ex- 
tractable radioactive materials isolated 
from the fish was 6.7 percent for “C- 
difluorobenzoy] dimilin and 5.3 percent 
for “C-chloropheny! dimilin, indicating 
again close agreement in the data for 
the two “C labels. While dimilin 
amounted to a small percentage of the 
extractable radioactive materials in the 
fish, the fractions of dimilin were con- 
siderably higher (46-98 percent) in 
the radioactive materials isolated from 
the rest of the organisms. 


Chlordimeform, or N-(4-chloro-o- 
tolyl)-N,N-dimethylforamidine, is one 
of the newer insecticides and appears 
to be effective in controlling cotton 
pests. In the model ecosystem only the 
snail contained-residues of this insecti- 
cide, with a concentration of 0.0710 
ppm (Table 32). The fraction of 
chlordimeform in the extractable radio- 
active materials isolated from the snail 
was about 40 percent. The water con- 
tained numerous breakdown products 
of chlordimeform, clearly indicating the 
lability of this insecticide in the model 
ecosystem. 

Banamite®, or benzoylchloride-2,4,6- 
trichlorophenylhydrazone, is a new 
pesticide that has found use on citrus 
for the control of mites (Table 49). 
Only the crab (0.0156 ppm), aquatic 
plant (0.041 ppm), and mosquito 
(0.0736 ppm) contained residues of 
this pesticide. The EM values for bana- 
mite in these organisms were 839 for 


Inuinots NaturAL Hisrory SURVEY BULLETIN 


Vol. 31, Art. 9 


the crab, 2,204 for the aquatic plant, 
and 3,957 for the mosquito. The amount 
of banamite in the extractable radio- 
active materials from these organisms 
ranged from 1 to 2 percent. Though 
neither the fish nor the snail contained 
residues of banamite, they contained 
an unidentified degradation product, 
designated II, that was magnified about 
20,000 times in the snail and about 
3,000 times in the fish. It does not ap- 
pear that continued use of this pesticide 
will lead to problems of aquatic food- 
chain accumulation, but perhaps more 
detailed analysis of the chemical struc- 
ture of some of the degradative prod- 
ucts should be undertaken. 


ORGANOCHLORINE 
INSECTICIDE 
TEST RESULTS 


The organochlorines, especially the 
cyclodienes aldrin, heptachlor, and 
chlordane, have been used extensively 
in Illinois since they were introduced 
in 1954 for the control of underground 
insect pests of corn, particularly the 
corn rootworms Diabrotica longicornis 
and D. undecimpunctata howardi (Big- 
ger & Blanchard 1959). Their use as 
soil treatments increased from about 
125,000 acres (5.06 x 10* ha) treated 
in 1954 to a maximum of 5,601,572 acres 
(2.27 x 10° ha) treated in 1966 and 
slowly declined to about 2,100,000 acres 
(8.51 < 10° ha) treated in 1974 (Petty 
1974). The average treatment rate is 
about 1.6 pounds per acre (1.76 kg 
per ha) of technical material for aldrin 
and 2.0 pounds (2.2 kg per ha) for 
heptachlor (U.S. EPA 1972a). It is 
estimated that over the 20-year period 
more than 82 million pounds (3.73 X 
10’ kg) of these chemicals have been 
applied to Illinois farm soils (Illinois 
Natural History Survey data). The ap- 
proximate farm acreages treated with 
the organochlorine insecticides in IIli- 
nois are presented in Table 1 (Illinois 
Cooperative Crop Reporting Service 
1973). 

The use of cyclodiene insecticides in 
Illinois has been complicated by the 


Aug., 1975 =Mercaur & SANBORN: PESTICIDES AND ENVIRONMENTAL QuALITY 


invasion of the western corn rootworm, 
D. virgifera, which now covers nearly 
all of the cornland of Illinois and is 
totally resistant to the toxic action 
of aldrin, heptachlor, and chlordane 
(Petty & Kuhlman 1972), and by the 
unpredictability of attacks by the black 
cutworm, Agrotis ipsilon. 


ENVIRONMENTAL PERSISTENCE 

The organochlorine insecticides in 
use in Illinois are generally environ- 
mentally persistent or are readily con- 
verted to environmentally persistent 
compounds by photochemical or mi- 
crobial action or in vivo in the tissues 
of plants and animals. This is particu- 
larly true of the oxidation of aldrin 
to its 6,7-epoxide, dieldrin; heptachlor 
to its 2,3-epoxide, heptachlor epoxide; 
and the cis- and trans-chlordane isomers 
to oxychlordane. The average times 
required for 95-percent “breakdown” of 
these compounds in the soil has been 
estimated as: DDT, 11 years; dieldrin, 
9.7 years; lindane, 6.7 years; chlordane, 
4.2 years; heptachlor, 3.5 years; and 
aldrin, 2.5 years (Edwards 1965). 
Therefore, because of extremely heavy 
use patterns, it is no surprise to find 
that Illinois soils have been relatively 
highly contaminated by these com- 
pounds. The National Soils Monitoring 
Program (Carey et al. 1973) has re- 
ported these concentrations in Illinois 
soils: aldrin, 0.01-0.83 (average 0.07) 
ppm; chlordane, 0.05-1.32 (average 
0.09) ppm; dieldrin, 0.01-1.08 (aver- 
age 0.14) ppm; and DDT(T), 0.06— 
0.12 (average >0.01) ppm. These resi- 
dues were among the highest found in 
the United States. 

DDT, or 2,2-bis-(p-chlorophenyl)- 
1,1,1-trichloroethane, has the highest 
potential for bioaccumulation, 84,500- 
fold from water to fish, of any of the 
compounds studied (Metcalf et al. 
1971). This tendency to accumulate 
is the result of DDT’s low water solu- 
bility (0.0012 ppm) and its environ- 
mental stability. DDT also accumulates 
because of its partial conversion by 
dehydrochlorination to DDE, 2,2-bis- 
(p-chloropheny] )- 1, 1-dichloroethylene 


395 


(water solubility 0.0013 ppm). In the 
fish at the top of the food chain DDT 
constituted 34.3 percent, DDE 53.9 
percent, and DDD 9.8 percent of the 
absorbed total “C-radiolabeled material 
(Table 33). This fact demonstrates the 
gravest environmental flaw in the use 
of DDT, ie., the conversion to and 
storage in animal lipids of the highly 
persistent DDE. DDE constituted 52.0 
percent of the total radioactive ma- 
terials in the snail, 58.4 percent in the 
mosquito, and 54.0 percent in the fish. 
The percentage of unextractable radio- 
active materials in the various orga- 
nisms, a measure of total environmental 
stability, was low, ranging from 0.25 
percent in the mosquito to 13.5 percent 
in the alga, and averaging 3.9 percent 
for all test organisms. As shown in 
Table 34, DDE in the model ecosystem 
was degraded slowly and showed high 
ecological magnification. 

Because of its persistence, degrada- 
tion to the even more stable DDE, 
bioaccumulation, and effectiveness in 
inducing mircosomal oxidase enzymes 
(Peakall 1970), DDT has been banned 
as an insecticide by both the U.S. and 
Illinois Environmental Protection Agen- 
cies. The high degree of bioconcentra- 
tion and the preponderance of storage 
as DDE found in the model ecosystem 
study are representative of the values 
found in nature, e.g., fatty tissues of 
humans in the USA contain an average 
of about 2.3-4.0 ppm of DDT and 
4.3-8.0 ppm of DDE (Durham 1969). 
DDT in Lake Michigan at a concen- 
tration of 0.000006 ppm is biomagnified 
in lake trout to levels of 10-28 ppm 
(U.S. EPA 1972b), and in herring gulls 
to 99 ppm (Hickey et al. 1966). The 
lake trout residues averaged 53 per- 
cent DDE, 15 percent DDD, and 32 
percent DDT (U.S. EPA 1972b). DDT 
applied to a marsh in New Jersey for 
mosquito control was found in fish 
at 0.17-2.07 ppm and in gulls at 75 
ppm (Woodwell et al. 1967). 

DDD, or 2,2-bis-(p-chloropheny] )- 
1,1-dichloroethane, exhibited similar 
model-ecosystem behavior to that of 


396 


DDT (Table 35) and is, in fact, a 
degradative product of DDT (Table 
33). DDD constituted 58.9 percent of 
the total extractable radioactive ma- 
terials in the snail, 59.0 percent in the 
mosquito, and 85.4 percent in the fish 
(Metcalf et al. 1971). Thus, although 
DDD is a step on the degradative 
pathway of DDT and does not form 
the environmentally recalcitrant DDE, 
DDD seems to offer only slight im- 
provement over DDT in regard to en- 
vironmental hazard. Its ultimate fate 
in higher animals is conversion to and 
excretion as DDA (4,4’-dichlordiphenyl 
acetic acid), but this is an extremely 
slow process. DDD applied to Clear 
Lake, California, to control the Clear 
Lake gnat, Chaoborus astictopus, was 
found to be bioconcentrated through 
food chains from 0.02 ppm in the water 
to 903 ppm in the fat of plankton-eating 
fish and to 2,690 ppm in the fat of 
carnivorous fish (Hunt & Bischoff 
1960). 

Methoxychlor, or 2,2-bis- (p-methoxy- 
phenyl) -1,1,1-trichloroethane, differs 
from DDT in two important ways. It 
is 500 times more soluble in water, and 
the aryl CH,O groups (degradophores ) 
are readily biodegradable to OH 
groups, further increasing the polarity 
and water solubility. Thus, as shown 
in Table 36, methoxychlor is much less 
accumulative than DDT is in most ani- 
mals. Methoxychlor amounted to 84.0 
percent of the total extractable radio- 
active materials in the snail and 51.5 
percent in the fish. In contrast to the 
ready conversion of DDT to DDE 
(Table 33) and the storage of the latter 
in animal tissues, only very small 
amounts of the corresponding methoxy- 
chlor ethylene are stored by animals. 
The principal degradation pathway for 
methoxychlor is through conversion to 
the mono-OH and di-OH derivatives, 
which are readily converted to polar 
conjugation products in animals (Met- 
calf et al. 1971). 

Methoxychlor is classed as a mod- 
erately persistent insecticide and does 


Iuurnors NaturAL History SURVEY BULLETIN 


Vol. 31, Art. 9 


not accumulate to high levels in most 
animal tissues or milk. 


It offers a severe toxic hazard to 
fish but is degraded in fish much more 
readily than is DDT (Reinbold et al. 
1971). When used for control of the 
elm bark beetle, Scolytus multistriatus, 
vector of Dutch elm disease, methoxy- 
chlor has not resulted in environmental 
problems of transfer from earthworms 
to birds, as has DDT (Hunt & Sacho 
1969). 

Aldrin, or 1,2,3,4,10,10-hexachloro- 
1,4,4a,5,8,8a-hexahydro-1,4-endo, exo-5, 
8-dimethanonaphthalene, is rapidly con- 
verted in the model ecosystem and its 
organisms to the very persistent 6,7- 
epoxide, dieldrin (Table 37). In the 
model ecosystem treated with aldrin, 
dieldrin was stored as 85.7 percent of 
the total extractable radioactive ma- 
terials in the alga, 91.6 percent in the 
snail, and 95.8 percent in the fish (Met- 
calf et al. 1973). The bioaccumulation 
of both aldrin and dieldrin is high, 
directly proportional to their water in- 
solubility, but not as high as that of 
DDT and DDE. Only minor amounts 
of two degradation products, 9-keto 
dieldrin and 9-hydroxy dieldrin, were 
found, attesting to the stability of 
dieldrin, and these two products were 
also concentrated in the alga, snail, and 
fish. The ultimate degradative path- 
way is through trans-dihydroxydihydro 
aldrin. Aldrin, because of its rapid con- 
version to the highly persistent dieldrin, 
its bioaccumulation, and its carcino- 
genicity (Walker et al. 1973), has been 
banned as an insecticide by the U.S. 
Environmental Protection Agency. 

Dieldrin. When the model-ecosystem 
evaluation of dieldrin, the 6,7-epoxide 
of aldrin, was begun (Table 38), little 
difference was found between it and 
the evaluation of aldrin (Table 37). 
Dieldrin is slightly more water soluble 
than aldrin and exhibited slightly lower 
bioconcentrations in the fish. The sta- 
bility of dieldrin was shown by the 
storage of dieldrin as 98.7 percent of 
the extractable radioactive materials in 


Aug., 1975 Mercatr & SANBORN: PEstICIDES AND ENVIRONMENTAL QUALITY 


the alga, 99.0 percent in the snail, and 
97.8 percent in the fish (Sanborn & 
Yu 1973). However, 9-OH and 9-C—O 
dieldrin were identified as important 
degradation products along with trans- 
dihydroxydihydro aldrin. 

The several thousandfold accumula- 
tion of dieldrin in the fish of the model 
ecosystem following the application of 
aldrin is in agreement with observa- 
tions in nature. Humans in the USA 
have average values of 0.29-0.31 ppm 
of dieldrin in fatty tissues (Durham 
1969). Dieldrin in Lake Michigan at 
a concentration of 0.000002 ppm in 
water is biomagnified in lake trout to 
levels of 0.14-0.45 ppm (U.S. EPA 
1972b). The average bioconcentration 
of dieldrin from the waters of Illinois 
farm ponds to the tissues of fish was 
5,000- to 20,000-fold (W. F. Childers 
& W.N. Bruce, Illinois Natural History 
Survey, unpublished data). 

Toxaphene has been shown to be a 
mixture of at least 177 components 
(Holmstead et al. 1974) about two- 
thirds of which are C,,H,,Cl,, C,,H,, 
Cl,, and C,,H,Cl, compounds. The 
highly insecticidal components are hep- 
tachlorobornanes (Casida et al. 1974). 
The “C-radiolabeled toxaphene used in 
the model-ecosystem experiments was 
supplied by the manufacturer as the 
chlorination product of -{8-“C}- cam- 
phene to 67-69 percent Cl (sample 
X19093-4-2K ) and is presumably repre- 
sentative of the technical product. As 
shown in Table 39, the “C-radiolabeled 
toxaphene behaved in a surprisingly 
homogenous fashion in the extracts 
from the organisms of the model eco- 
system. The major ingredients referred 
to as “toxaphene” (R,; 0.70) were highly 
persistent and accumulated to several 
thousandfold levels in the organisms 
of the system. “Toxaphene” constituted 
82.6 percent of the total extractable 
radioactive materials in the alga, $6.6 
percent in the snail, 62.7 percent in the 
mosquito, and 64.9 percent in the fish. 
The unextractable “C-labeled materials 
averaged 19 percent of the total radio- 


397 


active materials in all of the organisms. 
Thus, toxaphene exhibited model-eco- 
system behavior rather like that of 
endrin (Table 40). 

The behavior of toxaphene in the en- 
vironment is little known because its 
enormous number of constituents poses 
almost insurmountable analytical prob- 
lems. Toxaphene in Big Bear Lake, 
California, at 0.2 ppm was found to be 
biomagnified to 200 ppm in goldfish 
(Hunt & Keith 1963), and in Lake 
Poinsett, South Dakota, from 0.001 ppm 
in the water to 0.176 ppm in the tissue 
and 1.152 ppm in the fat of the carp, 
Cyprinus carpio (Hannon et al. 1970). 
These instances of thousandfold bio- 
magnification are in perfect agreement 
with the model ecosystem results. 

Endrin is a highly water-insoluble 
pesticide that was also bioconcentrated 
in the organisms of the model ecosys- 
tem to a high degree (Table 40). 
Endrin, or 1,2,3,4,10,10-hexachloro-6,7- 
epoxy-1,4,4a,5,6,7,8,8a-octahydro-1,4- 
endo,endo-5,8-dimethanonap hthalene, 
is the endo,endo-isomer of dieldrin and 
is less environmentally persistent than 
dieldrin. Endrin was stored as 84.9 
percent of the total extractable “C- 
labeled materials in the alga, $3.0 per- 
cent in the snail, and 75.9 percent in 
the fish. Degradation appeared to be 
largely through an unknown compound 
designated II, probably 9-OH endrin 
in analogy with dieldrin. Unknown 
compound III is probably 9-C—O 
endrin (Metcalf et al. 1973). 

Biological observations on the orga- 
nisms of the system were particularly 
informative. Endrin was not only highly 
toxic to the salt-marsh caterpillar, which 
had difficulty consuming the treated 
sorghum leaves, but repeatedly killed 
all the daphnia, mosquito larvae, and 
fish in the aquatic portion of the sys- 
tem. The high toxicity of the water 
phase persisted for more than 60 days 
from the beginning of the experiment 
and occurred at endrin concentrations 
of 0.001-0.002 ppm. Because of this 
toxicity the experiment was extended 


398 


to nearly twice the usual 33-day period, 
and thus the data in Table 40 were 
measured after 63 days. Fish added to 
the model system had violent convul- 
sions within 10-15 minutes after being 
placed in the contaminated water. 
These biological observations demon- 
strated the substantial predictive value 
of the model-ecosystem investigations 
and could have given a preview of the 
Mississippi River fish kills associated 
with the leaching of endrin wastes 
(Barthel et al. 1969). Endrin, because 
of its great bioaccumulation, persist- 
ence, and extremely high toxicity to a 
wide variety of organisms, is a highly 
dangerous insecticide. 

Lindane, or gamma-l,2,3,4,5,6-hexa- 
chlorocyclohexane, has a higher water 
solubility than many of the other or- 
ganochlorine insecticides and appears 
to be less readily bioconcentrated in 
animal tissues (Table 41). In the model 
ecosystem lindane was stored as 20.6 
percent of the total extractable radio- 
active materials in the snail and 91.7 
percent in the fish. None could be 
detected in the alga or the mosquito. 
The principal degradation product ap- 
peared to be gamma-pentachlorocyclo- 
hexene. Lindane is substantially more 
biodegradable than DDT and the cyclo- 
diene pesticides, and it appears to be 
degraded environmentally to a series 
of trichlorophenols (Metcalf et al. 
1973). 

BHC residues have been found widely 
distributed in human fatty tissues in 
the USA at 0.20-0.60 ppm (Durham 
1969). The beta-isomer (an ingredient 
of technical BHC insecticide) is the 
most persistent isomer of lindane, and 
the environmental persistence of the 
gamma-isomer (lindane) is not well 
understood. 

Mirex, dodecachloro-octahydro-1,3,4- 
metheno-2H -cyclabuta--{c,d}--pentalene, 
was one of the least degradable com- 
pounds that we evaluated and was 
stored as 97.8 percent of the total ex- 
tractable radioactive materials in the 
alga, 99.4 percent in the snail, 99.6 per- 
cent in the mosquito, and 98.6 percent 


Iuuinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 9 


in the fish (Table 42) (Metcalf et al. 
1973). It is clearly a highly persistent 
pollutant and showed a substantial de- 
gree of bioaccumulation. Mirex is of 
environmental importance, as it is one 
of the most effective inducers of mi- 
crosomal oxidase enzymes. Mirex, fol- 
lowing its widespread use as a bait for 
the fire ant, has been found in tissues 
of wild birds at levels of up to 3 ppm 
and in rodents at nearly 20 ppm (Un- 
published data). It has also been found 
in tissues of northern pike and long- 
nose gar from Lake Ontario at 0.020— 
0.050 ppm (Kaiser 1974). 

Heptachlor, or 1-exo-4,5,6,7,8,8-hep- 
tachloro-3a,4,7,7a,-tetrahydro - 4,7-meth- 
enoindene, has a low level of water 
solubility and a high potentiality for 
biozccumulation (Table 43). Hep- 
tachlor is rapidly converted in the 
model ecosystem and its organisms to 
the very persistent 2,3-epoxide, hep- 
tachlor epoxide. In the model eco- 
system heptachlor epoxide was stored 
as 59.1 percent of the total extractable 
redioactive materials in the alga, 45.6 
percent in the snail, and 60.6 percent 
in the fish. These values are consider- 
ably lower than the corresponding 
values for the storage of dieldrin after 
the treatment of crops with aldrin 
(Table 37) and reflect the existence 
of an alternate degradative pathway 
in heptachlor, the replacement of the 
1-Cl atom by OH to give 1-hydroxy- 
chlordene. This degradative product 
is more polar and water soluble than 
heptachlor and is not as highly ac- 
cumulative. It can also be epoxidized 
in vivo to the 2,3-epoxide, 1-hydroxy- 
chlordene epoxide, which was found 
stored in the snail, mosquito, and fish. 
This latter~degradative product could 
also be formed by hydrolysis of hepta- 
chlor epoxide. Heptachlor epoxide in 
the model ecosystem (Table 44) 
showed a persistence comparable to 
that of dieldrin (Table 38). 

In the heptachlor test the unextrac- 
table “C-labeled materials averaged 29 
percent of the total radioactive ma- 
terials in the various organisms. Hepta- 


Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


chlor epoxide is widely distributed in 
the environment, and the average level 
in the body fat of humans in the USA 
is 0.1-0.24 ppm (Durham 1969). Yel- 
low perch from Lake Michigan had 
heptachlor epoxide body residues rang- 
ing from 0.060 to 0.097 ppm (U.S. EPA 
1972b). Heptachlor and_heptachlor 
epoxide are under surveillance by the 
U.S. EPA because of their carcinogen- 


' icity (Carter 1974). 


Chlordane, or 1,2,4,5,6,7,8,8-octa- 
chloro -3a, 4,7, 7a-tetrahydro-4, 7 -metha- 
noindane, is chemically related to hep- 
tachlor except that the double bond 
has been chlorinated. The behavior of 
this insecticide in the model ecosystem 
clearly demonstrates its persistence and 
tendency to accumulate in the orga- 
nisms of this system (Table 45). The 
water of the model ecosystem contained 
only 5.98 percent chlordane, but the 
alga, snail, mosquito, and fish contained 
94.51, 91.17, 47.64, and 77.86 percent, 
respectively, of their radioactive ma- 
terials as chlordane. The EM values 
for chlordane for the alga, snail, mos- 
quito, and fish were 98,386, 132,613, 
6,132, and 8,261, respectively. Clearly, 
the continued use of chlordane, along 
with its minor contaminant, heptachlor, 
will lead to problems of accumulation 
in food chains, which can lead to resi- 
dues of these two pesticides in humans. 
Unpublished data accumulated by fed- 
eral monitoring agencies have indicated 
that 95 percent of the adipose tissue 
taken from humans in the United States 
contains residues of heptachlor. Fur- 
ther, nearly 70 percent of U.S. poultry, 
fish, and dairy products contain resi- 
dues of heptachlor. The data of this 
model-ecosystem experiment provide 
background information which explains 
the high incidence of heptachlor resi- 
dues in humans and food. 


FUNGICIDE 
TEST RESULTS 


Captan, or N-trichloromethylthio-4- 
cyclohexene-1,2-dicarboximide, is thc 
most versatile of the general foliar 


399 


fungicides for the treatment of fruits 
and vegetables. In the model eco- 
system it was found to be extensively 
degraded, producing at least 15 degra- 
dation products in the water phase 
(Table 46). No intact captan was 
identified in any of the organisms of 
the system, and only trace amounts of 
degradation products were found. Cap- 
tan appears not to offer any environ- 
mental problems following normal use. 

Hexachlorobenzene has had some 
use as a fungicide in seed treatment, 
replacing in part the organomercurial 
fungicides. In the model system it was 
extremely persistent and substantially 
bioaccumulative, the parent compound 
comprising 85.1 percent of the total 
extractable radioactive materials in the 
alga, 87.2 percent in the daphnia, 58.3 
percent in the mosquito, and 27.7 per- 
cent in the fish (Table 47) (Metcalf 
et al. 1973). EM values ranged from 
144 to 1,248. The degradation of hexa- 
chlorobenzene occurs through hydroly- 
sis to pentachlorophenol and other 
chlorophenols of increasing water solu- 
bility. 

Hexachlorobenzene used as a fun- 
gicide on wheat caused an epidemic 
of thousands of cases of cutaneous 
porphyrinuria in humans in Turkey 
(Schmid 1960), and the compound has 
been found in human tissues nearly 
everywhere, ranging up to 0.29 ppm 
in adipose tissues in Great Britain 
(Abbott et al. 1972). Hexachloroben- 
zene is clearly an undesirable environ- 
mental pollutant. 

Pentachlorophenol is the fungicide in 
largest scale use in the United States 
as a timber and paper pulp preservative 
and mildewproofer. It is also used as 
a soil and timber poison against ter- 
mites and as a nonselective herbicide. 
In the model ecosystem pentachloro- 
phenol accumulated in the various or- 
ganisms to a moderate degree (Table 
48). EM values were 5-205. Penta- 
chlorophenol constituted 15.1 percent 
of the total extractable radioactive ma- 
terials in the alga, 12.2 percent in the 
snail, 33.3 percent in the mosquito, 


400 


55.5 percent in daphnia, and 51.2 per- 
cent in the fish. It is apparently de- 
graded through a series of chlorinated 
phenols, and 10 degradation products 
were found in the water phase. 
Pentachlorophenol, because of its 
high toxicity to nearly all forms of life 
as an oxidative phosphorylation un- 
coupler and its stability, can be a 
dangerous environmental pollutant. Its 
use as an herbicide in Japan has re- 
sulted in its presence in almost all 
Japanese river waters at concentrations 


of 0.01-0.1 ppb (Goto 1971). 


DISCUSSION 


The data shown in the preceding 
tables, illustrating the fates of a variety 
of pesticides in the laboratory model 
ecosystem, can be used for predictive 
purposes in a number of ways. 


BIOLOGICAL EFFECTS 

The dosages applied in the model 
ecosystem are realistic in terms of those 
used in the field, ie., 0.2-1.0 pound 
per acre (0.22-1.1 kg per ha). There- 
fore, the biological results observed are 
meaningful as predictors of the en- 
vironmental impact of the pesticide 
studied. The most dramatic results on 
nontarget species were found with the 
organochlorine insecticides endrin, diel- 
drin, and heptachlor epoxide. Endrin 
applied at the equivalent of 0.2 pound 
per acre (0.22 kg per ha) repeatedly 
killed all daphnia and mosquitoes in 
the system, and the necessity for re- 
stocking delayed the termination of the 
experiment to over 60 days. Fish added 
to the endrin system showed violent 
convulsions within 10-15 minutes and 
died within a few hours. Similar re- 
sults were experienced with heptachlor 
epoxide, which killed daphnia and mos- 
quitoes for 56 days after having been 
applied at 0.2 pound per acre (0.22 kg 
per ha). Dieldrin was highly toxic 
to daphnia and mosquitoes, which did 
not survive at any time during the ex- 
periment. 

Temephos, the highly effective mos- 
quito larvicide, killed mosquito larvae 


TIutinois NATuRAL Hisrory SURVEY BULLETIN 


so persistently that the experiment was 
prolonged to 53 days. Chlorpyrifos and 
methyl chlorpyrifos even at the 1.0-mg 
dosage were highly toxic to daphnia, 
and chlorpyrifos adversely affected 
algae. 

The carbamate insecticides carbaryl 
and carbofuran were extremely toxic 
to daphnia in the initial stages of the 
experiments. 


Some of the herbicides, especially 
metrabuzin and bifenox, were highly 
toxic to algae in the model ecosystem. 
Surprisingly, the insecticide methoxy- 
chlor, or its degradation products, also 
affected algae adversely. 


DEGRADATIVE PRODUCTS 


This parameter is, of course, the di- 
rect measure of biodegradability. In 
general, the larger the number of 
degradative products in the water and 
in the organisms of the model eco- 
system, the lower the degree of eco- 
logical magnification and the higher 
the amount of unextractable radioac- 
tive materials. Thus, DDE with two 
degradation products and DDT with 
four were the worst offenders in eco- 
logical magnification in contrast to 
temephos, carbaryl, and metrabuzin, 
each with 11 degradative products, and 
chlordimeform with 13; each of the 
latter four compounds showed zero 
ecological magnification. Clearly, the 
relationship is not precise, because the 
variety of positions of radiolabeling 
limits the extent to which degradative 
products can be identified. Moreover, 
the formation of secondary toxicants, 
such as the epoxides, e.g., dieldrin 
from aldrin and heptachlor epoxide 
from heptachlor, provides products that 
are substantially more environmentally 
stable and ecologically magnified than 
are the parent compounds. 


Nevertheless, knowledge of the key 
degradative products of any pesticide 
is important in characterizing its en- 
vironmental impact. The model eco- 
system not only provides useful in- 
formation about the chemical nature 
of degradation products and about 


Vol. 31, Art. 9 


Aug., 1975 Merrca.r & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


degradative pathways, but also indi- 
cates potential rates and locations of 
storage and bioconcentration of pesti- 
cides and their degradation products. 
As examples, in addition to those of 
dieldrin and heptachlor epoxide, Bana- 
mite (Table 49) produced an unidenti- 
fied degradation product, designated 
II, which was ecologically magnified 
3,013-fold in fish and 19,824-fold in 
snails. Metrabuzin (Table 12) pro- 
duced an unidentified product, desig- 
nated II, which was ecologically mag- 
nified 175-fold in fish. Even the highly 
degradable malathion produced an 
unidentified product, designated II, 
which showed apparent ecological mag- 
nification of about 19,500-fold (Table 
20). 


ECOLOGICAL MAGNIFICATION 


The accumulation of lipid-soluble, 
water-insoluble pesticides in living or- 
ganisms is one of the most disturbing 


401 


pesticides. The laboratory model eco- 
system is particularly suitable for de- 
termining “ecological magnification,” or 
the pesticide concentration in an or- 
ganism divided by the pesticide con- 
centration in the water. When eco- 
logical magnification is considered for 
the fish (Gambusia), we find that the 
values from the data in the tables vary 
from 0 to 10°. Such ecological mag- 
nification is a function of the partition 
coefficient in lipid/water and the sta- 
bility of the pesticide and its metabo- 
lites in the animal. As shown in Fig. 
2, an effective approximation is ob- 
tained when the water solubility of 
the pesticide in parts per billion (ppb) 
is plotted as a log function against eco- 
logical magnification. There is clearly 
an inverse relationship, with the least 
water-soluble pesticides accumulating 
to the highest degree. This relationship 
is highly significant, with a correlation 


features of environmental pollution by coefficient of r — —0.76, and it is sub- 
107 : , 
<= : : 
n : : 
re : : 
! : : 
Zz : 
2 
ke : : 
105 : : 
S Pe 
rs 4 : : 
z : : 
= o “Oo : 
= o : 
"op @ "8 og 
< : @ 0% © 
5 © eo ® 
rss | : © © : 
=! : : 
3 X : 
uw 10 : © 2: 


103 
WATER SOLUBILITY ppb 


102 


10* 10 


® ont 
10S ~=107 =«108 = 09 


Fig. 2.—The relationship between the water solubility of pesticides, numbered as _in 
Tables 2-49, and the ecological magnification of parent compounds in the mosquito fish 
in the laboratory model ecosystem. A highly significant correlation (r = —0.76) exists. 


402 


stantially predictable. Thus, it is of 
great importance to know the water 
solubility of even the least soluble com- 
pounds. From this information it is 
possible to make a reliable estimate 
of the potentialities of new pesticides 
to accumulate in the tissues of fish and 
other aquatic’ organisms. Our study 
suggests a classification of pesticides as: 
1. water solubility<0.5 ppm, likely 
to be environmentally hazardous 
2. water solubility>50 ppm, likely 
to be environmentally nonhazard- 
ous 
3. water solubility from 0.5 to 50 
ppm, to be used with caution 
The lines of demarcation between the 
three classes obviously are not sharp, 
and the ultimate hazard also depends 
upon lipid partitioning, the rapidity 
of pesticide degradation in living ani- 
mals, use patterns, and amounts ap- 
plied. However, practical experience 
has already shown that most of the 


ECOLOGICAL MAGNIFICATION IN FISH 


Inuinors Natura History SURVEY BULLETIN 


Vol. 31, Art. 9 


pesticides with water solubilities of 
<0.5 ppm demonstrate bioaccumula- 
tion following field use and that most 
of those with water solubilities of >50 
ppm have not shown bioaccumulation. 
The large group of pesticides with 
water solubilities between 0.5 and 50 
ppm represent those which may demon- 
strate bioaccumulation under some con- 
ditions of use, e.g., in lakes or oceans 
with very cold water. Their use pat- 
terns should be judged accordingly. 


UNEXTRACTABLE 
RADIOACTIVE MATERIALS 


This parameter measures the conver- 
sion of the pesticide under investigation 
and its primary degradation products 
into simple degradation products which 
enter the metabolic pool of an orga- 
nism and are resynthesized into normal 
tissue ingredients. The percentage of 
unextractable radioactive materials can 
be determined for many of the pesticides 
investigated by adding the amount of 


PERCENTAGE OF UNEXTRACTABLE '4c IN FISH 


Fig. 3.—The relationship between the percentage of radioactive materials extractable from 
the mosquito fish of the laboratory model ecosystem and the total body accumulation of 


parent pesticide, numbered as in Tables 2-49, and all of its degradation products. 
is a highly significant correlation (r = —0.74). 


There 


Aug., 1975 Mercaur & SANBORN: PEsTICIDES AND ENVIRONMENTAL QUALITY 


unextractable radioactive materials to 
the total extractable radioactive mate- 
rials and determining the fraction. The 
values obtained in the fish (Gambusia), 
for example, range from 0.34 percent 
for DDE to about 90 percent for feni- 
trothion. As shown in Fig. 3, a highly 
significant correlation (r= —0.74) ex- 
ists between the percentage of unex- 
tractable radioactive materials and the 
in vivo stability of the pesticide and its 
principal degradation products as mea- 
sured by the total biomagnification of 
the radioactive materials from the water 
to the fish (or other organism). 


Considering that two different 


403 


methods for determining amounts of un- 
extractable radioactive materials were 
used, ie., total combustion analysis 
and solubilization, the results are sur- 
prisingly predictable. Clearly, pesti- 
cides and their degradation products 
which are highly lipid soluble in 
the tissues of organisms are almost 
quantitatively extractable and leave 
small amounts of unextractable radio- — 
active materials. As a tentative guide- 
line we suggest that pesticides which 
produce 40 percent or more of unex- 
tractable radioactive materials in the 
fish in the model ecosystem evaluation 
will not be likely to cause serious prob- 
lems with environmental quality. 


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PESTICIDES AND ENVIRONMENTAL QUALITY 


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Vol. 31, Art. 9 


It.tinois NATURAL History SuRVEY BULLETIN 


406 


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407 


PESTICIDES AND ENVIRONMENTAL QUALITY 


Aug., 1975 Mercatr & SANBORN 


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408 Intinors Natura History SuRvEY BULLETIN Vol. 31, Art. 9 


Table 7.—Rr values and amounts, in parts per million, of phenmedipham* and its degrada- 
tion products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
R;> Water (alga) (snail) (mosquito) (fish) 

Total 14C 0.028 4,22 2.69 1.312 0.545 
Phenmedipham eet 

Ac 0.64 0.0102 

B! tracet ia ore ee 

rT: 0.99 oi: 0.067 0.497 0.131 

II 0.96 aes 0.131 0.153 0.101 

Ill 0.76 8 oe 0.139 

IV 0.68 Siem 0.372 

Vv 0.35 nave 0.355 

VI 0.18 siete 0.506 5 meas ais 
Origin 0.00 0.0178 2.65 2.04 1.080 0.545 
Unextractable 4C 0.018 13.08 7.00 1.978 0.535 


a Methyl m-hydroxycarbanilate m-methylearbanilate, “C-ring UL. 

> Silica Gel GF-254, diethyl ether:petroleum ether :chloroform, 6:3:1 by volume. 
¢ A = N-(3-hydroxypheny1) -methy1! urethane. 

4B = 3-methylaniline. 

e Roman numerals indicate compounds whose chemical structures are unknown. 
f Determined by gas chromatography. 


Table 8—Rr values and amounts, in parts per million, of 2,4-D* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Elodea Physa Gambusia 
R? Water (alga) (aquatic plant) (snail) (fish) 
Total 4C 0.2048 5.498 2.752 0.757 0.0454 
Ie 0.97 Ser: 0.282 0.178 0.285 
II — 0.89 Bat 1.030 0.456 ane 
III 0.80 viet 0.477 0.477 0.301 
IV 0.65 eae 0.377 83 ror Ae 
V 0.58 0.0000641 0.295 ra aid 0.0431 
VI 0.63 0.00269 
VII 0.56 0.00212 
VIII 0.49 0.00226 
IX 0.39 0.000417 snd ae 
x 0.10 0.000474 1.675 0.768 
XI 0.067 0.000271 on = oes 
Origin 0.00 0.000185 1.362 0.873 0.171 0.00226 


Unextractable 4C 0.012 17.625 7.555 6.421 0.211 


a 2,4-dichlorophenoxyacetic acid, “C-ring UL. 
» Silica Gel GF-254, benzene-dioxane-acetic acid, 90:25:4 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


409 


PESTICIDES AND ENVIRONMENTAL QUALITY 


Aug., 1975 Metcatr & SANBORN 


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Aug., 1975 MercaLr & SANBORN: PESTICIDES AND ENVIRONMENTAL Quatiry 411 


Table 11.—Rr values and amounts, in parts per million, of trifluralin* and its degradation 
products found in the water and organisms of a model ecosystem. 


Daphnia Physa Culex Gambusia 
Rr Water (water flea) (snail) (mosquito) (fish) 

Total 4C 0.0489 0.445 6.663 0.238 0.767 
Trifluralin 0.74 0.000282 5.046 ae 0.261 

Ie 0.51 0.000066 Sieve 

At 0.39 0.000087 0.337 

Be 0.32 0.000374 bate 

II 0.24 0.000322 0.0399 

Iil 0.20 0.000139 0.216 

IV 0.17 0.000514 aa 

G 0.13 0.000803 oad 

V 0.11 0.000686 were aa 

VI 0.07 0.00141 Age 0.228 

VII 0.04 0.00203 BC Sa Sere 
Origin 0.00 0.0169 aie 0.796 wate 0.506 
Unextractable 4C 0.0253 1.017 6.648 0.520 1.011 


4 q,a,a-trifluoro-2,6-dinitro-N, N-dipropyl-p-toluidine, “C-ring UL. 
» Silica Gel GF-254, hexane-acetone-methanol, 90:10:2 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


4 A = q,a,a-trifluoro-2,6-dinitro-N-propyl-p-toluidine. 


¢ B = 2,b-dinitro-4-trifluoromethy] aniline. 


tC = 2-ethyl-5-trifluoromethyl-7-nitrobenzimidazole. 


Table 12.—Rer values and amounts, in parts per million, of metrabuzin* and its degradation 
products found in the water and organisms of a model ecosystem. 


Total 4C 
Ic 


XI 
Origin 
Unextractable 4C 


Rr? 


0.87 
0.83 
0.57 
0.35 
0.32 
0.24 
0.20 
0.17 
0.12 
0.09 
0.06 
0.04 
0.01 
0.00 


Water 


0.6524 
0.2911 
0.003118 
0.008965 
0.1292 
0.006435 
0.09676 
0.0005374 
0.001252 
0.005675 
0.001158 
0.00005275 
0.0005046 
0.002047 
0.005278 
0.1003 


Physa Culex Gambusia 
(snail) (mosquito) (fish ) 
1.2880 1.559 1.342 
0.762 1.307 0.307 
0.3920 oie 0.546 
0.291 
0.0746 
0.05217 0.05158 0.0140 
0.05217 0.2407 0.0769 
0.4578 3.7546 0.3498 


a 4-amino-6-tert-butyl-3-(methylthio) -as-triazin-5 (4H )-one 5-4C, 
> Chloroform: acetone, 9:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 
4 A = Desamino metrabuzin. 
¢ B = Desmercapto metrabuzin. 

fC = Desamino desmercapto metrabuzin. 


412 Txiinors NaTurAL History Survey BULLETIN Vol. 31, Art. 9 


Table 13.—Rr values and amounts, in parts per million, of bifenox* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 

Ri? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.0376 3.267 1.650 0.290 0.211 
ie 0.82 0.000108 sate 3.0 0.071 elie 
Bifenox 0.73 0.000745 3.189 1.203 0.219 0.156 
A‘ 0.66 0.000125 a 0.256 Sore rate 

II 0.60 0.000092 oe ee ett hate 
Til 0.45 0.000125 Be lege non aint 

B* 0.33 0.000325 e3 0.088 aan 0.020 
Origin 0.00 0.0301 0.78 0.103 trace 0.026 


Unextractable 4C 0.0061 


4 Methy1-5-(2’,4’-dichlorophenoxy ) -2-nitrobenzoate, “#C nitrophenyl ring UL. 

b Silica Gel GF-254, benzene :dioxane :acetic acid, 90:30:1 by volume. 

¢ Roman numerals indicate compounds whose chemical structures are unknown. 
a A = Methyl1-5-(2’,4’-dichlorophenoxy )-2-amino benzoate. 

e 2,4-dichlorophenoxy-2-nitro-5-benzoic acid. 


Table 14.—Rr values and amounts, in parts per million, of chlorpyrifos" and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 

Ry? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.00056 0.0261 0.158 0.063 0.0711 

Chlorpyrifos 0.76 0.00011 0.0079 0.076 0.005 0.0352 
A‘ 0.60 eh 538 ee a ab 

Ba 0.60 0.000115 0.0051 0.051 0.022 0.0207 

Origin 0.00 0.00005 0.0131 0.031 0.036 0.0152 

Unextractable 4C 0.00028 0.1507 0.0456 ne 0.0223 


4 0,0-diethyl-O-(3,5,6-trichloro-2-pyridyl) -phosphorothionate, “C-ring UL. 
b Silica Gel GF-254, benzene :dioxane:acetic acid, 90:15:1 by volume. Pyridinol (Rr — 0.17) 


and P=O ester (Rr — 0.90) separated with solvent system, acetonitrile :-hexane :acetone :NH,OH 
70:10:15 :5, by volume. 


¢ A = Chlorpyrifosoxon. 
4B = 3,5,6-trichloro-2-pyridol. 


Table 15.—Rr values and amounts, in parts per million, of chlorpyrifos-methyl* and its 
degradation products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 

R:? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.00262 0.0780 0.0882 0.220 0.0367 

Chlorpyrifos-methyl 0.75 0.00008 0.0382 0.0435 0.15 0.0076 
AC 0.54 aa. eg ork Ste Red 

Be 0.60 0.0002 0.0087 0.0082 0.037 0.0109 

Origin 0.00 0.00154 0.0051 0.0067 0.033 0.0101 

Unextractable 4C 0.0009 0.4703 0.2110 aeete 0.0398 


8 0,0-diethyl-O-(3,5,6-trichloropyridinyl) phosphorothionate, “C-ring UL. 
» Silica Gel GF-254, benzene: dioxane :acetic acid, 90:15:1 by volume. Pyridinol (Rr — 0.17) 


and P=O ester (Rr — 0.83) separated by solvent system, acetonitrile :hexane :acetone :NH,OH, 
70:10:15 :5, by volume. 


¢ A = Chlorpyrifosoxon methyl. 
4B = 3,5,6-trichloro-2-pyridol. 


Aug., 1975 Mercaur & SANBORN: PESTICIDES AND ENVIRONMENTAL Quauity 413 


Table 16.—Rsr values and amounts, in parts per million, of Counter®" and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
R? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.00259 0.1063 0.1556 0.1517 0.0427 

Te 0.83 0 aie aor 0.1156 says 
Counter@®) 0.77 0.00002 0.0035 0.0366 0.0072 0.0107 

At 0.66 0.00006 Aas Bac 

Be 0.54 0.000062 trace 0.0241 

ct 0.40 0.000357 st 

II 0.29 0.000046 0.0071 

III 0.18 0.000018 Sc eke 

Dé 0.13 0.000017 0.0106 trace 

IV 0.03 0.000007 0.0213 ere says iaid 
Origin 0.00 0.000283 0.0638 0.0494 0.0289 0.0320 
Unextractable 4C 0.00174 0.8913 0.778 0.4282 0.0813 


a 0,0-diethyl S-(tert-butylthio)-methyl phosphorodithioate, “C-tert-butyl. 
> Silica Gel GF-254, benzene :acetone, 4:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


iS) 
4A = (C,H;O):2 PSCH,S0.C (CHg)s. 
* B= (C2H;0).2 BSCH,SC(CH)s 
tC = (C.H;0)2 PSCH,S0,C (CHs)s.- 
=D = (C,H;0)>. bscESOC(CHs)s. 


Table 17.—Rsr values and amounts, in parts per million, of temephos* and its degradation 
products found in the water and organisms of a model ecosystem. 


x0 (0) ¥ ©) 0z 


Oedogonium Physa Gambusia 
x AG Z Re Water (alga) (snail) (fish) 

Total 3H 0.000280 0.00991 0.09161 0.00099 

(MeO).P=S Ss (MeO).P=S 0.0000013 0.00195 0.01876 

(MeO),P=S SO (Me0O).P=S 0.000002 0.00066 0.01483 

(MeO).P=S SO, (MeO).P=—S 0.0000007 0.00078 0.00396 

(MeO).P = O Ss (MeO),.P =O 0.00000014 0.00127 0.00785 

(MeO).P =O SO, (MeO).P=—O 0.00066 Sia 

(MeO).P—S SO, (MeO),.P=O note 0.00040 0.00698 

(MeO),.P=S Ss (MeO).P = O trace 0.00066 0.02705 

(MeO).P—S So, H 0.000002 0.00036 

(MeO).P=S s H 0.000002 0.00129 

H Ss H 0.000001 0.00045 pti 

H so, H 0.0000024 0.00046 0.00175 

(MeO),P =O Ss H 0.000008 0.00064 0.01178 

Origin 0.00019 0.00033 0.00436 

Unextractable 4C 0.000070 


2 0,0-dimethylphosphorothioate ester of 4,4’ dihydroxydiphenyl sulfide, *H-ring-labeled. 
» Silica Gel GF-254 three dimensional TLC: 1.toluene. 2.methanol :chloroform :toluene, 10:95: 


$5 by volume. 3.nitromethane :acetonitrile, 2 


9:65 :110 by volume. 


414 Intinors NATURAL History Survey BULLETIN Vol. 31, Art. 9 


Table 18.—Rr values and amounts, in parts per million, of fonofos* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusiu 
Rr? Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.1079 0.2977 0.2831 0.6863 0.08500 
Fonofos 0.92 0.0008866 0.09556 0.07635 0.6133 0.06845 
Ic 0.76 0.0000653 0.02203 
II 0.68 0.0002602 ahs 
Ill 0.62 0.001504 0.07905 
IV 0.37 0.002806 0.01887 
Vv 0.29 0.0005997 
vI 0.22 0.0008081 
Vil 0.19 0.0001472 
Vill 0.13 0.0003656 Pte 
IX 0.10 0.0001383 0.0401 
x 0.09 0.0000765 A rau ee 
XI 0.08 0.0002311 é 0.08748 ars 0.01193 
XII 0.04 0.0005152 0.01494 Ee aeho 
Origin 0.00 0.008277 0.02712 0.1193 0.07302 0.004624 
Unextractable 4C 0.09126 0.5247 2.2550 5.8578 0.2453 


“ Q-ethyl, S-phenyl ethylphosphonodithioate, “C-O-ethyl. 
> Silica Gel GF-254, chloroform :ethyl acetate, 4:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 19.—Rr values and amounts, in parts per million, of fenitrothion® and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
R:? Water (alga) (snail) (mosquito) (fish) 
Total 22P 0.1136 2.5579 5.270 0.0829 0.0545 
Ic 0.92 0.00238 ee BAS bee See 
Fenitrothion 0.81 0.00247 0.8632 saz 0.0055 0.0242 
II 0.73 0.00004 ers eke ae pers 
A‘ 0.52 0.00088 seks Fees oe 0.0057 
III 0.22 0.00030 
IV 0.13 0.00338 
Vv 0.06 0.00030 seek oe deh ae 
Origin 0.00 0.02438 1.2947 5.2700 0.0774 0.0246 
Unextractable 4C 0.07949 10.5993 1.5802 1.0983 8.9550 


* 0,O-dimethyl O-(3-methyl-4-nitrophenyl) phosphorothionate, #P. 

> Silica Gel GF-254, hexane (Skellysolve B) :ether, 4:1 by volume. 

© Roman numerals indicate compounds whose chemical structures are unknown. 
4A = Fenitroxon or dimethyl 3-methyl-4-nitrophenyl phosphate. 


Aug., 1975 Mercaur & SANEORN: PEsTICIDES AND ENVIRONMENTAL QUALITY 


415 


Table 20.—Rsr values and amounts, in parts per million, of malathion* and its degradation 
products found in the water and organisms of a model ecosystem. 


Ri? 
Total 4C 
1k 0.81 
II 0.75 
Ill 0.63-0.67 
IV 0.50 
Vv 0.36 
VI 0.31 
VII 0.15 
VIII 0.05 
Origin 0.00 


Oedogonium Physa Culler Gambusia 

Water (alga) (snail) (mosquito) (fish) 
0.01659 0.421 0.577 6.97 1.43 

eae 0.319 ans 1.82 ints 
0.0000447 0.338 3 oie 0.119 
0.0000335 2.34 0.655 
0.0000546 0.947 0.1033 
0.0000784 0.299 mies 

eae 0.0254 
0.0000345 an 0.0737 
0.0000754 yaa an6 0.275 0.0342 
0.003868 0.102 0.139 1.283 0.420 
0.0124 


Unextractable 4C 


a 0,O-dimethyl-S-(1,2-dicarboethoxyethyl) -phosphorodithioate, “C-O0,0-methyl. 
> Silica Gel GF-254, benzene :acetic acid, 4:1 by volume. 


© Roman numerals indicate compounds whose chemical structures are unknown. 


Vol. 31, Art. 9 


Intrors NaTurAL History SuRVEY BULLETIN 


416 


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Aug., 1975 Merrcaur & SANBORN: PESTICIDES AND ENVIRONMENTAL QuaLity 417 


Table 22.—Rr values and amounts, in parts per million, of leptophos* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Gambusia 
R Water (alga) (snail) (fish) 

Total 4C 0.180 31.637 53.696 1.866 
Leptophos 0.93 0.00108 13.221 52.270 1.559 

1 0.85 15.753 Bae aye 

II 0.25 0.105 0.0313 

Ill 0.24 sass 0.0235 

IV 0.22 ae 0.128 

V 0.20 sere 2.357 

VI 0.13 0.002712 

VII 0.12 0.00647 — 

VIII 0.10 0.000199 

IX 0.09 0.00392 

x 0.07 0.0000691 seers 

XI 0.05 0.009094 0.009 sists 

XII 0.03 0.000147 Bod des 0.0235 
Origin 0.00 0.02170 0.297 1.193 0.199 
Unextractable 4C 0.1351 57.241 11.612 1.555 


a O-(4-bromo-2,5-dichlorophenyl)-O-methyl phenylphosphonothionate, “C-O-methyl. 
> Silica Gel GF-254, benzene :chloroform, 1:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 23.—Rr values and amounts, in parts per million, of parathion® and its degradation 


products found in the water and organisms of a model ecosystem. 


Oedogonium Daphnia Physa Culex Gambusia 
R,? Water (alga) (water flea) (snail) (mosquito) (fish) 

Total 4C 0.003 0.3969 0.2987 0.2701 0.2031 0.1935 

ie 0.97 0.000200 0.0356 sisi 
Parathion 0.90 0.00030 0.1006 

Il 0.73 0.000060 ase 

A‘ 0.55 0.000136 0.0086 

III 0.33 0.00025 5 0.0222 

Be 0.25 0.0047 

IV 0.13 0.00049 

V 0.09 0.00274 ; Sr ahr aoe Bie 
Origin 0.00 0.00599 0.3613 0.2987 0.2701 0.2081 0.0621 
Unextractable 4C 0.0854 2.6284 0.3126 0.5818 0.4685 0.2055 


8 0,0-diethyl O-4-nitrophenyl phosphorothionate, “C-ring-2,6. 


> Silica Gel GF-254, ether-hexane, 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


aA = p-nitrophenol. 
¢ B = Paraoxon. 


7:3 by volume. 


. 31, Art. 9 


Iturino1s NaTurAL History SurvEY BULLETIN Vol 


418 


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419 


Prsvicipres AND ENVIRONMENTAL QUALITY 


Aug., 1975 Merca.r & SANBORN 


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Vol. 31, Art. 9 


Ituinois Natura History SurvEY BULLETIN 


420 


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Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL Quatiry 421 


Table 27.—Rzr values and amounts, in parts per million, of propoxur* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium 


R:? 

Total 4C 
it 0.92 
A‘ 0.74 
Propoxur 0.64 
Be 0.50 
II 0.38 
Ct 0.22 
Ill 0.10 
IV 0.08 
Origin 0.00 


Unextractable 4C 


Water 


0.00408 
0.000083 
0.00032 
0.000032 
0.00001 
0.000006 
0.00001 
0.000012 
0.00106 
0.00255 


(alga) 


0.4617 
0.2150 


0.0360 
0.0249 
0.0598 
0.1260 
3.9357 


Physa 
(snail) 
0.3946 
0.1330 
0.0406 
0.0928 
0.0236 


0.0300 


0.0746 
6.1600 


a 2-isopropoxyphenyl N-methylcarbamate, 4C-2-isopropoxy. 
b Silica Gel GF-254, chloroform :acetonitrile, 4:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


a A = 2-isopropoxyphenol. 


e B = 2-isopropoxyphenyl carbamate. 
tf C = 2-isopropoxyphenyl N-hydroxymethyl carbamate. 


Culer 
(mosquito) 


2.2913 
0.4312 


0.4441 
1.1520 


0.2640 
21.900 


Gambusia 
(fish) 


0.1173 
0.0252 
0.0468 
0.0180 


0.0273 
0.1053 


Table 28.—Rr values and amounts, in parts 
per million, of aldicarb* and its degradation 
products found in the water and organisms of 


a model ecosystem. 


Total 4C 
Aldicarb® 
Ad 


Origin 


R:? 


Culex 
(mos- 
Water quito) 


Gam- 
busia 
(fish) 


0.54 
0.42 
0.28 
0.14 
0.00 


0.16 17.0 
0.031 16.7 
trace 

0.04 

0.056 

0.025 0.3 


2.32 
1.31 
1.01 


a 2-methyl-2-methylthiopropionaldoximyl N- 
methylearbamate, 14C-tert-carbon. 
b Silica Gel GF-254, hexane: benzene: etha- 


nol, 2:2:1 by volume. 
¢ CH3SC (CH;) «CH = NOC (0) NHCHs. 


4 A = CH;S0.C (CH,),CH = NOH. 
¢ B = CH,SO.C (CH;)2CH = NOC (0) NHCHs. 
£C = CH,SOC(CH;)2CH = NOC(0O) NHCH3. 


422 Ittinors NaTuRAL History SuRvVEY BULLETIN Vol. 31, Art. 9 


Table 29.—Rr values and amounts, in parts per million, of formetanate” and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
R? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.11 44.98 2.10 1.61 1.17 

Ic 0.81 aes Sore 1.53 1.07 

II 0.75 SA 2.25 ae 

III 0.62 otis nae 0.32 

A‘ 0.35 0.0666 

IV 0.27 eis 2.70 

Vv 0.14 ae 4.05 ire Shave es 
Origin 0.00 0.0118 35.98 0.25 0.54 1.17 
Unextractable 4C 0.0316 22.10 9.02 5.59 1.71 


a 3-dimethylaminomethyleneiminophenyl N-methylcarbamateehydrochloride, “C-ring labeled. 
» Silica Gel GF-254, ethyl acetate. 

¢ Roman numerals indicate compounds whose chemical structures are unknown. 

4 A = N-formyl-3-aminophenol. 


Table 30.—Rr values and amounts, in parts per million, of methoprene* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Gambusia 
R, Water (alga) (snail) (fish) 
Total 4C 0.00556 4.626 4.885 0.070 
Ic 0.83 are 0.0990 0.1924 ays 
Methoprene" 0.76 0.000086 2.220 1.500 0.0176 
II 0.66 ciate 0.963 0.376 0.0305 
Ae 0.60 Si5 EA 1.5490 sr 
Bt 0.53 sisi 0.723 0.469 0.0181 
Ce 0.47 0.000075 re 0.0845 0.0017 
Other 0.00024 0.332 0.500 
Origin 0.00 0.000576 0.289 0.45 0.0021 


Unextractable 4C 0.00458 


« Tsopropyl-11-methoxy-3,7,11-trimethyldodeca-2,4-dienoate. 

» Silica Gel GF-254, benzene:ethyl acetate :acetic acid, 100:50 35 by volume. 

© Roman numerals indicate compounds whose chemical structures are unknown. 
4 Tsopropyl-11-methoxy-3,7,11-trimethyldodeca-2,4-dienoate (5-"4C), 

¢ A = 11-methoxy-3,7,11-trimethyldodeca-2,4-dienoic acid. 

f B = Isopropyl 11-hydroxy-3,7,11-trimethyldodeca-2,4-dienoate. 

&’ C = 11-hydroxy-3,7,11-trimethyldodeca-2, 4-dienoic acid. 


423 


PESTICIDES AND ENVIRONMENTAL QUALITY 


Aug., 1975 Mercatr & SANBORN 


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424 Iniinois NaTurAL History SuRvEY BULLETIN Vol. 31, Art. 9 


Table 32.—Rr values and amounts, in parts per million, of chlordimeform* and its degrada- 
tion products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
Ri? Water (alga) (snail) (mosquito) (fish) 

Total 4C 1 2 0.0427 0.9253 1.7682 0.3626 0.5259 
Chlordimeform 0.00, 0.65 er) vets 0.0710 Fe 

A‘ 0.33, 0.09 0.00075 Seve anes wags ests 

Bt 0.43, 0.00 0.00179 nae a at ais 

ce 0.53, 0.35 0.00016 eat 

Dt 0.10, 0.00 0.00085 0.109 35 tas. Reis 

EF 0.77, 0.71 0.00041 0.0933 0.255 %, 0.0553 

1 0.83, 0.00 0.00052 0.181! 

II 0.83, 0.66 0.00031 

III 0.73, 0.70 0.00026 aie 

IV 0.57, 0.90 a Sc a Re 0.0246 

Vv 0.53, 0.03 0.00070 fe. uke am 

VI 0.52, 0.27 0.000077 

VII 0.50, 0.23 0.00017 

VIII 0.33, 0.23 0.00025 

IX 0.23, 0.00 0.00077 5 oe Sees Ani 
Origin 0.00, 0.00 0.0125 0.542 1.442 Bh 0.446 
Unextractable 4C 0.0233 


a N’-(4-chloro-o-tolyl)-N,N-dimethylforamidine, “4C-tolyl. 
» Silica Gel GF-254 two dimensional tle: 1. benzene :dioxane :acetic acid, 90:30:1 by volume. 
2. benzene :diethylamine, 95:5 by volume. 


¢ A = 2-methyl-4-chloroformanilide. 

4B = 5-chloroanthranilic acid. 

¢ C = 2-methyl-4-chloroaniline. 

t D = 2-carboxy-4-chloroformanilide. 

& EH = 2,2’-dimethyl-4, 4’-dichloroazobenzene. 

h Roman numerals indicate compounds whose chemical structures are unknown. 
! Alga contained traces of unknowns totaling 0.181 ppm. 


Table 33.—Rr values and amounts, in parts per million, of DDT* and its degradation 
products found in the water and organisms of a model ecosystem. 


Physa Culex Gambusia 
R;? Water (snail) (mosquito) (fish ) 
Total 4C 0.004 22.9 8.9 54.2 
DDE 0.53 0.00026 12.0 5.2 29.2 
DDT 0.34 0.00022 7.6 1.8 18.6 
DDD 0.17 0.00012 1.6 0.4 5.3 
Origin 0.00 0.0032 0.98 1.5 0.85 


2 2,2-bis-(p-chlorophenyl]) -1,1,1-trichloroethane, “C-ring UL. 
» Silica Gel GF-254, petroleum ether solvent, b.p. 60—80°C, 
© Dry weight. 


Table 34.—Rr values and amounts, in parts per million, of DDE* and its degradation 
products found in the water and organisms of a model ecosystem. 


Physa Culex® Gambusia 
R, Water (snail) (mosquito) (fish ) 
Total 4C 0.008 121.6 168.9 149.8 
DDE 0.53 0.0053 103.5 159.5 145.0 
Origin 0.0 0.0027 18.1 9.4 4.8 


® 2,2-bis-(p-chloropheny1) -1,1-dichloroethylene, C-ring UL, 
» Silica Gel GF-254, petroleum ether solvent, b.p. 60--80°C. 


pe: se 


Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL Quatity 425 


Table 35.—Rzr values and amounts, in parts per million, of DDD" and its degradation 
products found in the water and organisms of a model ecosystem. 


Physa Culex Gambusia 

Ry Water (snail) (mosquito) (fish ) 

Total 4C 0.006 5.65 5.85 39.12 

A‘ 0.53 nie.» 0.24 wh 2.08 

Ie 0.47 af 0.14 BCS 1.54 
DDD 0.17 0.0004 3.3 3.43 33.4 

II 0.05 Ac 0.87 aren ae 
Origin 0.00 0.0056 ileal Ba 2.0 


a 2,2-bis-(p-chloropheny1) -1,1-dichloroethane, “C-ring UL. 

b Silica Gel GF-254, hexane (Skellysolve B). 

¢ A = CICgHyC = CCl.»CgHiCl. 

4@Roman numerals indicate compounds whose chemical structures are unknown. 


Table 36.—Rr values and amounts, in parts per million, of methoxychlor* and its degrada- 
tion products found in the water and organisms of a model ecosystem. 


Physa Culex® Gambusia 

Ry? Water (snail) (mosquito) (fish ) 

Total 3H 0.0016 ay 0.48 0.33 
Ad 0.32 Nae 0.7 sth hod 
Methoxychlor 0.25 0.00011 13.2 shin 0.17 
Be 0.07 0.00013 1.0 Aeie trace 

if 0.00 0.00003 trace nels trace 
Ds 0.00 0.00003 eke ake 
Unknowns trace 0.00009 trace Hat trace 
Origin 0.00 0.00125 0.8 see 0.16 


a 2,2-bis-(p-methoxypheny]) -1,1,1-trichloroethane, *H-ring labeled. 
b Silica Gel GF-254, petroleum ether solvent, b.p. 60—-80°C. 

¢ Dry weight. 

4 A = CH,OCsHC = CCl,CsH,OCHs3. 

¢ B = CH;0C,H,HCC1;C;H,OH. 

£C = HOC,HyHCC1;,C,H,OH. 

& D => HOC,H.C = CCl,CsH,OH. 


Table 37.—Rzr values and amounts, in parts per million, of aldrin* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 


RR Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.0117 19.70 57.20 1.13 29.21 
Aldrin 0.81 0.00005 1.95 2.23 ae 0.157 
Dieldrin 0.71 0.0047 16.88 52.40 1.10 28.00 
ie 0.63 sake 0.57 2.05 oe cc 0.612 
Aa 0.45 0.00052 0.12 0.17 oe 0.322 
Be 0.34 0.0004 0.079 0.217 ates 0.088 
ct 0.08 0.00039 0.015 aks — aie 
Origin 0.00 0.0040 0.015 0.097 Bone 0.004 


Unextractable 4C 0.00155 


_ #1,2,3,4,10,10-hexachloro-1,4,4a,5,8,8a-hexahydro-1,4-endo, exo-5,8-dimethanonaphthalene, uC. 
ring. 
b Silica Gel GF-254, n-hexane :diethyl ether, 1:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 
a A = 9-hydroxy dieldrin. 
eB = 9-keto dieldrin. 


Vol. 31, Art. 9 


Intinois NATURAL History SURVEY BULLETIN 


426 


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Aug., 1975 Mercar & SANBORN: PESTICIDES AND ENVIRONMENTAL Qua.ity 427 


Table 39.—Rr values and amounts, in parts per million, of toxaphene" and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 

Ri? Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.04441 13.2941 17.6198 2.2570 10.3977 
Toxaphene 0.70 0.00159 10.9743 15.2637 1.4147 6.7523 
Ic 0.57 0.00106 1.7535 1.8360 0.2359 2.4923 
II 0.51 0.00076 Slave 0.2961 aan 0.5022 
III 0.45 0.00099 0.3589 0.0863 one 0.3161 
IV 0.34 0.00164 0.1130 0.0585 trace 0.1487 

V (strip) 0.00429 Ae Nate 0.4042 ote 
VI 0.03 0.00078 dese re tas 0.0187 
Origin 0.00 0.02002 0.0944 0.0211 0.2022 0.1674 
Unextractable 4C 0.01328 2.2156 1.1153 1.1245 4.2264 


8 CyoHioCls (67-69% chlorinated camphene), 8-14C. 
» Silica Gel GF-254, Skellysolve B (b.p. 68°C) : diethyl ether :acetone, 80:20:10 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 40.—Rr values and amounts, in parts per million, of endrin* and its degradation 
products found in the water and organisms of a model ecosystem”. 


Oedogonium Physa Culex" Gambusia 
Rio Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.0135 13.62 150.58 ass 4.48 
fe 0.81 ne 0.48 5.07 ea an 
Endrin 0.73 0.00254 11.56 125.00 mice 3.40 
II 0.53 0.00885 1.58 6.55 Pitas 1.04 
III 0.42 trace trace 5.87 
IV 0.31 trace trace 2.69 Syous ene 
Origin 0.00 0.00436 See 1.85 sree 0.04 


Unextractable 4C 0.0027 


8 1,2,3,4,10,10-hexachloro-6,7-epoxy-1,4,4a,5,6,7,8,8a-octahydro-1,4-endo, endo-5,8-dimethano- 
naphthalene, C-ring. 


> Experiment terminated after 63 days. 

¢ Silica Gel GF-254, n-hexane :diethyl ether, 1:1 by volume. 

4 Mosquito larvae killed throughout experiment. 

¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 41.—Rr values and amounts, in parts per million, of lindane* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
Ri? Water (alga) (snail) (mosquito) (fish) 

Total 4C 0.0232 0.375 3.70 0.75 1.02 

AS 0.55 Boe 560 2.50 rete 3350 
Lindane 0.47 0.00167 ss 0.762 36x 0.935 

if 0.27 0.000084 

II 0.19 0.00304 

III 0.14 0.00276 x6 305 

IV 0.09 0.00636 avale 0.248 Saxe Sjere 
Origin 0.00 0.00877 0.375 0.185 Sa 0.085 


* gamma-1,2,3,4,5,6-hexachlorocyclohexane, “C-ring. 

» Silica Gel GF-254, n-hexane-acetone, 9:1 by volume. 

¢ A = gamma-pentachlorocyclohexene. 

4 Roman numerals indicate cempounds whose chemical structures are unknown. 


428 Intinors NaturAL History Survey BULLETIN Vol. 31, Art. 9 


Table 42.—Rr values and amounts, in parts per million, of mirex* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
R? Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.018 9.70 18.40 13.60 3.50 
Mirex 0.95 0.0157 9.49 18.29 13.54 3.45 
Origin 0.00 0.0023 0.21 0.11 0.06 0.05 


a Dodecachloro-octahydro-1,3,4-metheno-2-H-cyclabuta-[c,d]-pentalene, “C-ring. 
b Silica Gel GF-254, chloroform. 


Table 43.—Rr values and amounts, in parts per million, of heptachlor* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
RR Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.02225 0.8448 2.7515 3.1258 2.0603 
Heptachlor 0.64 0.00003 0.6219 1.1146 0.9421 0.1146 
Heptachlor epoxide 0.56 0.00021 0.1877 1.0659 1.5332 1.6293 
Ie 0.43 0.00002 Bt 0.0217 0.0434 
II 0.37 0.00001 icane 0.1142 0.0328 
Ill 0.32 0.00005 a 0.0490 0.0244 Same 
1-hydroxychlordene 0.21 0.00040 sheus 0.0597 0.0791 0.0471 
1-hydroxychlordene 
epoxide 0.14 0.00659 sats 0.2066 0.2694 0.1211 
IV 0.07 0.00036 ve 0.0272 0.0763 0.1010 
V 0.03 0.00026 on 0.0055 0.0244 ake 
Origin 0.00 0.00677 0.0352 0.0871 0.1007 0.0472 
Unextractable 4C 0.00755 0.4079 0.1646 0.2363 1.5479 


4 1-exo0-4,5,6,7,8,8-heptachloro-3a,4,7,7a-tetrahydro-4,7-methanoindene, “C-ring. 
b Silica Gel GF-254, cyclohexane :diethyl ether, 80:20 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 44.—Rr values and amounts, in parts per million, of heptachlor epoxide* and its 
degradation products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex® Gambusia 
Ry? Water (alga) (snail) (mosquito) (fish) 
Total 4C 0.00638 2.2620 101.9105 ae 8.8807 
Heptachlor epoxide 0.63 0.00125 2.0618 83.0774 seat 6.1100 
Aa 0.18 0.00036 0.0800 8.8663 at 1.7114 
Origin 0.0 0.00200 0.1202 9.9668 aos 1.0595 
Unextractable 4C 0.00277 1.1602 0.1110 e.5 0.8554 


* |-exo-4,5,6,7,8,8-heptachloro-exo-2,3-epoxy-3a,4,7,7a-tetrahydro-4,7-methanoindane, ™“C-ring. 
> Silica Gel GF-254, cyclohexane, diethylether, $0:20 by volume. 

¢ All killed during the experiment. 

4 A = 1-hydroxychlordene epoxide. 


——— 


Aug., 1975 Mercarr & SANBORN: PEsTICIDES AND ENVIRONMENTAL QuaLity 429 


Table 45.—Rr values and amounts, in parts per million, of chlordane* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Physa Culex Gambusia 
Ri? Water (alga) (snail) (mosquito) (fish) 
Total 4c 0.017718 110.3415 154.189 13.645 11.243 
if 0.92 a 0.974 2.199 sa 0.451 
II 0.90 sere 0.664 oar ies aoe 
Ill 0.84 0.000059 1.905 5.751 3.602 1.405 
IV 0.78 a 1.708 ae 5.502 ae 
Vv 0.73 airehs ore tas Meld 0.658 
Chlordane+ 0.70 0.00106 104.289 140.570 6.500 8.754 
VI 0.64 sere irr 1.541 
VII 0.55 0.0000135 Preva 0.244 
VIII 0.47 0.0000027 he ae 538 
IX 0.28 0.00127 0.474 2.088 0.509 0.217 
xX 0.23 0.000176 SS Rate Bate sae 
XI 0.19 0.0000939 ae feted ae 0.0260 
XII 0.17 0.000264 iat Sine See exfice 
XIII 0.15 0.000415 0.358 0.478 0.339 0.0223 
XIV 0.12 0.000438 0.180 0.398 iste ee 
XV 0.10 Sate ate 0.884 Brea 0.0744 
XVI 0.06 0.000689 
XVII 0.04 0.000501 
XVIII 0.03 0.009305 or 
XIX 0.02 ar 0.325 
xX 0.01 ate 0.238 oe ae Sate 
Origin 0.00 0.0123 0.324 1.516 2.169 0.205 
Unextractable 4C 0.00752 100.0847 1.752 6.920 2.450 
cis: trans 4.02 3.08 5.39 iets 6.98 


41,2,4,5,6,7,8,8-octachloro-3a,4,7,7a-tetrahydro-4,7-methanoindane (cis:trans, 3:1), “C-ring. 
b Silica Gel GF-254, n-hexane :ethyl acetate, 9:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


430 Iuuno1s NATURAL History SURVEY BULLETIN Vol. 31, Art. 9 


Table 46.—Rxr values and amounts, in parts per million, of captan* and its degradation 
products found in the water and organisms of a model ecosystem. 


Oedogonium Daphnia Physa Culex Gambusia 
R,? Water (alga) (water flea) (snail) (mosquito) (fish) 
Total 4C 0.001789 0.865 0.393 0.301 0.0462 0.0522 
Ts 0.93 aie 0.0278 cee 0.0592 
II 0.85 ives 0.0077 tats 0.0795 wit sists 
Ill 0.81 eer aes ware 0.0679 wont 0.0492 
IV 0.79 Bs 0.0166 
V 0.68 S50 
VI 0.39 ee 0.0105 
VII 0.35 0.00000426 eaha 
VIII 0.33 ae 0.0142 
IX 0.26 0.0000096 ane 
x 0.25 0.00000893 0.0608 
XI 0.18 0.00000456 
XII 0.14 0.0000109 Ns 
XIII 0.10 0.00000365 0.590 ats oat Pas a 
XIV 0.053 0.0000891 0.0159 dR, 35. sic 0.00215 
Origin 0.00 0.0000353 0.122 3s 0.0940 sates 0.000861 
Unextractable 4C 0.001623 0.967 0.338 0.0998 0.0584 0.0158 


4 N-trichloromethylthio-4-cyclohexene-1,2-dicarboximide, “C-trichloromethyl. 
» Silica Gel GF-254, petroleum ether-acetone, 4:1 by volume. 
¢ Roman numerals indicate compounds whose chemical structures are unknown. 


Table 47.—Rr values and amounts, in parts per million, of hexachlorobenzene" and its 
degradation products found in the water and organisms of a model ecosystem. 


Oedogonium Daphnia Physa Culex Gambusia 

R, Water (alga) (water flea) (snail) (mosquito) (fish) 

Total 4C 0.00695 1.827 0.696 4.098 0.737 3.155 

Hexachlorobenzene 0.80 0.00298 1.556 0.598 3.72 0.429 0.857 

AS 0.50 0.00034 aan eke care as Aa 

Ja 0.10 0.00023 Bae Herc ne ap 0.446 

II 0.05 ward Wee ae 0.0385 0.857 

Origin 0.00 0.000143 0.271 0.098 0.378 0.269 0.995 
Unextractable 4C 0.00197 


“1,2,3,4,5,6-hexachlorobenzene, “C-ring UL. 

» Silica Gel GF-254, benzene :acetone, 1:1 by volume. 

¢ A = Pentachlorophenol. 

“Roman numerals indicate compounds whose chemical structures are unknown. 


431 


PESTICIDES AND ENVIRONMENTAL QUALITY 


Aug., 1975 Mercatr & SANBORN 


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LITERATURE CITED 


Assott, D. C., G. B. Cottins, and R. Gouxp- 
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556. 

Azovu-Don1a, M. B., M. A. OrHMmaAN, G. 
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AHARONSON, N., and A. Ben-Aziz. 1974. 
Persistence of residues of Velsicol VCS- 
506 and two of its metabolites in tomatoes 
and grapes. Journal of Agricultural and 
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AnonyMous. 1974. EPA refuses to raise 
permissible dieldrin level for contami- 
nated chickens. Pesticide Chemical News 
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BartHet, W. F., J. C. Hawruorne, J. H. 
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E. H. Grissincer, and D. A. Parsons. 
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Biccer, J. H., and R. A. BLANCHARD. 1959. 
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BootnH, G. M., C. C. Yu, and D. J. HANSEN. 
1973. Fate, metabolism, and toxicity of 
3-isopropyl-1H-2,1,3-benzothiadiazin-4 
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411. 

Carzy, A. E., G. B. Wiersma, H. Tat, and 
W. G. MitcHeELL. 1973. Pesticides in soil. 
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Carter, L. J. 1974. Cancer and the environ- 
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Casipa, J. E., R. L. Hotmsteap, 8S. KHALIFA, 
J. R. Knox, T. Osawa, K. J. PAcMer, and 
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a complex biodegradable mixture. Science 
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DurHAmM, W. H. 1969. Body burden of 
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Epwarps, C. A. 1965. Effects of pesticide 
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Fowter, D. L., and J. N. Manan. 1972. 
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Agriculture, Agricultural Stabilization 
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433 


Freeman, L. 1953. A standardized method 
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Goro, M. 1971. Organochlorine compounds 
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Hannon, M. R., Y. A. GrercHus, R. L. 
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Hickey, J. J., J. A. Kerry, and F. B. Coon. 
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Lake Michigan ecosystem. Pages 141-154 
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Hotmsteap, R. L., T. R. FuKuto, and R. B. 
Marcu. 1973. The metabolism of O-(4- 
bromo-2,5-dichlorophenyl) O-methyl phe- 
nylphosphonothioate (leptophos) in white 
mice and on cotton plants. Archives of 
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cology 1:133-147. 

, S. Kwara, and J. E. Casmpa, 1974. 
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939-944. 

Hunt, E. G., and A. I. BiscHorr. 1960. 
Clinical effects on wildlife of periodic 
DDD applications to Clear Lake. Cali- 
fornia Fish and Game 46:91-106. 

_ and J. O. KeirH. 1963. Pesticide- 
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1962. Proceedings of the Second Con- 
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Hunt, L. B., and R. J. Saco. 1969. Re- 
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chlor. Journal of Wildlife Management 
33:336-345. 

ILLINOIS COOPERATIVE Crop REPORTING SER- 
vick. 1970. Pesticide use by Illinois 
farmers, 1969. Illinois Department of 
Agriculture and U.S. Department of Agri- 
culture Bulletin 70-4. Springfield, Illinois. 

1973. Illinois pesticide use by Illi- 

nois farmers 1972. Illinois Department 

of Agriculture and U.S. Department of 

Agriculture Bulletin 73-3. Springfield, 

Illinois. 


434 


Katser, K. L. BE. 1974. Mirex: an un- 
recognized contaminant of fishes from 
Lake Ontario. Science 185:523-525. 

Ketiy, R. G., E. A. Peers, S. Gorpon, and 
D. A. Buyske. 1961. Determination of 
C4 and H? in biological samples by 
Schéniger combustion and liquid scintil- 
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McGuiamery, M. D., E. Knaxe, and F. W. 
Suire. 1974. 1974 field crops weed control 
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school. ‘Cooperative Extension Service, 
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Mercatr, R. L., T. R. Fuxuto, C. CoLiins, 
K. Borck, J. Burk, H. T. ReyNo.ps, and 
M. F. Osman. 1966. Metabolism of 2- 
methyl - 2 4 (methylthio)- propionaldehyde 
O-(methylearbamoyl)-oxime in plant and 
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1971. Model ecosystem for the evaluation 

of pesticide biodegradability and eco- 

logical magnification. Environmental Sci- 

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3 ie 2P) KAPOOR, Pe Ye plo: Coase 

ScuurH, and P. SHERMAN. 1973. Model 

ecosystem studies of the environmental 

fate of six organochlorine pesticides. En- 

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PeakaLL, D. B. 1970. p,p’-DDT: effect on 
calcium metabolism and concentration of 
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594. 


Perry, H. B. 1974. Soil insecticide use in 
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, and D. E. Kuntman, 1972. Root- 
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fourth Illinois custom spray operators 
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Agriculture in cooperation with the Illi- 
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TIuurnors NaturAL History SuRVEY BULLETIN 


Vol. 31, Art. 9 


M. J. Forster, I. N. Oxa, O. D. SHOLEs, 
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Progst, G. W., and J. B. Tere. 1969. Tri- 
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Quistap, G. B., L. E. Sraicer, and D. A. 
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of Agricultural and Food Chemistry 22: 
582-589. 


; , and . 1975. Environ- 
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Rergoip, K. A., I. P. Kapoor, W. F. 
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degradability of DDT and methoxychlor 
by aquatic organisms. Illinois Natural 
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Sangorn, J. R., and C. C. Yu. 1973. The 
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Bulletin of Environmental Contamination 
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Scumip, R. 1960. Cutaneous porphyria in 
Turkey. New England Journal of Medi- 
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4F)-11-methoxy-3,7,11-trimethyl-2,4-do- 
decadienoate). II. Metabolism by aquatic 
microorganisms. Journal of Agricultural 
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Sura, K. P. 1974. Nerve damage. The re- 
turn of the “ginger jake?” Environment 
16 (9) :6—-10. 

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1972a. Pesticide use on the nonirrigated 
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Series 4, TS-00-72-08. Washington, D.C. 


1972b. An evaluation of DDT and 
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Von RiimxKer, R., and F. Horay. 1972. 
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i 
a 
Ki 
fe 

; 


Aug., 1975 Mercatr & SANBORN: PESTICIDES AND ENVIRONMENTAL QUALITY 


WALKER, A.I.T., E. THorPr, and D. HE. 
STEVENSON. 1973. The toxicology of diel- 
drin (HEOD). I. Long-term oral toxicity 
studies in mice. Food and Cosmetics 
Toxicology 11:415—-432. 


Wiersma, G. B., H. Tar, and P. F. Sanp. 
1972. Pesticide residue levels in soils, 
FY 1969-National Soils Monitoring Pro- 
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6:194—-228. 

WoopweLL, G. M., C. F. Wurster, Jr., and 
P.A. Isaacson. 1967. DDT residues in an 
East Coast estuary: a case of biological 


435 


concentration of a persistent insecticide. 
Science 156:821—-824. 


Yu, C. C., G. M. Boorn, D. J. HANsEN, and 
J. R. Larsen. 1974. Fate of carbofuran in 
a model ecosystem. Journal of Agricul- 
tural and Food Chemistry 22:431-434. 

, D. J. Hansen, and G. M. Booru. 

1975a. Fate of dicamba in a model eco- 

system. Bulletin of Environmental Con- 

tamination and Toxicology 13:280—-283. 

, G. M. Boorn, D. J. Hansen, and 

J. R. Larsen. 1975b. Fate of pyrazon 

in a model ecosystem. Journal of Agri- 

cultural and Food Chemistry 23:309-311. 


INDEX 


A 


Abate@®, 390, 413 

Acephate, 391, 416 

Alachlor, 386, 404 

Aldicarb, 393, 421 

Aldrin, 383, 384, 394, 395, 396, 425 
Atrazine, 386, 405 


B 
Banamite@®, 394, 452 
Benefit-risk of pesticide use, 383 
Bentazon, 386-387, 405 
Bifenox, 388-389, 412 
Biological effects, 400 


Cc 
Captan, 382, 399, 430 
Carbamate insecticide test results, 392 
Carbaryl, 392, 419 
Carbofuran, 382, 392, 420 
Chlordane, 382, 384, 389, 394, 395, 399, 429 
Chlordimeform, 394, 424 
Chlorpyrifos, 389, 412 
Chlorpyrifos-methyl, 389, 412 
Corn rootworms, 394, 395 
Counter@®, 389-390, 413 
Crop yields in Illinois, 381 
Cyanazine, 387, 406 


D 
DDD, 395-396, 425 
DDE, 395, 396, 403, 424 
DDT, 395, 396, 424 
Degradation products, 400-401 
Diabrotica 
longicornis (see corn rootworms) 
undecimpunctata howardi (see corn root- 
worms) 
virgifera (see corn rootworms) 
Dicamba, 387, 407 
Dieldrin, 382, 383, 384, 395, 396, 426 
Dimilin, 393-394, 423 
Dyfonate®, 390, 414 


E 
Early-warning technology, 383-384 
Ecological magnification vs. water solubil- 
ity, 401-402 
Endrin, 397-398, 427 


F 
Fenitrothion, 390, 403, 414 
Fonofos, 390, 414 
Formetanate, 393, 422 
Fungicide test results, 399 


H 
Heptachlor, 384, 389, 394, 395, 398-399, 428 
Heptachlor epoxide, 389, 395, 398-399, 428 
Herbicide test results, 386 
Herbicide use, 381-383, 386 
Hexachlorobenzene, 399, 430 


I 
Insecticide use, 381-382, 383 
Introduction of new pesticides, 382 


L 
Leptophos, 391, 417 
Lindane, 395, 398, 427 


M 
Malathion, 391, 415 
Maneb, 382 
Metalkamate, 392, 418 
Methoprene, 393, 422 
Methoxychlor, 396, 425 
Metrabuzin, 388, 411 
Mirex, 398, 428 
Miscellaneous insecticide test results, 393 
Mixtures of herbicides used in Illinois, 383 
Mixtures of pesticides, 383 
Model-ecosystem technology, 385-386 


Oo 


Organochlorine insecticide persistence, 395 

Organophosphorus insecticide test results, 
389 

Orthene@®, 391, 416 

Oxychlordane, 395 


P 
Parathion, 391-392, 417 
Pentachlorophenol, 399-400, 431 
Phenmedipham, 387, 408 
Phosvel@®, 391, 417 
Propachlor, 387-388, 409 
Propanil, 382 
Propoxur, 392-393, 421 
Pyrazon, 388, 410 


T 
Temephos, 390, 413 
Toxaphene, 382, 397, 427 
Trifluralin, 382, 388, 411 
2,4-D, 387, 408 


U 


Unextractable radioactive materials and 
ecological magnification correlation, 403 


Zz 
Zineb, 382 


436 


Some Publications of the ILLINOIS NATURAL HISTORY SURVEY _ 


BULLETIN 


Volume 31, Article 3—Nutritional Respon- 
ses of Pheasants to Corn, with Special 
Reference to High-Lysine Corn. By Ron- 
ald F. Labisky and William L. Anderson. 
July, 1973. 26 p., index. 


Volume 31, Article 4—An Urban Epiphy- 
totic of Phloem Necrosis and Dutch Elm 
Disease, 1944-1972. By J. Cedric Carter 
and Lucile Rogers Carter. May, 1974. 31 
p., index. 


Volume 31, Article 5.—Larvae of the Seri- 
cothripini (Thysanoptera: Thripidae), 
with Reference to Other Larvae of the 
Terebrantia, of Illinois. By Thomas C. 
Vance. August, 1974. 64 p., index. 


Volume 31, Article 6.—Root Infection of 
Woody Hosts with Verticillium albo- 
atrum. By Gerald L. Born. August, 1974. 
41 p., index. 


Volume 31, Article 7—The Mecoptera, or 
Scorpionflies, of Illinois. By Donald W. 
Webb, Norman D. Penny, and John C. 
Marlin. August, 1975. 66 p., index. 


Volume 31, Article 8—An HElectrofishing 
Survey of the Illinois River, 1959-1974. 
By Richard BH. Sparks and William C. 
Starrett. August, 1975. 64 p., index. 


BIOLOGICAL NOTES 


88.—Illinois Birds: Laniidae. By Richard 
R. Graber, Jean W. Graber, and Ethelyn 
L. Kirk, June, 1973. 18 p. 


84.—Interactions of Intensive Cultures of 
Channel Catfish with Largemouth Bass in 
1-Acre Ponds. By D. Homer Buck, Rich- 
ard J. Baur, and C. Russell Rose. Febru- 
ary, 1974. 8 p. 


85.—The Literature of Arthropods Associ- 
ated with Soybeans. III. A Bibliography 
of the Bean Leaf Beetle, Cerotoma trifur- 
cata (Forster) and C. rujicornis (Olivier) 
(Coleoptera: Chrysomelidae). By M. P. 
Nichols, M. Kogan, and G. P. Waldbauer. 
February, 1974. 16 p. 


86.—Illinois Birds: Tyrannidae. By Rich- 
ard R. Graber, Jean W. Graber, and 
Ethelyn L. Kirk. February, 1974. 56 p. 

87.—The Literature of Arthropods Associ- 
ated with Alfalfa. I. A Bibliography of 
the Spotted Alfalfa Aphid, Therioaphis 
maculata (Buckton) (Homoptera: Aphi- 


dae). By D. W. Davis, M. P. Nichol 
E. J. Armbrust. February, 1974. — 


88.—The Literature of Arthropods As 
ated with Alfalfa, II. A Biblio 
the Sitona Species (Coleoptera: 
lionidae). By W. P. Morrison, B. 
M. P. Nichols, and E. J. Arne 
ruary, 1974. 24 p. 


89.—The,Life History of the aise 
er, Etheostoma squamiceps, in 
Illinois, and Ferguson Creek, K 
By Lawrence M. Page. May, 1974. 
90.—A Bibliography of the Northert 
Rootworm, Diabrotica longicorn 
and the Western Corn Rootworm, 
brotica virgifera LeConte (Co 
Chrysomelidae). By W. H. Luel 
H. C. Chiang, EB. E. Ortman, and | 
P. Nichols. April, 1974. 15 p. 


91.—The Distribution of Periodical 
in Illinois. By Lewis J. St 7 
February, 1975. 12 p. a - 
t 


92.—The Literature of Arthropods 
ated with Soybeans. IV. A Biblio; 
of the Velvetbean Caterpillar Ar 
gemmatalis Hiibner (Lepidoptera: 
tuidae). By B. J. Ford, J. R. y 
Reid, and G. L. Godfrey. ro 
15 p. 

93.—The Life History of the Str 
Darter, Etheostoma kennicotti, 
Creek, Illinois. By Lawrence M. 
February, LOT GD, 


94.—Illinois Pheasants: Their Distr 
and Abundance, 1958-1973. By Ronald 
Labisky. February, 1975. 11 p. 


95.—The Nest Biology of the Bee An 
(Ptilandrena) erigeniae. Robertson - 
menoptera: Andrenidae). By Lio} 
Davis, Jr. and Wallace E. Lalor 
1975. 16 p. 


CIRCULAR 
51.—Illinois Trees: Selection, Plantin 
Care. By J. Cedric Carter. August, | 2 
123 p. 


52.—Fertilizing and Watering Trees, 
Dan Neely and E. B. Himelick. De 
ber, 1971. (Third printing.) 20 p. 


54—Corn Rootworm Pest Manage’ 
Canning Sweet Corn. By W. Hi.) 
mann, J. -T. Shaw, D. BE. Kuhlm 
Randell, and C. D. LeSar. March 
10 p. 


List of available publications mailed on request 


No charge is made for publications of the ILt1noIs NATURAL History Survey. A Si 
copy of most publications will be sent free to anyone requesting it until the supply be 
low. Costly publications, more than one copy of a publication, and publications in 
supply are subjects for special correspondence, Such correspondence should ident 
writer and explain the use to be made of the publication or publications, 


Address orders and correspondence to the Chief 
Illinois Natural History Survey 
Natural Resources Building, Urbana, Illinois 61801 


ILLINOIS 


History Survey 
BULLETIN 


The Bantam Sunfish, Lepomis 
symmetricus: Systematics and 
Distribution, and Life History 
in Wolf Lake, Illinois 


oks M. Burr 


; 


A 


E OF ILLINOIS 
ARTMENT OF REGISTRATION AND EDUCATION 
4 


§ 

TURAL HISTORY SURVEY DIVISION “ 

BANA, ILLINOIS | 

} NATORAL BRSTORY SURVEY 

| VOLUME 31, ARTICLE 10 
ae Pe ects 977 SEPTEMBER, 1977 


The Bantam Sunfish, Lepomis 
symmetricus: Systematics and 
Distribution, and Life History 
in Wolf Lake, Illinois 


oks M. Burr 


E OF ILLINOIS 
RTMENT OF REGISTRATION AND EDUCATION 


URAL HISTORY SURVEY DIVISION 
ANA, ILLINOIS 


VOLUME 31, ARTICLE 10 
SEPTEMBER, 1977 


STATE OF ILLINOIS 


DEPARTMENT OF REGISTRATION AND EDUCATION 


BOARD OF NATURAL RESOURCES AND CONSERVATION 


Joan G. ANDERSON, B.S. Chairman; THOMAS PARK, Ph.D., Biology; 


L. L. Suoss, Ph.D., Geology; 


H. S. Gurowsky, Ph.D., Chemistry; Ropert H. ANDERSON, B.S.C.E., Engineering; STANLEY K, SHAPIRO, 


Ph.D., Forestry; W. L. Everitt, E.E., 


Ph.D., Representing the President of the University of Illinois; 


JOHN C. Guyon, Ph.D., Representing the President of Southern Illinois University. 


NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois 
SCIENTIFIC AND TECHNICAL STAFF 
GEORGE SPRUGEL, JR., Ph.D., Chief 
ALICE K, ADAMS, Secretary to the Chief 


Section of Economic Entomology 


WitiiaM H. LucKMANN, Ph.D., Entomologist and 
Head 

JAMES E, APPLEBY, Ph.D., Entomologist 

Marcos KoGan, Ph.D., Entomologist 

Ronautp H. MEYER, Ph.D., Entomologist 

STEVENSON Moore, III, Ph.D., Entomologist, Ex- 
tension 

Epwarp J. ARMBRUST, Ph.D., Associate Entomolo- 
gist 

JOSEPH V. MaAppDox, Ph.D., Associate Entomologist 

RoBertT D. PauscH, Ph.D., Associate Entomologist 

RALPH E. SECHRIEST, Ph.D., Associate Entomolo- 
ist 

Ses K. BousEMAN, M.S., Assistant Entomologist 

CHARLES D. BREMER, M.S., Assistant Entomologist, 
Extension 

MricHaeL E. Irwin, Ph.D., Assistant Entomologist 

Donautp E. KUHLMAN, Ph.D., Associate Professor, 
Extension 

RoscorE RANDELL, Ph.D., Associate Professor, Ex- 
tension 

WILLIAM G. RUESINK, Ph.D., Assistant Entomolo- 
gist 

DovueGuas K. SELL, Ph.D., Assistant Entomologist 

JOHN L. WEDBERG, Ph.D., Assistant Entomologist, 
Extension 

CLARENCE E. WHITE, B.S., Assistant Entomologist 

Kevin D, BuAck, M.S., Assistant Specialist, Exten- 
ston 

Davin A. GENTRY, M.S., Assistant Specialist, Ex- 
tension 

STEVEN TROESTER, M.E., Assistant Systems Engineer 

JEAN G. WILSON, B.A., Supervisory Assistant 

CATHERINE EASTMAN, Ph.D., Assistant Professional 
Scientist 

Joun T. SHAw, B.S., Assistant Professional Scientist 

DANIEL SHERROD, M.S., Assistant Professional Scientist 

Lester WEI, Ph.D., Assistant Professional Scientist 

CHARLES G. HELM, M.S., Junior Professional Sci- 
entist 

LINDA ISENHOWER, Junior Professional Scientist 

STEPHEN ROBERTS, B.S., Junior Professional Sci- 
entist 

Li-CHUN CHIO, Ph.D., Research Assistant 

ELIZABETH ALLISON, B.S., Research Assistant 

MARGARET ANDERSON, B.S., Research Assistant 

ROBERT J. BARNEY, B.S., Research Assistant 

Tzu-Suan CuHu, M.S., Research Assistant 

MARION FARRIS, M.S., Research Assistant 

JANET Harry, B.S., Research Assistant 

BONNIE IRWIN, M.S., Research Assistant 

Louis JAcKAal, M.S., Research Assistant 

JENNY KoGAn, M.S., Research Assistant 

PATRICIA MACKEY, B.S., Research Assistant 

BRIAN MELLIN, B.S., Research Assistant 

Mary MILsraTH, M.S., Research Assistant 

JuDY MOLLETT, B.S., Research Assistant 

LYNN PAUTLER, B.S., Research Assistant 

CELIA SHIH, M.S., Research Assistant 

BARBARA STANGER, B.S., Research Assistant 

LEE ANNE TURNER, M.S., Research Assistant 

JO ANN AUBLE, Technical Assistant 

CHARLOTTE JOHNSON, B.S., Technical Assistant 


Section of Botany and Plant Pathology 
NS RH Ph.D., Plant Physiologist and 
ea 

EUGENE B. HIMELICK, Ph.D., Plant Pathologist 

DAN NEELY, Ph.D., Plant Pathologist 

D. F. SCHOENEWEISS, Ph.D., Plant Pathologist 

J. LELAND CRANE, Ph.D., Associate Mycologist 

KENNETH R. ROBERTSON, Ph.D., Assistant Taxono- 
mist 

Betty S. NELSON, Junior Professional Scientist 

GENE FE, RED, Junior Professional Scientist 

JAMES E. SERGENT, Greenhouse Superintendent 

RICHARD WILSON, Technical Assistant 


Section of Aquatic Biology 


D. HOMER Buck, Ph.D., Aquatic Biologist 

WILLIAM F. CHILDERS, Ph.D., Aquatic Biologist 

R. WELDON LARIMORE, Ph.D., Aquatic Biologist 

ROBERT C, HILTIBRAN, Ph.D., Biochemist 

ALLISON BRIGHAM, Ph.D., Associate Aquatic Biol- 
ogist 

WARREN U. BRIGHAM, Ph.D., Associate Aquatic Bi- 
ologist 

RICHARD E. SPARKS, PhD., Associate Aquatic Biologist 

Tep W. SToRCK, Ph.D., Assistant Aquatic Biologist 

JOHN TRANQUILLI, Ph.D., Assistant Aquatic Biolo- 
gist 

RicHaArD J. BAuR, M.S., Junior Professional Scientist 

CarL M. THOMPSON, B.S., Junior Professional Sci- 
entist 

JANA LEE WaAITE, M.S., Junior Professional Scien- 
tist 

DONALD W. DUFFORD, M.S., Research Associate 

JOHN M. MCNuRNEY, M.S., Research Associate 

Davin P. Puitipp, Ph.D., Research Associate 

Harry W. BERGMANN, B.S., Research Assistant 

WARNER D. BRIGHAM, B.S., Research Assistant 

Kurt T. CLEMENT, B.S., Research Assistant 

LarrRY W. CouTant, M.S., Research Assistant 

DONALD R. HALFFIELD, M.S., Research Assistant 

EARL THOMAS Joy, JR., M.S., Research Assistant 

ROBERT MoRAN, M.S., Research Assistant 

MICHAEL J. SULE, M.S., Research Assistant 

STEPHEN O. SWADENER, M.S., Research Assistant 

STEPHEN W. WAITE, M.S., Research Assistant 

Cart ALDE, B.S., Technical Assistant 

Paut Beaty, M.S., Technical Assistant 

KATHRYN EwINe, B.S., Technical Assistant 

Jerr Hutton, B.S., Technical Assistant 

GeorcE Lewis, M.S., Technical Assistant 

SuE Moran, Technical Assistant 

Don Myrick, B.S., Technical Assistant 

JENS SANDBERGER, M.S., Technical Assistant 

JOHN J. SuLowAY, B.S., Technical Assistant 

LIANN SuLoway, M.S., Technical Assistant 

RosBert THomas, B.S., Technical Assistant 

GARY L. WARREN, B.S., Technical Assistant 

Lowe Lt Davis, Field Assistant 

C. RussELu Rose, Field Assistant 


Section of Faunistic Surveys and 
Insect Identification 


PuHitip W. SMITH, Ph.D., Taxonomist and Head 
WALLACE E. LABERGE, Ph.D., Taxonomist 

GEORGE L. GopFREY, Ph.D., Associate Taxonomist 
JOHN D. UNZICKER, Ph.D., Associate Taxonomist 
DONALD W. WEBB, M.S., Associate Taxonomist 
Larry M. Pace, Ph.D., Associate Taxonomist 
BERNICE P. SWEENEY, Junior Professional Scientist 


Section of Wildlife Research 
ony 3 SANDERSON, Ph.D., Wildlife Specialist and 
ea 

FRANK C. BELLROSE, Sc.D., Wildlife Specialist 

WILLIAM R, Epwarps, Ph.D., Wildlife Specialist 

JEAN W. GRABER, Ph.D., Wildlife Specialist 

RICHARD R. GRABER, Ph.D., Wildlife Specialist 

HAROLD C. HANSON, Ph.D., Wildlife Specialist 

W. W. CocHRAN,~JR., B.S., Associate Wildlife 
Specialist 

CHARLES M. NIXON, M.S., Associate Wildlife 
Specialist 

KENNETH E, SMITH, Ph.D., Associate Chemist 

RONALD L. WESTEMEIER, M.S., Associate Wildlife 
Specialist 

LONNIE P. HANSEN, Ph.D., Assistant Wildlife Specialist 

STEPHEN P. HAvERA, M.S., Assistant Wildlife 
Specialist 

DAvipD R. VANCE, M.S., Assistant Wildlife Specialist 

RICHARD E, WARNER, M.S., Assistant Wildlife Specialist 

RONALD E. DuzAn, Junior Professional Scientist 

esp C. ScHuutTz, M.A., Junior Professional 

cientist 


Continued on page 466 


CONTENTS 


ACKNOWLEDGMENTS 
METHODS AND MATERIALS 


SYSTEMATICS 
Synonymy 
Types . 5 
ID¥apnosis. -......00-.c::- 
Description 
Variation 
Sexual 
Allometric . 
Geographic 
Relationships 
Specimens Studied . 
Ohio River Drainage 
Mississippi River Drainage 
Gulf Coast Drainage 


DISTRIBUTION 
CONSERVATION STATUS 


Lire History iN WotF LAKE 
Study Area 
Habitat 
Reproduction 
Reproductive Cycle of the Male 


Reproductive Cycle of the Female 


Spawning 
Development and Growth 
Demography 

Density 

Composition 

Survival 
Diet ; 
Interaction with Other Organisms 

Competition 

Predation 

Hybridization 

Parasitism 
Summary 


LITERATURE CITED 


INDEX 


437 
438 
439 


_. 439 


440 


a 


443 


. 445 


| 445 


445 
445 
446 
446 
446 
446 


. 447 


.. 447 


448 


449 
449 
449 


+ ey) 


450 
451 
452 
453 
454 


aba 


455 
455 
455 
457 
457 
458 
459 
459 
460 


461 
465 


This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It is 
a contribution from the Section of Faunistic Surveys and Insect Identification of the Illinois 


Natural History Survey. 


Brooks M. Burr is a former Research Assistant, Illinois Natural History Survey. He is 
presently an assistant professor, Department of Zoology, Southern Illinois University, Carbondale. 


GB 2 
(01284—2M-8-77) 


Fig. 1—Distribution of Lepomis symmetricus in relation to the Coastal Boundary (solid black line). 
circles represent recent localities (1938 to the present); large open circles represent old records (pre-19 
where the species is presumably extinct. The most northern open circle also represents the type-loca 
The life-history study area is enclosed within the square. 


The Bantam Sunfish, Lepomis symmetricus: Systematics 
and Distribution, and Life History in Wolf Lake, Illinois 


The bantam sunfish, described as 
Lepomis symmetricus by Stephen A. 
Forbes in 1883, is one of the least 
known species in the genus, probably 
because of its small size, rarity over 
parts of its range, occurrence in rather 
inaccessible swamp habitats, and drab 
and nondescript appearance. This ef- 
fort to remedy the gaps in our knowl- 
edge of the species reviews all published 
references to L. symmetricus. To sup- 
plement the meager information avail- 
able, this report includes an analysis 
of morphological variation based on 
the study of museum specimens, an 
assessment of the species’ distribution, 
and a life-history study based on peri- 
odic collections made at a study site in 
southern Illinois. 


ACKNOWLEDGMENTS 

For aid in the literature search, I am 
indebted to Philip W. Smith and IIli- 
nois Natural History Survey librarian 
Doris L. Sublette; for help in collecting 
specimens, to present and former asso- 
ciates John A. Boyd, Lloyd R. Davis, 
Larry M. Page, Philip W. Smith, and 
Roger D. Wrisberg. Larry M. Page 
and Philip W. Smith provided counsel 
on numerous matters. For identifying 
trematode and acanthocephalan _par- 
asites, I am grateful to David F. 
Oetinger and for advice on other par- 
asitological problems, to Mary H. 
Pritchard, both of the Nebraska State 
Museum. E. Donald McKay III of the 
Illinois State Geological Survey sup- 
plied information on the age of Wolf 
Lake. 

For information about the location 
of syntypes and other pertinent study 
material, I am indebted to museum 
officials Reeve M. Bailey; James E. 


Brooks M. Burr 


Bohlke; Garrett S. Glodek; Tomio 
Iwamoto, California Academy of Sci- 
ences, including the Stanford Univer- 
sity (SU) collection; Robert K. John- 
son; Craig E. Nelson; Donn E. Rosen, 
Robert Schoknecht, Museum of Com- 
parative Zoology (MCZ) at Harvard 
University; and Keith §. Thompson. 
For providing laboratory space and/or 
the loan of specimens I thank Reeve 
M. Bailey, University of Michigan Mu- 
seum of Zoology (UMMZ); Thomas 
M. Buchanan, Westark Community 
College at Fort Smith, Arkansas 
(ARP); Neil H. Douglas, Northeast 
Louisiana University (NLU); David 
A. Etnier, University of Tennessee 
(UT); Robert K. Johnson, Field Mu- 
seum of Natural History (FMNH); 
Ernest A. Lachner, U.S. National Mu- 
seum of Natural History (USNM) ; 
Robert F. Martin, Texas Natural His- 
tory Collection at Austin (TNHC) ; 
John D. McEachran, Texas Cooper- 
ative Wildlife Collection at College 
Station (TCWC) ; William D. Pearson, 
University of Louisville, Kentucky 
(UL); Henry W. Robison, Southern 
State College at Magnolia, Arkansas 
(HWR); and Royal D. Suttkus and 
Michael M. Stevenson, Tulane Univer- 
sity, Louisiana (TU). For providing 
lists of locality records and/or distri- 
bution maps I thank Thomas M. Bu- 
chanan; B. E. Gandy, Mississippi Mu- 
seum of Natural Science; Henry W. 
Robison; Morgan E. Sisk, Murray 
State University, Kentucky; Neil H. 
Douglas; and Glenn H. Clemmer, Mis- 
sissippi State University. 

Unless stated otherwise, the majority 
of the specimens used in this study are 
deposited at the Illinois Natural His- 
tory Survey (INHS). 


437 


438 


Most of the illustrations for this 
paper were prepared by Larry Farlow, 
Technical Photographer; Lloyd Le- 
Mere, Technical Illustrator; and Craig 
Ronto, all of the Illinois Natural His- 
tory Survey; the drawing of the sub- 
adult was done by Alice A. Prickett of 
the University of Illinois School of Life 
Sciences. Computer analysis of some of 
the data was undertaken by Stephen D. 
Cowan of the Survey. The manuscript 
was edited by Shirley McClellan, As- 
sistant Technical Editor at the Survey, 
and Dr. Neil H. Douglas, Northeast 
Louisiana University, served as guest 
reviewer. Partial support for the field 
work was provided by the U.S. Depart- 
ment of Agriculture Forest Service; the 
Illinois Natural History Survey ren- 
dered the other support. Special per- 
mission to collect specimens of the 
bantam sunfish, which is protected by 
the Illinois Fish Code, was given by 
the Department of Conservation. Per- 
mits to take specimens in the National 
Park were issued by Joe L. Newcomb 
of the Forest Service. Paul Brown of 
the Trojan Powder Plant granted per- 
mission to collect on powder plant 


property. 


METHODS AND MATERIALS 


An attempt was made to compile as 
complete a synonymy as possible for 
Lepomis symmetricus, and it is be- 
lieved that virtually all published ref- 
erences to it have been examined. 
Morphological data were taken on se- 
lected series that could be expected to 
show geographical variation, allomet- 
ric variation, or sexual dimorphism in 
the species. Meristic and morphomet- 
ric data were taken in the conventional 
manner of Hubbs & Lagler (1964: 19- 
26). One-way analysis of variance tests 
were run to determine significant dif- 
ferences in means of samples deter- 
mined by sex. Unless stated otherwise, 
measurements are standard lengths 
(SL). 


ILLINOIS NATURAL HISTORY SURVEY BULLETIN 


Vol. 31, Art. 10 


Observations and minnow-seine col- 
lections were made in Wolf Lake in 
Union County, Illinois, at approx- 
imately I-month intervals, except dur- 
ing the spawning season, when more 
frequent observations were needed. 
The life-history study began 2 June 
1973 and ended 27 May 1975. 

Specimens were preserved in 10-per- 
cent formalin and were returned to 
the laboratory for study. In all, 233 
specimens from Wolf Lake were pre- 
served and examined. Because the spe- 
cies is protected by the Illinois Fish 
Code, usually no more than 20 spec- 
imens were taken on one visit even 
when the species was commonly en- 
countered, so as not to seriously dec 
imate the population. Collecting was 
done by bag seine; minnow seine; dip 
net; and, in one instance, by electro- 
fishing. Potential predators of the ban- 
tam sunfish were occasionally collected 
for examination of stomach contents. 
Field notes were routinely taken. In 
the laboratory, specimens were sexed, 
measured, and aged, and their gonads 
and stomachs were excised and studied. 
During the spawning season, breeding 
adults were brought to the laboratory 
and placed in observation tanks. 

Aging to year class was done by 


‘counting scale annuli removed from 


the dorsum. Aging to month was done 
by using May, the month of greatest 
breeding activity in Wolf Lake, as 
month zero. Thus, a sunfish collected 
in October with one scale annulus was 
estimated to have lived 1 year and 5 
months. For certain comparisons sun- 
fishes were divided into young (through 
12 months) and adult (over 12 months) 
age groups. 

Weights of the ovaries of 30 females 
were obtained and recorded as a pro- 
portion of the adjusted body weight 
(the specimen minus the ovaries, stom- 
ach, intestine, and liver) of the female. 
Mature ova from 14 preserved breed- 
ing females were counted. Indicators 
used for ascertaining probable spawn- 


Sept., 1977 Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 439 


ing periods in other localities and 
other years were that males exhibited 
breeding color patterns and that fe- 
males were heavy with ova. 

The relative survival of ‘each year 
class of the study population was cal- 
culated by expressing the number of 
individuals in that year class as a pro- 
portion of the number of individuals 
in a younger year Class. 


SYSTEMATICS 
SYNONYMY 


Lepomis symmetricus Forbes 


Lepomis symmetricus McKay 1882: 88 
(nomen nudum); Forbes in Jordan 
& Gilbert 1883: 473-474 (original de- 
scription, Illinois River [at Pekin] 
Illinois) ; Forbes 1884: 68 (Illinois 
range) ; Jordan 1884: 320-321 (rede- 
scription, museum specimens cited) ; 
Jordan 1888:117  (redescription) ; 
Bollman 1892: 566, 571 (key, range) ; 
Evermann & Kendall 1894: 84, 93, 
111 (redescription, Texas records) ; 
Hay 1894: 255, 261 (redescription, 
key, not taken in Indiana) ; Richard- 
son 1904: 31, 33 (relationships, key, 
Illinois range); Forbes & Richard- 
son 1908: 251-252 _ (redescription, 
key, Illinois range); Forbes 1909: 
388 (Illinois range) ; Pratt 1923: 118 
(key, range) ; Greene 1927: 309 (not 
in Wisconsin); Hildebrand & 
Towers 1927: 133-134 (Greenwood, 
Mississippi, food habits in Missis- 
sippi); Summers 1937: 434 (new 
trematode parasite, Baton Rouge, 
Louisiana) ; Mizelle 1938: 160 (tre- 
matode transferred to new genus) ; 
Mizelle & Hughes 1938: 351  (tre- 
matode parasite cited); Summers & 
Bennett 1938: 248 (trematode para- 
site cited); Kuhne 1939: 110 (Ten- 
nessee list, redescription, sexes fig- 
ured); Lamb 1941:45 (Willow 
Creek, San Jacinto drainage, Texas) ; 
Fowler 1945: 364, 370 (Louisiana 
and Mississippi records, figure erro- 
neous) ; Gerking 1945: 115 (possible 


in Indiana) ; Seamster 1948: 165, 168 
(trematode parasite cited); Baugh- 
man 1950: 247 (Texas list) ; Moore 
& Cross 1950: 146 (recorded from 
Oklahoma) ; Reeves & Moore 1951: 
42. (Oklahoma Coastal Plain) ; 
Bohlke — 1953: 71 (SU — syntypes 
listed) ; Moore 1952: n.p. (Oklahoma 
list); Jurgens & Hubbs 1953: 15 
(Texas list); Knapp 1953: 115 (key, 
Texas range); Gunning & Lewis 
1955: 556 (habitat, food habits in 
Illinois) ; Gunning & Lewis 1956: 24 
(Wolf Lake and Pine Hills, Illinois) ; 
Eddy 1957: 191 (key, range) ; Hubbs 
1957a: 97 (Texas range); Hubbs 
1957b:9 (Texas list); Moore in 
Blair e¢ al. 1957: 170 (key, range) ; 
Hubbs 1958: 10 (Texas list); Bou- 
dreaux et al. 1959: 8, 10 (Sour Lake, 
Hardin County, Texas) ; Cook 1959: 
180 (redescription, ecology, Missis- 
sippi range); Bailey et al. 1960: 27 
(list); Smith & Bridges 1960: 254 
(INHS syntypes); Hubbs 1961: 10 
(Texas range); Branson & Moore 
L9G 23 OF WD, 2a 229s Ol oo, Ales, 
65, 72, 91, 99 (relationships, acous- 
tico-lateralis system) ; Clay 1962: 119 
(Kentucky range); Collette 1962: 
146, 177 (associate of slough darter 
and swamp darter); Lambou 1962: 
78 (Lake Bistineau, Louisiana) ; 
Walker 1962: 40 (Jackson, Lincoln, 
and Bienville parishes, Louisiana) ; 
Walker 1963: 48 (Choudrant Bayou, 
Louisiana) ; Sharma 1964: 533 (mu- 
cus cells in canal linings) ; Burton & 
Douglas 1965: 94 (Bayou De Siard, 
Louisiana) ; Smith 1965: 9 (Illinois 
range); Pflieger 1966: 53 (Missouri 
key); Breder & Rosen 1966: 413 
(breeding habits unknown); Chil- 
ders 1967: 160 (tribe Lepomini) ; 
Douglas & Davis 1967: 23 (Louisi- 
ana list); Hoffman 1967: 340 
(known parasites) ; Pflieger 1968: 54 
(Missouri key); Moore in Blair 
et al. 1968: 128-129 (key, range) ; 
Whitaker 1968: 96-97 (key, range) ; 
Eddy 1969: 217 (key, range); 


440 


Smith & Sisk 1969: 66 (Obion 
Creek, Kentucky); Bailey et al. 
1970: 36 (list); Jenkins et al. 
1971: 74 (possibly present in lower 
Tennessee or Cumberland rivers) ; 
Pflieger 1971: 413-414 (habitat, zoo- 
geography, Missouri range); Smith 
et al. 1971: 10 (not in upper Missis- 
sippi River) ; Hubbs 1972: 6 (Texas 
range) ; Miller 1972: 244 (threatened 
in Illinois and Missouri) ; Rozenburg 
et al. 1972: ili, 22, 28, 30, 32, 33, 36, 
40, 45, 51, 82, 111 (Navasota River, 
Texas); Buchanan 1973a: 29 (Ar- 
kansas list); Buchanan 19730: 51 
(key, Arkansas range); Miller & 
Robison 1973: 184-185 (key, rede- 
scription, ecology, Oklahoma range) ; 
Moore 1973: 6 (McCurtain County, 
Oklahoma) ; Smith 1973: 33 (Illinois 
key) ; Lopinot & Smith 1973: 46-47 
(status in Illinois) ; Buchanan 1974: 
89 (status undetermined in Arkan- 
sas); Douglas 1974: 312-313 (rede- 
scription, Louisiana range) ; Pflieger 
in Holt et al. 1974: n.p. (rare in 
Missouri) ; Ackerman 1975: 10 (en- 
dangered in Illinois); Boyd et al. 
1975: 11, 21 (status in Illinois) ; Clay 


ILLINo1Is NATURAL History SURVEY BULLETIN Vol. 31, Art. 10 


spawning); Webb & Sisk 1975: 63, 
67, 69 (Bayou de Chien, Kentucky, 
endangered in Kentucky); Hubbs 
1976: 6 (Texas list); Hubbs & Pigg 
1976: 116 (indeterminate status in 
Oklahoma); Seehorn 1976: 21 (South- 
eastern National Forest list) . 

Apomotis symmetricus: Boulenger 
1895: 21 (redescription); Jordan & 
Evermann 1896: 998-999 (redescrip- 
tion); Evermann 1899: 310 (Lake 
Lapourde, Louisiana); Large 1903: 
24 (Illinois range); Jordan e¢ al. 
1930: 299 (list, range); Gowanloch 
1933: 348, 351 (Louisiana range) ; 
Schlaikjer 1937:12 (phylogeny) ; 
Schrenkeisen 1938: 243-244  (rede- 
scription, range) . 

Lethogrammus symmetricus: Hubbs in 
Jordan 1929: 147 (transfer to new 
genus erected by C. L. Hubbs); 
Greene 1935: 220 (not in Wiscon- 
sin) ; O'Donnell 1935: 486 (Illinois 
range); Breder 1936: 28 (breeding 
habits unknown); Baker 1937: 48 
(redescription, rare at Reelfoot 
Lake); Baker & Parker 1938: 162 
(Reelfoot Lake list); Baker 1939a: 
34 (redescription, sexes figured, com- 


1975: 267, 276, 280 (redescription, mon at Reelfoot Lake); Baker 
key, Kentucky range); Douglas & 1939b: 45 (Reelfoot Lake key). 
Davis 1975: 23 (Louisiana list) ; Mc- 

Reynolds 1975: 253 (LaRue Swamp, TYPES 

Illinois) ; Pflieger 1975: 254, 265 (fig- Lepomis symmetricus was described 
ure, key, redescription, Missouri by Forbes in Jordan & Gilbert (1883: 
range) ; Robison 1975: 54, 56 (Saline 473-474) from a syntypic series consist- 
River, Arkansas, evidence of recent ing of 15 specimens collected 16 April 


Table 1.—Frequency distribution for number of caudal peduncle scales in selected pop- 
ulations of Lepomis symmetricus. 


‘ Number of Scales Standard Coefficient 
Drainage N Mean Deva of 

17 18 19 20 21 22 eviation Variation 
Illinois R., Ill. Le laren? peda.) ee 11 19.5 0.69 3.5 
Wabash R., II. cS: ESC RES 12 20.9 0.67 3.2 
Mississippi R.., Ill., Mo., Ky. Meese are liam wet) 51 20.2 1.21 6.0 
Mississippi R., Tenn. re, Catal Oe ply White Sam 19.5 0.51 2.6 
Mississippi R., Ark., La. WG NO By 25 19.3 0.95 4.9 
Ouachita R., Ark., La. 4° beara 252 26 18.8 1.13 6.0 
Red R., Okla., Tex., Ark., La. Yaar S. It) NL 33 18.5 1.03 5.6 
Gulf Slope, Tex., La. Te 7) 10h £9 Gar 39 19.0 1.34 7.1 


Sept., 1977 Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 441 


Table 2.—Frequency distribution for number of lateral line scales in selected populations 
of Lepomis symmetricus. 


Number of Scales Coefficient 
Drainage N Mean aoe of 
30 31 32 33 34 35 36 37 38 Sree B OD  ariation 
Illinois R., Il. OA ti aN ee ah MEN gece 11 32.5 0.82 2.5 
Wabash R., III. Ae ine bei ete eed Meee ee S| | ee 35.4 1.51 43 
Mississippi R., 
Ill., Mo., Ky. rer A IPA Ge ei (iis ZO elt 51 33.6 2.87 8.5 
Mississippi R., 
Tenn. ee eG gga AP eos el ee 25 33.5 1.39 4.2 
Mississippi R., 
Ark., La. [Vl IP POLY (Pr 8%, Sil” TO, ia eee oe 25 33.4 1.58 4.7 
Ouachita R., 
Ark., La. he NLS ilar MR La la ed Bile 26 33.3 1.95 5.9 
Red R., 
Okla., Tex., 
Ark., La. Gat LO ge Ora Cee oo 58 33 32.0 1.39 4.3 
Gulf Slope, 


Tex., La. eee RO Bi See Soe Mess sels 39 32.6 1.57 4.8 


Table 3.—Frequency distribution for number of dorsal soft rays in selected populations 
of Lepomis symmetricus. 


Number of Rays Stdard Coefficient 
Drainage arr = ora N Mean Deviation oe 

Die tO Ail S12 Variation 
Illinois R., Ill. os 7 a) al 1] 10.5 0.69 6.6 
Wabash R., Il. 2 9 1 12 9.9 0.51 52 
Mississippi R.., Ill., Mo., Ky. Pa ABS APA 4 51 10.6 0.40 3.8 
Mississippi R., Tenn. So ey Ga 2 25 10.4 0.65 6.3 
Mississippi R., Ark., La. 3) 18 4 Z 25 10.0 0.54 5.4 
Ouachita R., Ark., La. 3) lg 3 1 26 10.1 0.63 6.2 
Red R., Okla., Tex., Ark., La. 7 a) Vous By ee 33 10.2 0.66 6.5 
Gulf Slope, Tex., La. aie Spee UI) i: 39 10.3 0.77 75 


Table 4.—Frequency distribution for number of anal soft rays in selected populations of 
Lepomis symmetricus. 


Number of Rays : - Standand ee 
i Oe ean See ° 

me Pie (ee eee Deviation Variation 
Illinois R.., Ill. an pelle Scar TH 11 10.2 0.60 5.9 
Wabash R.), II. 1 Gi) BE Aa 12 10.1 0.51 5.0 
Mississippi R.., Ill., Mo., Ky. MY) 22) ae 1 51 10.2 0.61 6.0 
Mississippi R., Tenn. aT, 1 25 9.8 0.52 5.3 
Mississippi R., Ark., La. Ses: 2 25 9.8 0.60 6.1 
Ouachita R., Ark., La. 8 17 1 26 9.7 0.53 5.5 
Red R., Okla., Tex., Ark., La. ee | 9 33 10.2 0.58 5.7 
Gulf Slope, Tex., La. ee 3 39 9.8 0.54 55 


442 


and 2 June 1880 from the Illinois 
River (Mississippi drainage) at Pekin, 
Tazewell County, Illinois (Fig. 1). All 
15 of the original syntypes are extant: 
INHS 220 (8, 32.7-39.5 mm _ SL); 
INHS 226 (2, 50.1-51.2 mm SL) ; MCZ 
25014 (1, 49.5 mm SL); SU 1276 (3, 
49.8-56.9 mm SL); USNM 29864 (1, 
51.0 mm SL). All 15 are in a good state 
of preservation. To preserve customary 
nomenclature and in accordance with 
the International Code of Zoological 
Nomenclature Article 74, recommenda- 
tion 74D, a lectotype of L. symmetricus 
Forbes is herewith designated (INHS 
75004, 39.5 mm SL). The specimen, 
a juvenile, conforms to the character- 
ization of the species given under De- 
scription and in Tables 1-4. The in- 
complete lateral line has 34 scales with 
6 scales above and 13 scales below the 
lateral line. There are 19 caudal pe- 
duncle scales, 5 cheek row scales, and 
6 branchiostegal rays. Fin ray counts 
are: dorsal spines, 10; anal spines, 3; 
pectoral rays, 12-12; dorsal soft rays, 
10; anal soft rays, 10. The nine other 
specimens originally accessioned as 


ILLINoIs NATURAL HISTORY SURVEY BULLETIN 


Vol. 31, Art. 10 


INHS 220 and 226 are paralectotypes, 
now INHS 75005 and INHS 75006, re- 
spectively. The USNM, SU, and MCZ 
syntypes also became paralectotypes, 
keeping their original catalogue num- 
bers. 

It is unlikely that the original mate- 
rial of L. symmetricus collected by 
Forbes and associates was captured 
from the Illinois River proper. Al- 
though the Illinois River has changed 
rather drastically since Forbes’s era, it 
probably never maintained habitat 
suitable for L. symmetricus. More 
likely the specimens came from one of 
the natural floodplain lakes in the 
Pekin area, where favorable habitat 
has been present in past years. 


DIAGNOSIS 


The most diminutive species of Le- 
pomis (the largest specimen measured 
is 75.5 mm SL) is distinguished from 
other members of the genus by this 
combination of characters: Lateral 
line incomplete (1-18 scales unpored) 
or interrupted (as many as 6 times). 
Gill rakers long (longest in the genus, 


Table 5.—Proportional measurements of Lepomis symmetricus from throughout the range, 


expressed in thousandths of standard length." 


10 Males (54-64 mm SL) 


10 Females (50-64 mm SL) 


Coeffi Coeffi 
Stan : Stan- —. 

Measurement sea CSR Beer COM 
Range Mean ‘ of Range Mean : of 

Devi aaa Devi- 5 

= ari- 5 Vari- 

ation ation ation ation 
Head length 375-423 396 =: 013 3.5 361-403 390 _ 0125 3s 
Body depth 471-531 491 018 3.6 468-527 494 9017 3.4 
Caudal-peduncle depth 150-169 160 006 3.6 142-192 163 014 8.8 
Pectoral fin length 245-285 263 014 5.4 248-291 263 «6012. 4.7 
Pelvic fin length 227-255 238 8=©009 Ss 33.9 212-243 225 (O10. 4.6 
Longest dorsal spine 116-153 139 010 Tees 126-164 140 014 9.7 
Head width 180-219 203 = 013 6.5 191-234 213 013 6.0 
Bony interorbital width 078-096 087 006 866.6 074-093 084 = 8=—007 7.8 
Snout length 071-086 078 005 6.7 074-088 081 006 «7.4 
Upper jaw length 123-151 140 009 6.2 124-158 133 010 7.8 
Predorsal length 439-480 459 014 = 33.0 445-483 462 014 29 
Base dorsal fin length 461-508 478 015 3.1 455-517 480 017 3.6 
Longest anal spine 120-151 137 010 3=6.9 124-157 139° (012) 8.7 
Base anal fin length 216-297 244 6022, 8.8 206-257 234 «015 6.6 
Orbit length 087-105 095 005 5.7 083-105 095 0027.8 


2 Based on NLU 29918, 12804, 1954; UT 90.116, 


75022, 75023, 18151, 18143, 17547. 


90.140; TCWC 3643; HWR 74-8; INHS 75020, 75021, 


Sept., 1977 


longest rakers 2.3-2.9 mm), and slen- 
der (0.3-0.5 mm wide, 7—9 times longer 
than wide) , numbering 12-15, modally 
13. Opercle stiff to its bony margin, 
the dark opercular spot slightly diffuse 
on narrow, bordering membrane. Dor- 
sal coloration dusky with dark coffee- 
colored spots on body, spots occasion- 
ally forming irregular vertical bands. 
Head and cheeks darkened and with- 
out patterns. Juveniles often more ver- 
tically barred than adults and have a 
prominent black blotch in the poste- 
rior rays of the soft dorsal fin, becom- 
ing less intense with age. Branson & 
Moore (1962) showed these additional 
characters to be distinctive: only one 
posterior pore on the post-temporal, 
lateralis ending under the soft dorsal 
fin, preopercle angle 110° to 115°, 
lachrymal bone nearly twice as tall as 
wide, supramaxilla shorter than max- 
illa, and no teeth on tongue or ptery- 
goids. 


DESCRIPTION 
Forbes (im Jordan & Gilbert 1883: 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


443 


473-474) and Forbes & Richardson 
(1908: 251-252) adequately described 
the specimens available to them. The 
following description is an amplifica- 
tion, which includes additional meris- 
tic and morphometric data, and a more 
comprehensive description of colora- 
tion. Body proportion yalues are 
presented in Table 5. When no geo- 
graphic variation was noted, the var- 
iation data from throughout the range 
of the species are merely summarized. 
When geographic variation was noted, 
the ranges and modes are given in 
the description, but their frequencies 
are discussed under Variation. Counts 
of lateral-line scales, caudal-peduncle 
scales, dorsal soft rays, and anal soft 
rays, all of which show slight clinal 
variation, are presented in Tables 1-4. 
General physiognomy and pigmenta- 
tion of adults and juveniles are shown 
in Fig. 2 and 3. 

Lateral line scales 30-38, modally 32 
(Table 2). Bailey (1938) reported one 
specimen with 40 lateral line scales. 
Scales above the lateral line 5 (in 7 


Fig. 2—Breeding male Lepomis symmetricus 53.6 mm in standard length collected in 
Wolf Lake on 27 May 1975. Pigmentation in the fins is somewhat subdued by preservation. 


444 


specimens), 6 (92), 7 (11), x=6.1. 
Scales below the lateral line 12 (in 48 
specimens) , 13 (52), 14 (15), x = 12.7. 
Caudal peduncle scales 17-22, modally 
19 (Table 1). Scales on cheek 4-6, 
modally 5. Scales well developed on 
preopercle, subopercle, interopercle, 
and opercle, all such scales about the 
same size and shape. No scales on top 
of head. 

Dorsal spines 9 (in 22 specimens) , 
10 (133), 11 (6), x=9.9. Dorsal soft 
rays 9-12, modally 10 (Table 3). Anal 
spines 2 (in 1 specimen), 4 (2), 3 in 
all others. Anal soft rays 9-12, modally 
10 (Table 4). Pectoral rays 11 (in 8 
specimens) , 12 (66), 13 (32), x= 12.2. 
All pelvic fins counted had 1 spine. 
Pelvic rays 4-4 (in 1 specimen), 4—5 
(2), 5-5 (42). Principal caudal rays 
17 (in 41 specimens), 18 (1). 

Gill rakers on first arch (all rudi- 
ments counted) 12 (in 12 specimens) , 
13 (36), 14 (22), 15 (6), x=13.3. 
Rakers long and slender (see Diagno- 
sis). Rudimentary rakers (usually 3-5) 
are shorter and more blunt. The lat- 
eral line on the body is incomplete or 
interrupted (see Diagnosis and Fig. 3). 
The cephalic lateral-line system was 
described in detail by Branson & 
Moore (1962). Caudal fin slightly 
emarginate. No teeth on tongue and 
pterygoids. Teeth present on vomers 
and palatines. Pharyngeal arches nar- 
row with many small, blunt subconical 
teeth present (Richardson 1904). Peri- 
toneal color is usually fleshy with many 
scattered melanophores, but occasional 
specimens have a more silvery ground 
color with melanophores scattered 
throughout. 

Dorsal coloration is dusky olive- 
brown or, in life, dark green with a 
somewhat lighter venter of yellowish 
brown. Many dark coffee-colored spots 
occur over the body, often one spot per 
scale, creating vague, irregular vertical 
bands or longitudinal rows. The belly, 
breast, throat, and chin have many 
tiny, dark melanophores. Some spec- 


Ivtinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 10 


Fig. 3—Lepcmis symmetricus prejuvenile 
12.0 mm in standard length (above) and ju- 
venile 30.0 mm in standard length (below). 


imens are almost solid black on the 
midbody with discrete black punctate 
marks on the cheeks. The fins, except 
the pectorals, are dusky overall with 
the soft dorsal and anal usually having 
several light spots. The pectoral fin 
rays are outlined by melanophores but 
are otherwise clear. The cheeks and 
head are very dark and have no pat- 
terns. The dark opercular spot is 
usually bordered with a light area on 
its posterior margin. 

Juveniles contrast with adults in 
generally having more distinct vertical 
bands, in always having a black spot 
in the soft dorsal, and in having some 
red-orange pigmentation in both the 
soft dorsal and soft anal fins. Juveniles 
are lighter overall than adults and gen- 
erally have seven to nine rather distinct 
vertical bands that are darker (brown) 
than the overall light greenish ground 
color. The vertical barring is occasion- 
ally obscured by flecks of darker pig- 
ment over the body, giving it a spotted 
appearance. The juveniles of both 
sexes have a distinct black blotch on 
the last five to eight rays of the soft 


Sept., 1977 


dorsal fin; the pigment is distributed 
both on the radial and _ interradial 
membranes. Rarely, there is a black 
spot in the soft anal fin (NLU 2907, 
2 of 21 specimens; INHS 18151, 1 of 
45 specimens) on the last three rays, 
and again the pigment is both on the 
radial and interradial membranes. Red- 
orange pigmentation is also present in 
the soft dorsal and soft anal fins of 
both sexes, on the radial and inter- 
radial membranes, and is very prom- 
inent in specimens collected during the 
fall and winter months. The belly, 
breast, throat, and chin sometimes are 
marked with discrete, tiny brown mel- 
anophores. Jordan (1884:320—321) re- 
marked that small specimens from New 
Orleans had faint blue spots on the 
sides of their heads. Breeding color- 
ation is discussed under Reproductive 
Cycles of both sexes. 


VARIATION 
Sexual 


No sexual variation in meristic char- 
acters was noted, but some dimorphism 
in one proportional character and in 


iff S ry VY q ‘ a3 
c NC} D 

Fig. 4—Genital papillae of Lepomis sym- 
metricus. A, nonbreeding male; B, breeding 
male; C, nonbreeding female; D, breeding 
female. The nonbreeding specimens were 1+ 
years old, collected on 19 October 1973; the 


breeding specimens were 2 years old, collected 
on 27 April 1974. 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


445 


sex organs was evident. Pelvic fin 
length is significantly greater at the 
0.05 level (F = 8.29) in the male than 
in the female (Table 5). The urogen- 
ital papilla of the adult female is en- 
larged and protruding during the 
spawning season, whereas that of the 
adult male is only slightly enlarged 
(Fig. 4). The male is not appreciably 
larger than the female. The largest 
individuals from the study area were 
females (61 and 63 mm SL), the 
largest specimen examined from 
throughout the range was a female 
(75.5 mm SL). The largest male was 
73.5 mm SL. 


Allometric 


No allometric variation in meristic 
characters was found. Although allo- 
metric variation in morphometric char- 
acters was not investigated, adults are 
more robust than juveniles, as in other 
sunfishes. Moreover, it is the adult that 
is symmetrical in shape and thus is re- 
sponsible for the trivial name of the 
species. Juveniles have body propor- 
tions similar to those of other juvenile 
sunfishes. The number of vertical 
bands, if present at all, is the same in 
the juvenile and adult. The most no- 
table allometric variation is the ten- 
dency for the black spot in the soft dor- 
sal fin to become more diffuse and 
weak with age. It is prominent in the 
smallest young and absent in the adult, 
except in an occasional female. The 
soft anal and soft dorsal fins have a 
red-orange coloration that disappears 
when the fish becomes adult. 


Geographic 

Geographic variation in some meris- 
tic characters was evident when samples 
were grouped according to major river 
systems and arranged in a_ north-to- 
south order from the Mississippi drain- 
age of Illinois; through the Ouachita 
and Red river drainages of Arkansas, 
Oklahoma, Louisiana, and Texas; to 


446 


the Gulf Coast drainages of Texas and 
Louisiana (Tables 1-4). 

No significant geographic variation 
was found in any of the body propor- 
tions measured (Table 5). In fact, in 
this respect L. symmetricus is remark- 
ably conservative for a species with a 
rather long north-to-south distribution 
(Fig. 1). These meristic characters 
varied clinally: numbers of caudal- 
peduncle scales, lateral-line scales, and 
anal soft rays. The number of dorsal 
soft rays showed a slight but somewhat 
irregular trend toward more ray ele- 
ments in the north (Table 3). The 
Mississippi drainage samples from Ar- 
kansas, Louisiana, Tennessee, Missouri, 
Illinois, and Kentucky were intermedi- 
ate in caudal-peduncle and lateral-line 
scale counts between Red River-Gulf 
Coast samples and those from the Wa- 
bash drainage of Illinois. In these 
counts (Tables 1 and 2) the samples 
showed a gradual increase toward the 
north, whereas the soft-ray counts (Ta- 
bles 3 and 4) were more discordant, 
with specimens from the Red River- 
Gulf Coast samples having means close 
to that of the Illinois River specimens. 

The most aberrant samples are those 
that formerly occurred in oxbow ponds 
along the Wabash River in White 
County, Illinois. They have a slightly 
higher mean number of lateral-line 
scales and slightly higher mean num- 
ber of caudal-peduncle scales, but they 
have lower means for the soft fin ray 
counts than samples from the upper 
Mississippi drainage (Tables 1-4). 

No apparent geographic trends in 
coloration or pattern could be _per- 
ceived. Individual variation occurs in 
the prominence of the vertical bars and 
overall darkness, due perhaps in part 
to the strength of the preservative and 
age of the individuals. 


RELATIONSHIPS 

Because of various features of mor- 
phology, cytology, and paleontology, 
L. symmetricus has been considered to 


ILLINOIS NATURAL HISTORY SURVEY BULLETIN 


Vol. 31, Art. 10 


be most closely related to L. cyanellus 
(Bailey 1938; Branson & Moore 1962) 
as a highly specialized congener with 
several unique characters. Hubbs (in 
Jordan 1929) considered L. symmetricus 
distinctive enough to warrant place- 
ment in a new monotypic genus, Le- 
thogrammus, and Bailey (1938), adopt- 
ing the use of subgenera, placed L. 
symmetricus in the subgenus Lethog- 
rammus. 

More recent studies on species of 
Lepomis using the techniques of elec- 
trophoresis (Avise & Smith 1974), hy- 
bridization (Hester 1970), and chro- 
mosome analysis (Roberts 1964) have 
not included specimens of L. symmetri- 
cus. Thus, it is not known where the 
species would be placed in the classi- 
fication schemes presented by these 
authors. 


SPECIMENS STUDIED 


The following list includes only 
those collections of L. symmetricus that 
were used for meristic and morpho- 
metric features. Others were used for 
the assessment of distribution, descrip- 
tive features, and life-history data. Col- 
lections are listed generally from north 
to south. The number of specimens ex- 
amined is given in parentheses follow- 
ing the catalog number. Specific local- 
ity data may be obtained upon request 
from the author. 


Ohio River Drainage 


WabBasH RIVER SYSTEM.—ILLINOIS, 
White County: 2 October 1882, INHS 


75008 (1); 1 October 1882, INHS 
75009 (1); 3 October 1882, INHS 
75007 (10) . 


Mississippi River Drainage 

ILLiNnois RIVER sysTEM.—ILLINOIS, 
Tazewell County: 16 April 1880, 
INHS 75004 (1), INHS 75005 (7), 
USNM 29864 (1); 2 June 1880, INHS 
75006 (2). 

CLEAR CREEK sYSTEM.—ILLINOIS, 


Sept., 1977 


Union County: 18 July 1883, INHS 
75102. (5); +16 September 1959, 
INHS 17547 (6) ; 27 April 1963, INHS 
17566 (1); 27 May 1965, INHS 17583 


(1) ; 31 August 1970, INHS-17557 (1); - 


21 June 1973, INHS 18143 (5); 25 July 
1973, INHS 18151 (2); 24 January 
1974, INHS 75025 (6); 28 March 1974, 
INHS 75022 (1); 30 May 1974, INHS 
75021 (1); 27 May 1975, INHS 75020 
(1). 

OBION CREEK SYSTEM.—KEN- 
TUCKY, Hickman County: 21 Jan- 
uary 1964, INHS 75024 (1); no date, 
UL 5617 (10). Fulton County: 15 
June 1948, UL 10691 (4). 

SAINT FRANCIS RIVER sysTEM.—MIS- 
SOURI, Stoddard County: 25 October 
1973, INHS 75023 (10). 

NATURAL LAKES AND BACKWATERS.— 
TENNESSEE, Lake County: 11-13 
March 1968, UT 90.27 (8); 8 April 
1950, FMNH 80532 (2). Lauderdale 
County: 9 October 1972, UT 90.102 
(2). ARKANSAS, Chicot County: 17 
August 1974, HWR 74-35 (8). 

FORKED DEER RIVER sySTEM.—TEN- 
NESSEE, Haywood County: 3 Novem- 
bemlo7a Ue 90138) (1) 527 April 
1974, UT 90.140 (6). Gibson County: 
19 October 1973, UT 90.139 (10) . 

L’ANGVILLE RIVER SYSTEM.—AR- 
KANSAS, St. Francis County: 7 Au- 
gust 1939, UMMZ 128537 (2). 

ARKANSAS RIVER SYSTEM.—ARKAN- 
SAS, Arkansas County: 13 August 
1974, ARP-79 (10). 

OuacniTa RIvER sySTEM.—ARKAN- 
SAS, Bradley County: 23 May 1974, 
UT 90.116 (1), HWR 74-8 (1); 10 Au- 
gust 1974, HWR 74-26 (7). Calhoun 
County: 6 October 1974, JLS 74-14 
(2). Union County: 25 April 1975, 
NLU 31455 (10). LOUISIANA, Oua- 
chita Parish: 17 October 1964, NLU 
$94) (5). 

Rep RIVER 
Little River County: 


SYSTEM.—ARKANSAS, 
13 September 


1940, UMMZ 170879 (1). OKLA- 
HOMA, McCurtain County: 20 Au- 
gust 1948, UMMZ 155830 (1). LOU- 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


447 


ISIANA, Red River Parish: 22 June 
1965, NLU 1954 (7). Winn Parish: 
23 June 1965, NLU 1989 (7). Caddo 
Parish: 22 February 1969, NLU 12804 
(5). TEXAS, Bowie County: 24 May 


1957, TNHC 4984 (10). Harrison 
County: 17 March 1972, TOCWC 
4068.14 (2). 

LAKE PONTCHARTRAIN.—LOUISI- 


ANA, Orleans Parish: 
NLU 29918 (5). 


15 April 1974, 


Gulf Coast Drainage 


CaALcasiEU RIVER sySTEM.—LOUISI- 
ANA, Calcasieu Parish: 10 August 
1965, NLU 2534 (6). Allen Parish: 10 
August 1965, NLU 2907 (5). Jefferson 
Davis Parish: 10 August 1965, NLU 
2909 (4). 

MERMENTAU-TECHE RIVER SYSTEM.— 
LOUISIANA, Avoyelles Parish: 20 
April 1975; NEU 31572 (3) 

NECHES RIVER SySTEM.—TEXAS, Jef- 
ferson County: 2 May 1970, TCWC 
3643 (14). Hardin County: August 
1950, TNHC 585 (1). Newton County: 
7 June 1952, TNHC 2889 (3). 

TRINITY RIVER SYSTEM.—TEXAS, 
Chambers County: 14 July 1953, 
MNELE 3873) \(2)y. 

SAN JACINTO RIVER SYSTEM.—TEXAS, 
Montgomery County: 23 March 1951, 
TNHC 1211 (1). 


DISTRIBUTION 


All known locality records for L. 
symmetricus are plotted in Fig. 1. 
Along the Gulf Coast the species ex- 
tends from Eagle Lake (Colorado River 
drainage, UMMZ 129793) in Texas 
east to marshes of the Jordan River sys- 
tem in Mississippi. In the Mississippi 
Valley it presently extends north to the 
bottomland oxbow lakes and swamps 
of southern Illinois. A published rec- 
ord for the St. Joseph River of Mich- 
igan (Dolley 1933) is clearly based on 
a misidentification, as Michigan is far 
out of the range of the bantam sunfish. 

L. symmetricus is now almost en- 


448 


tirely restricted to the Coastal Plain. It 
formerly traversed the Coastal Plain 
boundary far northward to the Illi- 
nois River (at Pekin) and backwater 
ponds and sloughs of the Wabash River 
system in White County, Illinois (Fig. 
1). The species has not been collected 
from the type-locality since 1880, a fact 
which Richardson (1904) noted only 
24 years after its original description. 
Indeed, it was collected only twice from 
Pekin. It has not been collected from 
the Wabash valley since 1882, whence 
it was known from three localities and 
12 specimens (INHS 75007, 75008, 
75009). The distribution of L. sym- 
metricus has thus changed rather dra- 
matically in Illinois, the decimation 
probably being the result of radical 
changes brought on by human modifi- 
cations, notably the stocking of non- 
native sunfishes, a reduction in aquatic 
vegetation, draining of lowland swamps 
and sloughs, and various forms of ag- 
ricultural and industrial pollution 
(Smith 1971). Mills et al. (1966) 
clearly demonstrated the effects of hu- 
man modification on the fauna and 
flora of the Illinois River, and the fac- 
tors listed above almost surely caused 
the extirpation of the species from the 
Pekin area. It is also possible that the 
relatively short life span of the species 
(3+ years) is somehow associated with 
its fairly rapid extirpation from dis- 
turbed or polluted areas in the Missis- 
sippi Valley of Illinois, Missouri, and 
Kentucky. 

The species is virtually absent east of 
the Mississippi River in Mississippi. 
Perhaps the Mississippi River has been 
an effective barrier to dispersal in this 
region, or the species’ apparent absence 
there may be because collectors tend to 
avoid swamps, sloughs, and lowland 
streams. ‘The species is statewide in 
occurrence in Louisiana, where it is 
common, and it is rather common in 
eastern Texas, southern Arkansas, and 
parts of western Tennessee (Fig. 1) . 


ILLino1s NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 10 


The distribution of L. symmetricus 
suggests that it is autochthonous to the 
lower Mississippi River valley (Pflieger 
1971:413-414). It apparently dispersed 
through oxbow lakes, swamps, and 
sloughs, created by varying water levels 
during the history of the Mississippi 
River. (Pflieger (1971:414) suggested 
that L. symmetricus may have had its 
origin in the lower Mississippi valley, 
dispersed northward to central Illinois 
during the postglacial Climatic Opti- 
mum, and become disjunct in its north- 
ern distribution subsequently. 


CONSERVATION STATUS 


Miller (1972) listed L. symmetricus 
as rare in both Illinois and Missouri in 
a compilation of threatened fishes of 
the United States. At that time it was 
known in those states from only two 
localities: the LaRue-Pine Hills area 
of southwestern Illinois (Union County) 
and the Duck Creek Wildlife Area of 
southeastern Missouri (Bollinger 
County) , where it has been reported to 
be common (Pflieger 1971:413). It has 
since been found to be common in 
Wolf Lake, Illinois, and Mingo Na- 
tional Wildlife Refuge, Missouri (Pflie- 
ger 1975:265). The species is on the 
protected list of both states but not 
presently endangered in either because 
its habitat is now rigidly protected in 
refuges. Recently, Webb & Sisk (1975: 
69) recommended that L. symmetricus 
be placed on Kentucky's rare and en- 
dangered species list in view of its rar- 
ity in Kentucky. 

In Oklahoma the species is found 
only inthe swamps of McCurtain 
County in the southeastern corner of 
the state (Fig. 1). L. symmetricus was 
not considered threatened by Robison 
et al. (1974) in their list of threatened 
Oklahoma fishes, but it may presently 
be reduced in numbers according to 
Hubbs & Pigg (1976:116). In Arkan- 
sas the status of the species was listed 
as indeterminate by Buchanan (1974), 


Sept., 1977 


but L. symmetricus was not cited by 
Robison (1974) in his list of threat- 
ened Arkansas fishes. The species is 
apparently in no danger in southern 
Arkansas (Fig. 1), where it is known 
from many localities. 


LIFE HISTORY IN WOLF LAKE 
STUDY AREA 

Wolf Lake is a long (ca. 1.9 km), 
narrow (ca. 0.1 km), and ancient ox- 
bow of the Big Muddy River (Missis- 
sippi drainage) situated south of the 
LaRue-Pine Hills Ecological Area to 
which it is connected by bottomland 
swamp. The lake is apparently still in 
a fairly natural, undisturbed condition 
and is estimated to be at least 2,000 
years old (E. Donald McKay IJ, per- 
sonal communication). The northern 
portion of the lake was recently ac- 
quired by the U. S. Forest Service, 
whereas the southern portion of the 
lake is privately owned by the Trojan 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


449 


Powder Plant. Most observations and 
collections in Wolf Lake were made 
near the powder plant bridge, where 
access to the lake was easy although 
other portions of the lake were 
sampled. 


HABITAT 


Wolf Lake is characterized by two 
predominant habitats: a heavily veg- 
etated shoreline with many submerged 
logs and stumps (Fig. 5) and an open 
deepwater area in the center of the lake 
free from vegetation and submerged ob- 
jects. The lake is not shaded and the 
water is usually turbid. The vegetated 
shoreline, where L. symmetricus occurs 
(Fig. 5), is dominated by spatterdock 
(Nymphaea advena), American lotus 
(Nelumbo lutea), common arrowhead 
(Sagittaria latifolia), coontail (Cera- 
tophyllum demersum) and duckweed 
(Lemna spp., Wolffia spp.). The bot- 


tom consists mostly of decomposed veg- 


Fig. 5—Vegetated margin of Wolf Lake, Union County, Illinois, illustrating the preferred 


habitat of Lepomis symmetricus. 


Photo taken in May 1974. 


450 


etation, silt, and mud, with some sand. 
Water depth ranges from 300 mm to 18 
meters. Dissolved oxygen averages 9.0 
ppm; temperatures range from 4° to 
8° C from December to February and 
are as high as 29° C in July and August. 

L. symmetricus was found in similar 
habitat during a I-year study of fishes 
in the adjacent LaRue-Pine Hills 
swamp (Boyd et al. 1975). During the 
fall and winter months L. symmetricus 
was characteristically found at a depth 
of 150-300 mm usually near the shore- 
line in Wolf Lake. During the summer 
months the species could be found at 
depths of 600-1200 mm but still within 
the vegetated periphery of the lake. 

Elsewhere in its range L. symmetri- 
cus is invariably found in lentic waters 
characterized by standing timber, sub- 
merged logs and stumps, and rich vege- 
tation. Sloughs, oxbows, ponds, back- 
waters, lakes, and swamps typical of 
the undisturbed portions of the Coastal 
Plain are optimal habitat. L. symmet- 
ricus is found in greatest numbers over 
substrates consisting of mud, detritus, 
and decayed plant material. 

Although L. symmetricus is syntopic 
with several other species of Lepomis 
in Wolf Lake, it was almost always col- 
lected by itself in the areas mentioned. 
The other Lepomis were usually taken 
in more open areas and generally in 
deeper water. In Wolf Lake the fishes 
most often found with L. symmetricus 
in descending order of association were 
L. macrochirus, L. gulosus, Pomoxis 
nigromaculatus, Notemigonus crysoleu- 
cas, Gambusia affinis, Micropterus sal- 
moides, Elassoma zonatum, and Etheos- 
toma gracile. Other inhabitants of the 
habitat of L. symmetricus occurring in 
less frequent numbers are Lepisosteus 
oculatus, L. platostomus, Dorosoma ce- 
pedianum, Umbra limi, Cyprinus car- 
pio, Ictiobus cyprinellus, Ictalurus nat- 


alis, I. nebulosus, Fundulus dispar, 
Aphredoderus sayanus, Centrarchus 
macropterus, Lepomis microlophus, 


and L. punctatus. 


ILtinois NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 10 


REPRODUCTION 
Reproductive Cycle of the Male 

The genital papilla (Fig. 4) of ripe L. 
symmetricus males enlarged slightly as 
the spawning season approached. The 
testes, normally small, translucent, and 
elongate, became large, opaque white, 
and thickened. 

Breeding males (Fig. 2), in contrast 
to non-breeding males and females 
(which were nearly identical in color 
and pattern) , became very dark on the 
head, and the irregular vertical cross 
bars grew subdued. The venter from 
the chin and throat to the anterior rays 
of the anal fin became grayish black. 
Many small greenish flecks were present 
on the head and opercle, and the dark 
opercular spot was outlined by a silvery- 
cream color with a hint of suffused red. 
The pectoral fins were relatively dusky 
overall but with no definite patterns. 
The posterior edges of the pelvic fins 
were almost solid black with the re- 
mainder of the fins cream color. The 
dorsal fin had many light spots sur- 
rounded by dusky brown or black areas. 
The iris of the eye was brilliant red 
with a distinct black transverse bar 
through it. 

Because of the silty darkly-stained 
water of Wolf Lake, no nests of L. sym- 
metricus could be observed in nature, 
and nothing is known of territory size. 
However, Robison (1975:56) reported 
that on 23 May 1974 in a roadside pool, 
Saline County, Arkansas, L. symmelrt- 
cus had recently spawned, inasmuch as 
“depressions in the mud and leaf litter 
substrate were filled with numerous 
eggs.” Since males were observed to be 
highly aggressive toward females and 
other sunfishes are known to be terri- 
torial (Larimore 1957), it is assumed 
that L. symmetricus defends an area in 
nature. An aquarium-held male col- 
lected in May was seen on several occa- 
sions to form a shallow nest by rapidly 
swimming forward, then turning his 
body straight up in a vertical position 


Sept., 1977 


and descending, sweeping his tail vig- 
orously back and forth until a nest de- 
pression was formed. Such nests were 
formed over both sand and gravel sub- 
strates. These nests were approximately 
90-120 mm in diameter. It is likely 
that L. symmetricus males build shal- 
low depressions in the mud bottom of 
Wolf Lake along the shallow edges 
close to the vegetation where ege at- 
tachment may take place. This behav- 
ior has been described for L. cyanellus 
(Hankinson 1908:210-211). 

Only large males developed the 
breeding patterns, the slightly enlarged 
genital papilla, and the enlargement of 
the testes. Only males of at least 1+ 
years and 40 mm or longer appeared to 
be sexually mature, according to color- 
ation and condition of the testes. The 
largest males probably do most or all 
of the spawning. 


Reproductive Cycle of the Female 


Generally the largest females devel- 
oped the earliest mature ova and prob- 
ably contributed most to the spawning 
effort. Females as short as or shorter 
than 34 mm and | year of age devel- 
oped mature ova and were potential 
spawners. 

Females underwent some changes in 
coloration associated with the breeding 
season. In contrast to males and non- 
breeding individuals, the breeding fe- 
male had 9 or 10 distinct vertical bars 
of a dark bluish-purple color with light 
greenish flecks in the spaces between 
the bars. The cheek and opercle con- 
tained bright spots of golden green, but 
the fins were relatively clear and not 
dusky. Some females retained a diffuse 
ocellus in the posterior rays of the dor- 
sal fin. As in males, the iris was bright 
red. Other marked morphological 
changes were the distended belly caused 
by the maturing ova and the enlarge- 
ment of the genital papilla (Fig. 4). 
Enlargement of the papilla was notice- 
able only in ripe females. 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


451 


Small white ova were present in fe- 
males 1+ years of age and 35 mm long 
as early as September but were difficult 
to distinguish in younger and smaller 
females. By January and February 
larger yellowish ova were found in 1+- 
year females of 40 mm or longer. Large, 
coarse, maturing orange ova were pres- 
ent from March to May in larger and 
older females and in some smaller fe- 
males over 54 mm and approaching 1 
year of age. Just prior to spawning 
time, the mature ova became a translu- 
cent orange. 

The largest and oldest females pro- 
duced the largest number of mature 
ova. In 14 ripe females collected in 
April and May the number of ova var- 
ied from 219 to approximately 1,600 
(Table 6). For these females the rela- 
tionship between the number of ma- 
ture ova (F) and the adjusted body 
weight (W) was F = —50.94 + 210.70W, 
with r = 0.818, and between the num- 
ber of mature ova and the standard 
length (L) was log F = —2.785 + 3.383 
log L, with r = 0.663. 

Ovaries of postspawning females col- 
lected in June were smaller than those 
of females collected in April and May. 
They averaged slightly heavier than 
ovaries from females collected in 
March. Ovaries from females taken in 
July and August were small. A relative 
increase In ovary size was evident by 
late fall and continued to the spawning 
period the following spring (Fig. 6). 
For the females examined, the relation- 
ship between the weight of the ovaries 
divided by the adjusted body weight 
(Y) and the month (X), with July = 
1 and May = 11, was log Y = 0.699 + 
0.099X, with r = 0.782 (Fig. 6). The 
proportionally largest ovaries (equaling 
30.8 percent of the adjusted body 
weight) were found in a 5l-mm, 2- 
year-old female collected on 27 April 
1974 (UT 90.140). In the 14 females 
represented in Table 6, overy-weight- 
to-adjusted-body-weight ratios ranged 
from 0.070 to 0.308 and averaged 0.107. 


452 


ILLINOIS NATURAL HISTORY SURVEY BULLETIN 


Vol. 31, Art. 10 


Table 6.—Relationship between size, age, and ovary weight of Lepomis symmetricus fe- 
males and the number of mature ova produced. An age of 1 year = 11—13 months, 2 years = 


23-25 months. 
Wolf Lake, are included. 


Adjusted 


Standard Bod \ 
Length Weigh t in 
Ee in Grams* 

34 1.42 1 
34 1.49 1 
36 1.78 1 
37 1.65 1 
37 2.06 1 
38 2.32 1 
39 2.11 1 
40 2.43 ] 
42 2.44 1 
43 2.75 1 
45 3.22 1 
45 3.34 1 
51 4.51 2 
52 7.57 2 


Age 


Years 


Data from TCWC 3643, UT 90.140, and INHS 17583, as well as that from 


Number of 
Ovary Mature 

: orange or 

Weight ee 

pee 0.6-0.9 mm) 
Ova 
0.10 326 
0.12 219 
0.13 491 
0.20 368 
0.18 403 
0.20 330 
0.21 432 
0.18 417 
0.26 421 
0.20 374 
0.33 364 
0.31 378 
1.39 ca 1600 
0.88 ca 1400 


= Adjusted body weight is the specimen’s weight after removal of the ovaries, stomach, intestine, and liver. 


500 


LOG Y=0.699+0.099X 


WEIGHT OF OVARIES X 1000/ADJUSTED BODY WEIGHT 


JULY SEPT. NOV, JAN, MAR, MAY 


Fig. 6—Monthly variations in ovarian 
weight relative to adjusted body weight of 
Lepomis symmetricus. The vertical axis is on 
a logarithmic scale. Ovaries from specimens 
collected June to April were from all age 
classes, but ovaries from specimens collected 
in May were from 2-year-old (24 months) 
fish. 


Spawning 

In Wolf Lake breeding individuals 
were captured as early as 24 April and 
as late as 30 May. Most spawning prob- 
ably occurred in May when water tem- 


peratures ranged from 18° to 22° C. 
Field observations and examination of 
museum specimens collected during all 
months of the year indicated that mid- 
April to early June was the typical 
spawning period for the species 
throughout its range (Table 7) . 
Although spawning was not observed 
in the study area, ripe aquarium-held 
individuals collected 27 May 1975 en- 
gaged in prespawning activity for 7 
days at water temperatures varying 
from 24° to 26° C. After presumed 
stimulation from a recent feeding the 
male began to court the female by 
nudging her with his snout along the 
posterior regions of her body and con- 
tinually nipping at her caudal fin. The 
female did not respond to these actions 
but the male continued to nip at her 
fins and nudged the female with his 
snout between the pelvic fins while 
chasing her. The female remained un- 
responsive. After 3 days of this behay- 
ior the male began to charge the fe- 
male at rapid speeds with his opercles 
flared out and with the irises of his 
eyes more intense in color than before. 


Sept., 1977 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


453 


Table 7.—Collections of breeding Lepomis symmetricus. 


Locality 


Collection Date 


Remarks 


Wolf Lake, Union Co., Ill. 
(INHS 75020, 75021) 


Pine Hills Swamp, Union Co., Ill. 
(INHS 17583) 


Illinois River, Tazewell Co., Ill. 
(INHS 75006) 

Swamp, Haywood Co., Tenn. 
(UT 90.140) 

Reelfoot Lake, Lake Co., Tenn. 
(FMNH 80532) 


Roadside Ditch, Bradley Co., Ark. 
(UT 90.116) (HWR 74-8) 

Ouachita River, Union Co., Ark. 
(NLU 31455) 

Big Hill Oil Field, Jefferson Co., Tex. 
(TCWC 3643) 

Marsh, Orleans Parish, La. 
(NLU 29918) 


Creek, Avoyelles Parish, La. 
(NLU 31572) 


24 April-30 May 1974 
27 May 1965 

2 June 1880 

27 April 1974 

8 April 1950 

23 May 1974 

25 April 1975 

2 May 1970 

15 April 1974 


20 April 1975 


Males and females in extreme 
breeding condition. 


Female in breeding condition. 
Females in breeding condition. 


Males and females in extreme 
breeding condition. 


Males and females in breeding 
condition. 


Male and female in breeding 
condition. 


Females in extreme breeding 
condition. 


Males and females in breeding 
condition. 


Males and females in breeding 
condition. 


Males in breeding condition. 


When he approached the female, he 
abruptly turned himself to a vertical 
position (with his snout pointing up- 
ward) and gently swam around her in 
a close circle while fanning his tail. 
Similar courtship patterns were de- 
scribed by Larimore (1957) for L. gu- 
losus. After 7 days of constant nipping, 
nudging, badgering, and displaying 
other prenuptial behavior, the male 
had succeeded in completely mutilat- 
ing the uncooperative female’s caudal 
fin, and on the 8th day the female was 
found dead. Even though an actual 
egg-laying session did not take place, it 
is evident that the nest building and 
prespawning behavior of L. symmetri- 
cus does not vary greatly from that de- 
scribed for other species of Lepomis 
summarized by Breder & Rosen (1966) . 


DEVELOPMENT AND GROWTH 


Mature ova ranged in size from 0.6 
to 0.9 mm in diameter, were translu- 
cent orange, and contained a single oil 
droplet. No data are available on incu- 
bation temperatures of eggs, the length 


of time required for hatching, or the 
morphology of hatchlings. 

The smallest L. symmetricus individ- 
ual from the study area was 12 mm, 
collected 21 June 1973 (Fig. 3). At 
this size the nape, breast, and sides of 
the head were the only regions incom- 
pletely scaled, but no definite pigment 
pattern was present. Small melano- 
phores outlined the scale borders on 
the body and some of the fin rays but 
were concentrated heavily on the top 
of the head, on the lips, and around 
the eye. The soft dorsal fin ocellus was 
just beginning to develop (Fig. 3). 

A series of 43 young L. symmetricus 
from 14.0 mm to 25.0 mm was collected 
in the study area on 25 July 1973. At 
14 mm squamation patterns were like 
that at 12 mm, but many more melano- 
phores were present in the fins and 
they began to form patterns on the 
body. The ocellus was dark at this size. 
At 19 mm vague vertical bars had 
formed, and squamation was nearly 
complete. At 25 mm the lateralis sys- 
tem was developed, and the overall pig- 
ment pattern was similar to that of 


454 


adults. Squamation was complete at 
this stage. At a slightly larger size ju- 
veniles began to take on the form, 
pattern, and coloration illustrated in 
Fic, 3. 

L. symmetricus from Wolf Lake grew 
at a decreasing rate (Fig. 7) and reached 


Y=5.91+ 32.97 LOG X a 


STANDARD LENGTH, sm 


ipa 


02" 4 6 8 18 20 22 24 34 36 38 


10 12 14 16 
MONTHS OF AGE 


Fig. 7—Size distribution by age of Lepomis 
symmetricus collected in Wolf Lake between 
21 June 1973 and 30 May 1974. Data from 
27 May 1975 and 12 December 1974 are 
included. Black dots represent sample means 
for both sexes combined. In total, 233 speci- 
mens are represented. 


one-half of the first year’s mean growth 
in approximately 10 weeks. The rela- 
tionship between standard length (Y) 
and age in months (X) expressed for 
the sexes combined is Y = 5.91 + 32.97 
log X, with r = 0.943. Males grew at a 
slightly more rapid rate than females 
but were not significantly larger than 
females. At 13-18 months males aver- 
aged 45.9 mm and females averaged 
42.7 mm (¢ = 1.39, df =11). At 19-24 
months males averaged 49.3 mm and 
females averaged 47.5 mm (¢t = 1.00, 
df=14). The largest specimen exam- 
ined from Wolf Lake was a 63.0-mm 
female collected 25 July 1973. In other 
parts of its range L. symmetricus is 
known to attain a greater length, and 
specimens as long as 75.5 mm_ have 
been collected (TU 148—St. Tam- 
many Parish, Louisiana). Based on 
the collections examined, such large 
size is unusual, with most adults rang- 
ing between 55 and 60 mm. 


ILLINOIS NATURAL History SURVEY BULLETIN 


Vol. 31, Art. 10 


DEMOGRAPHY 
Density 


The nature of the habitat of L. sym- 
metricus made population density mea- 
surements difficult, since submerged 
logs, brush, and vegetation prevented 
thorough sampling of a given area. 
However, on two occasions approx- 
imately 5 months apart quantitative 
samples of L. symmetricus were taken 
in Wolf Lake by repeatedly seining a 
measured shallow margin of the lake 
until no more individuals could be 
collected. The number collected was 
translated into the number per square 
meter. The greatest density found for 
L. symmetricus in Wolf Lake was 0.69 
sunfish per square meter (Table 8). 

In the nearby LaRue-Pine Hills 
swamp, the density of L. symmetricus 
may approach 0.72 sunfish per square 
meter (Table 8) or, at best, 1 individ- 


Table 8.—Number of Lepomis symmetricus 
per square meter collected in vegetated mar- 
gins of Wolf Lake and LaRue-Pine Hills swamp. 


Number 
of L. 
symmetricus 


Number per square 
Date Collected meter in 
Wolf Lake 
and 
Pine Hills 
25 October 1973, 

Wolf Lake 9 0.313 
28 March 1974, 

Wolf Lake 20 0.694 
Mean 0.504 
24 October 1973, 

Pine Hills 6 0.723 
28 November 1973, 

Pine Hills 4 0.542 
27 March 1974, 

Pine Hills 1 0.114 
Mean 0.460 


ual per 3 square meters in optimal 
habitat (Boyd et al. 1975). Of 31 in- 
dividuals collected during 9 months at 
several collecting sites in Pine Hills, L. 
symmetricus made up 2.3 percent of 
the total sample of fishes captured. 


Sept., 1977 


However, more than 80 percent of the 
individuals were captured at one site 
where the habitat was judged to be op- 
timal (Boyd et al. 1975). Gunning & 
Lewis (1955) found that L. symmetri- 
cus made up 5 percent of their total 
sample of fishes at Pine Hills. 


Composition 

Of the 233 L. symmetricus collected 
in Wolf Lake, 85.4 percent were up to 
1 year of age, 12.4 percent were over 1 
and up to 2 years of age, 0.8 percent 
were over 2 and up to 3 years of age, 
and 1.2 percent were over 3 years of 
age (Table 9). 


Table 9.—Distribution of sexes and year 
classes in samples of Lepomis symmetricus 
collected in Wolf Lake between 21 June 1973 
and 30 May 1974, and on 27 May 1975 and 
12 December 1974. 


Number by Year Class 


Sex Total 
—l 1+ 2+ 3+ 

Males 81 16 1 an 98 

Females 118 13 1 3 135 

Total 199 29 2 3 233 


Females predominated in the young- 
of-the-year (—1) age class [1.5 females 
to 1 male (x? = 6.87; P < 0.01) ], and 
in the total sample (N = 233) the ratio 
was 1.4 females to 1 male (y? = 5.97; 
P < 0.025). Although predominating 
significantly in the —1 age class and in 
the total, females were slightly less 
common than males in the 1+ age 
class. 


Survival 


Relative survival values (Table 10) 
for each year of life were calculated for 
males, females, and the total sample of 
L. symmetricus, using the data in Ta- 
ble 9. It was assumed that each age 
class was collected in proportion to its 
relative number in the population, that 
the population was neither increasing 
nor decreasing, and that the number of 
fry entering the population each year 
was constant. 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


455 


Table 10.—Relative survival of year classes 
of Lepomis symmetricus in Wolf Lake ex- 
pressed as proportions of the —1 year class 
(1 x") and the 1+ year class (1 x). 


Number Survival 
Sample Year of : 
Class Speci- 
Tere Ine xe 
Males —l 81 1.000 bane 
1+ 16 0.198 1.000 
2: 1 0.012 0.063 
3+ fe iads 
Females —l 118 1.000 xara 
1+ 13 0.110 1.000 
2+ 1 0.008 0.077 
3+ 3 0.025 0.231 
Total 
sample =I 199 1.000 vee 
1+ 29 0.146 1.000 
2+ 2 0.010 0.069 
3+ 3 0.015 0.103 


Because of the difficulty in collecting 
in Wolf Lake, the numbers of 1+ and 
older individuals in Tables 9 and 10 
are probably lower than their actual 
proportion in the population. 

The shapes of the survival curves for 
males, females, and total sample were 
quite similar. All showed a very low 
survival rate after the Ist year of life. 
Only three individuals 3 years or older 
were found. The oldest L. symmetricus 
from Wolf Lake examined was a fe- 
male 3 years and 2 months old (as- 
suming May hatching) collected 25 
July 1973. 

Specimens from throughout the 
range further confirm a 3+ -year life 
span for the species: INHS 17547— 
from LaRue-Pine Hills Ecological Area 
collected 16 September 1959, contain- 
ing three individuals all 3 years 
and 4 months of age (assuming May 
hatching); NLU 31572—collected 20 
April 1975 from a creek, Avoyelles Par- 
ish, Louisiana, containing three indi- 
viduals 3 years of age. Most other spe- 
cies of Lepomis are much longer lived. 


DIET 


Stomach contents of 176 L. symmet- 
ricus from Wolf Lake were examined. 


456 


‘Twenty-nine of these contained no food 
items and eight contained green algal 
material. A large variety of food orga- 
nisms was found (Tables 11-14). The 
predominant food items of the Wolf 
Lake population were gastropods, cla- 
docerans, ostracods, amphipods, dragon- 
fly naiads, chironomids, and ceratopo- 
gonids. 

Small L. symmetricus (less than 21 
mm) fed predominantly on microcrus- 
tacea, dragonfly naiads, and chirono- 
mids; large individuals (more than 40 
mm) fed primarily on gastropods, 
dragonfly naiads, and amphipods (Ta- 
bles 11 and 12). Some seasonal vari- 


ILLiNnois NATURAL HISTORY SURVEY BULLETIN 


Vol. 31, Art. 10 


ation in diet (Tables 13 and 14) was 
evident. Gastropods were eaten in the 
winter and spring months. The largest 
percentages of most food items, includ- 
ing gastropods, stratiomyids, chirono- 
mids, and some microcrustacea, were 
eaten in the months prior to and dur- 
ing the spawning season, presumably 
reflecting an increase in consumption 
associated with spawning preparedness 
(Page 1974:17). Aquatic Hemiptera 
were eaten exclusively in the summer 
months, when they were most abun- 
dant. The presence in the diet of the 
exclusively terrestrial hemiteran family 
Fulgoridae reflects surface feeding by 


Table 11.—Stomach contents of Lepomis symmetricus from Wolf Lake, by size class of 
sunfish. Figures in parentheses are numbers of stomachs examined. 


Percent of Stomachs in Which Food Organism Occurred 


Food Organism <2] 21-30 31-40 41-50 51-60 >60 
mm mm mm mm mm mm 
(25) (44) (56) (19) (5) (5) 
Gastropoda 12.5 31.6 4.0 4.0 
Arachnida 
Araneae a 2.2 1.8 
Acarina 10.7 
Crustacea 
Cladocera 4.0 40.9 66.1 5.2 
Ostracoda 12.0 34.1 42.9 
Copepoda 8.0 18.2 32.1 otc oe 
Amphipoda 56.0 20.0 17.9 15.8 20.0 
Insecta 
Odonata 36.0 29.5 16.1 10.5 20.0 60.0 
Coleoptera 2 
Helodidae bY 2.2 20.0 
Noteridae yy 4.5 a 
Haliplidae 10.7 
Diptera 
Psychodidae 3.6 
Chaoboridae 1.8 
Tipulidae 1.8 ee ae 
Stratiomyidae aS AN 8.9 15.8 40.0 
Ceratopogonidae si¢ 45 Gige = 5.3 20.0 
Culicidae Se be 1.8 
Chironomidae 52.0 4.5 25.0 xo 20 Seg 
Ephemeroptera 26 6.8 7.1 Be an 20.0 
Trichoptera 5.4 5.3 
Hemiptera 
Corixidae re 9.1 8.9 21.1 
Naucoridae ar 6.8 1.8 ire te Bo 
Fulgoridae 36 be 1.8 = vs 20.0 
Pleidae 8.0 2.2 40.0 


Mesoveliidae a 2.2 


Sept., 1977 Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 457 


Table 12.—Stomach contents of Lepomis symmetricus from Wolf Lake, by size class of 
sunfish. Figures in parentheses are numbers of stomachs examined. 


Mean Number of Food Organisms Per Stomach 


Food Organism <21 21-30 31-40 41-50 51-60 >60 
mm mm mm mm mm mm 
(25) (44) (56) (19) (5) (5) 
Gastropoda 0.29 0.84 0.60 2.40 
Arachnida 
Araneae 0.02 0.02 
Acarina 0.32 
Crustacea 
Cladocera 0.36 6.41 2.14 1.26 
Ostracoda 0.48 2.57 0.07 
Copepoda 0.12 0.91 1.25 5e a 
Amphipoda 1.92 0.91 1.20 0.47 12.8 
Insecta 
Odonata 0.52 0.45 0.32 0.11 0.20 1.60 
Coleoptera 
Helodidae 0.05 0.20 
Noteridae 0.07 aa 
Haliplidae 0.54 
Diptera 
Psychodidae 0.02 
Chaoboridae 0.52 
Tipulidae 0.04 5d ae 
Stratiomyidae a 0.25 0.21 4.00 
Ceratopogonidae 0.02 0.07 0.05 3.00 
Culicidae ate Re 0.03 
Chironomidae 0.08 0.16 0.52 ia 
Ephemeroptera 0.07 0.07 oe 0.20 
Trichoptera 0.05 0.05 
Hemiptera 
Corixidae 0.18 0.29 0.26 
Naucoridae 0.06 0.02 an 
Fulgoridae we a 0.02 0.20 
Pleidae 0.08 0.02 0.60 
Mesoveliidae as 0.02 a 
L. symmetricus when these insects L. symmetricus has been reported to 


alight on the water surface. 

Aquarium-held L. symmetricus fed 
in the typical Lepomis manner. When 
food was dropped into the water near 
them, they sucked it in or swam up 
near the food item and gulped it down 
before the food item fell to the bottom 
of the aquarium. Occasionally they 
fed off the bottom by sucking up food 
items. Spawning males and other indi- 
viduals fed readily on dragonfly naiads, 
chironomids, and live and frozen earth- 
worms. Miller & Robison (1973:184) 
reported aquarium-held specimens from 
Oklahoma feeding on “daphnia and 
small earthworms.” 


eat “dragon-fly nymphs and midge lar- 
vae” near Greenwood, Mississippi (Hil- 
debrand & Towers 1927:134; Cook 
1959:180). In 22 specimens from LaRue- 
Pine Hills, Illinois, the major food 
items were “‘aquatic snails, green algae, 
amphipods, and miscellaneous insects 
and insect larvae” (Gunning & Lewis 
1955:556) . 


INTERACTION WITH OTHER 
ORGANISMS 


Competition 


L. symmetricus occurs syntopically 
with all other described species of Le- 


458 


pomis (including the introduced L. 
auritus) except L. gibbosus, from which 
it is geographically separated. Because 
of its preferred habitat of heavily vege- 
tated, shallow, lentic or slow-moving 
water and its relative abundance there, 
it is doubtful that the species is geo- 
graphically limited to a great degree by 
its several congeners. 


Predation 


There are no literature reports of 
predation on L. symmetricus and no 
evidence of such predation was found 
in the Wolf Lake study. As potential 
predators five Muicropterus salmoides 


ILLiNo1s NATURAL HistoRY SURVEY BULLETIN 


Vol. 31, Art. 10 


(71.6-240.3 mm SL), four Pomoxis 
nigromaculatus (76.4-144.8 mm SL), 
one P. annularis (141.1 mm SL), five 
Lepomis gulosus (19.7-127.4 mm SL), 
four L. macrochirus (131.8-140.1 mm 
SL), one Centrarchus macropterus (82.4 
mm SL), and one Ictalurus natalis 
(124.8 mm SL) were preserved and 
later examined for ingested L. symmet- 
ricus. These predators were collected 
from all months of the year except July 
and December. A number of large gar 
(Lepisosteus oculatus, L. platostomus) 
were seen during the summer and fall 
months but were not collected. Per- 
haps these large, relatively common 


Table 13.—Stomach contents of Lepomis symmetricus from Wolf Lake by month of col- 


lection.* Figures in parentheses are numbers of stomachs examined. 


Percent of Stomachs in Which Food Organism Occurred 


Food Organism Feb. 


(18) 


Mar. 
(15) 


Jan. 
(17) 


April June 


Oct. 
(9) 


Nov. Dec. 
(13) (18) 


July Aug. Sept. 


(9) (7) (43), (17) (10) 


Gastropoda 294 I1.1 13.3 


Arachnida 
Araneae 
Acarina 


Crustacea 
Cladocera 
Ostracoda 
Copepoda 
Amphipoda 


17.6 44.4 


Insecta 

Odonata 

Coleoptera 
Helodidae 
Noteridae 
Haliplidae 

Diptera 
Psychodidae 
Chaoboridae 
Tipulidae 
Stratiomyidae 
Ceratopogonidae 
Culicidae 
Chironomidae 

Ephemeroptera 

Trichoptera 

Hemiptera 
Corixidae 
Naucoridae 
Fulgoridae 
Pleidae 
Mesoveliidae 


5.9 


29.4 se 
6.6 


5.9 


55.6 
22.2 


33.3 
22.2 
33.3 
11.1 


55.6 


33.3 


14.3 11.1 


7.7 5.6 


76.5 
52.9 

5.9 
35.3 


90.0 77.7 
11.1 


16.7 
16.7 
33.3 
11.1 


61.5 
23.1 
23.1 

7.7 


10.0 
10.0 


64.7 30.8 


11.8 


66.7 


1 


33.3 
11.1 


14.3 1.7 
11.1 
38.8 
5.6 


Oe 20.0 
14.3 


57.1 
14.3 


70.0 


143 


Stomach contents were not examined for May-collected specimens. 


Sept., 1977 


predators take some toll on the Wolf 
Lake population of L. symmetricus. 


Hybridization 

Schwartz (1972) did not report any 
accounts of hybridization involving L. 
symmetricus. No evidence of hybrid- 
ization was found in the Wolf Lake 
study area or in specimens examined 
from elsewhere. The small size of L. 
symmetricus, its preference for shallow, 
vegetated water, and its distinct breed- 
ing coloration probably preclude mis- 
mating of the parental species. Since 
there is ample habitat available in 
Wolf Lake and the fishes are presum- 
ably not unduly crowded, chances of 


Burr: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 


459 


hybridization are small (Hubbs 1955: 
ZrlB\ye 


Parasitism 


The Wolf Lake study population was 
rather heavily parasitized by plerocer- 
coids of the cestode Haplobothrium 
globuliforme. These plerocercoids oc- 
curred in a total of 44 of 176 stomachs 
(25 percent) examined. From one to 
five plerocercoids were found in each 
stomach. Usually the highest numbers 
occurred in stomachs of the —1 year 
class. Specimens were found during all 
months of the year except May and 
June. The plerocercoid stage of H. 
globuliforme normally encysts in the 


Table 14.—Stomach contents of Lepomis symmetricus from Wolf Lake by month of col- 


lection.* 


Figures in parentheses are numbers of stomachs examined. 


Mean Number of Food Organisms Per Stomach 


Food Organism Jan. Feb. 


(17) (18) 


Mar. April June 
(15) (9) 


Nov. Dec. 
(13) (18) 


July 
(7) (48) 


Aug. Sept. Oct. 
(17) (10) 9) 


Gastropoda OF 07) 70:93. 


Arachnida 
Araneae 


1.56 


0.29 0.11 


0.08 0.06 


Acarina 


Crustacea 
Cladocera 
Ostracoda 
Copepoda 
Amphipoda 

Insecta 
Odonata 
Coleoptera 

Helodidae 
Noteridae 
Haliplidae 
Diptera 
Psychodidae 
Chaoboridae 
Tipulidae 
Stratiomyidae 
Ceratopogonidae 
Culicidae 
Chironomidae 
Ephemeroptera 
Trichoptera 
Hemiptera 
Corixidae 
Naucoridae 
Fulgoridae 
Pleidae 
Mesoveliidae 


0.06 3.61 
7.11 


0.06 1.00 


0.06 0.11 


0.05 
1.61 
0.11 
0.06 


1.41 


0.41 0.39 


0.93 0.44 
0.93 
10.40 
3.40 
0.13 


1.67 
0.56 
1.00 
0.11 


11.35 
3.00 
0.06 
2.59 


20.00 13.22 


0.44 0.11 
0.07 


1.79 


0.40 
0.10 
0.73 0.42 = 1.47 
0.07 et 
ee 0.18 
3.33 


0.12 
0.12 
0.06 


0.06 O.11 2.14 
O09 0.40 
0.14 


1.00 
0.14 


0.14 
0.02 
0.02 
0.02 
0.02 


me lOO) 
0.12 
ae 0.14 
0.14 


17.23 
0.38 
0.69 
0.08 


0.31 


0.15 


0.77 


0.72 
0.78 
2.11 
0.11 


0.22 
0.39 
0.06 


4 Stomach contents were not examined for May-collected specimens. 


460 


liver of fishes and has been reported 
from a number of other fishes in both 
this and the adult stage (Hoffman 1967: 
233). No adults were found in the 
study population. 

One adult specimen of the acantho- 
cephalan Pomphorynchus bulbicolli 
was found in the stomach of an L. sym- 
metricus collected 24 April 1974 at 
Wolf Lake. Neither the cestode nor 
the acanthocephalan had been known 
to parasitize L. symmetricus. 

Dolley (1933) reported cestodes and 
trematodes from “Lepomis symmetricus 
in the St. Joseph River of Michigan,” 
but the misidentification of the host 
species is obvious, since L. symmetricus 
has never occurred in Michigan. Hoff- 
man (1967), who compiled a list of 
fish parasites, cited for L. symmetricus 
the trematodes Actinocleidus symmetri- 
cus, Cleidodiscus diversus, and Ancho- 
radiscus triangularis. Dr. Mary H. 
Pritchard informed me that Hoffman 
(1967) evidently cited as the species 


ILtino1s NATURAL HisToRY SURVEY BULLETIN 


Vol. 31, Art. 10 


name (A. symmetricus) that of the host 
instead of the parasite and that “A. 
symmetricus” does not exist. She also 
noted that Cleidodiscus diversus was 
described from Lepomis cyanellus and 
that its listing for L. symmetricus was 
an error in the 1964 Index-Catalogue, 
Trematoda and Trematode Diseases, 
Part 2, that was perpetuated by Hoff- 
man (1967) and the 1969 Index- 
Catalogue. 

One collection examined during this 
study from Texas (TCWC 3643) col- 
lected 2 May 1970 was heavily infested 
(all 32 specimens in the lot) with a 
monogenetic trematode, presumably 
Anchoradiscus triangularis. No exter- 
nal parasites were observed during the 
present study. 


SUMMARY 


The life-history information on L. 
symmetricus collected in Wolf Lake be- 
tween 2 June 1973 and 27 May 1975 is 
summarized in Table 15. 


Table 15.—Summary of life-history information on Wolf Lake Lepomis symmetricus. 


Characteristics 


Principal habitat 


Age at reaching sexual maturity 
Size at reaching sexual maturity 
Sexual dimorphism 


Number of mature ova in preserved females 
Description of egg 

Spawning period 

Spawning habitat 


Spawning site 


Influence of sex on growth rate 
Density 

Sex ratio among young 
Longevity 

Maximum size 

Principal diet 


Life-History Data 


Shallow, heavily vegetated margins of standing 
water 


1 year 
Females about 34 mm; males about 40 mm 


Adult males are darker on the head and body 
have duskier pelvic fins and longer pelvic fins; 
females tend to have more distinct vertical bars 


219-1,600 

About 0.8 mm in diameter, translucent orange 

From mid-April to early June 

Presumably in shallow water, over soft mud bot- 
tom, near plant material 


Shallow nest depression, about 90-120 mm in di- 
ameter 


Virtually none 

Up to 0.69 sunfish per square meter 
1.5 females : 1 male 

3+ years 

63.0 mm standard length 


Aquatic gastropods, insect immatures, and micro- 
crustaceans 


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actions 30:60-68. 

SmirH, P. W. 1965. A preliminary annotated 
list of the lampreys and fishes of Illinois. 
Illinois Natural History Survey Biological 
Notes 54. 12 p. 

1971. Illinois streams: a classification 

based on their fishes and an analysis of fac- 

tors responsible for disappearance of native 
species. Illinois Natural History Survey Bio- 

logical Notes 76. 14 p. 

. [1973.] A key to the fishes of Illinois. 

Illinois Department of Conservation Fishery 

Bulletin 6. 43 p. 

, and D. W. Brivces. 1960. Ichthyolog- 

ical type specimens extant from the old Illi- 

nois State Laboratory of Natural History. 

Copeia 1960:253-254. 

, A. C. Lopinot, and W. L. PFLIEGER. 
1971. A distributional atlas of upper Missis- 
sippi River fishes. Illinois Natural History 
Survey Biological Notes 73. 20 p. 

Summers, W. A. 1937. A new species of Tetra- 
onchinae from Lepomis symmetricus. Jour- 
nal of Parasitology 23:432-434. 

, and H. J. BENNETT. 1938. A prelimi- 
nary survey of the trematodes from the gills 
of Louisiana fishes. Louisiana Academy of 
Science Proceedings 1:247-248. (Abstract) . 

WALKER, J. M. 1962. Fishes in north Louisi- 
ana. Louisiana Academy of Science Proceed- 
ings 25:35-41. 

1963. Fishes in Choudrant Bayou. 
Louisiana Academy of Science Proceedings 
26:45-48. 

Wess, D. H., and M. E. Sisk. 1975. The fishes 
of west Kentucky. III. The fishes of Bayou 
de Chien. Kentucky Academy of Science 
Transactions 36:63-70. 

WuitakeER, J. O., JR. 1968. Keys to the verte- 
brates of the eastern United States, excluding 
birds. Burgess Publishing Company, Minne- 
apolis. iii + 256 p. 


A 


Adjusted body weight, 438, 452 

Age attained (see longevity) 

Age composition, 455 

Aging method, 438 

Associated species (see species associates) 


Breeding dates (see locations and dates of 
breeding) 
Breeding male, 443, 450-451 


Cc 


Collecting methods, 438 
Coloration (see description) 
Competition, 457-458 
Conservation status, 448-449 


D 


Density per square meter, 454, 455 
Demography, 454-455 
Description 

coloration, 443-445 

general, 443-445 
Development and growth, 453-454 
Diagnosis, 442-443 
Diet 

general, 455-459 

of young, 456-457 

seasonal variation, 456, 458-459 
Distribution, geographic, (iv) 
Drainages, 445-447 


E 
Eggs 
number of, 451-452 


G 


Genital papillae, 445, 450-451 

Geographic distribution (see distribution, 
geographic) 

Growth (see development and growth) 


H 


Habitat, 449-450 
Hatching (see development and growth) 
Hybridization, 459 


Illinois Fish Code, 438 


Interactions with other organisms, 457-460 


INDEX 


Juvenile, 444 


L 


Largest specimen examined, 442, 445 
Lectotype designation, 442 

Life history in Wolf Lake, 449-460 
Locations and dates of breeding, 452-453 
Longevity, 455 


M 


Museum abbreviations, 437 


oO 


Ovarian size and weight variation, 451-452 


P 
Parasitism, 459-460 
Predation, 458-459 
Prejuvenile, 444 

R 


Relationships, 446, 450 
Relative survival, 455 
Relative survival calculation, 439 
Reproductive cycle 
of female, 451 
of male, 450-451 


S 


Sexual differences (see variation, sexual) 
Size distribution, 454 

Spawning, 452-453 

Species associates, 450 

Specimens studied, 446-447 

Study area, 449 

Survival (see relative survival) 
Synonymies and synonyms, 439-440 
Syntopic species of sunfishes, 457-458 


Types, 440, 442 


Variation 
allometric, 445 
geographic, 440-442, 445-446 
meristic, 440-441, 445 
morphometric, 442 
ontogenetic, 444-445 
sexual, 445 


465 


466 ILLINoIs NATURAL History SURVEY BULLETIN Vol. 31, Art. 10 


ELEANORE WILSON, Junior Professional Scientist MELVIN E. ScHWARTZ, Fiscal Officer 

Roser? D. Crompton, Field Assistant DENNIS WALLER, Stockroom Manager 
JAMES W. SEETS, Technical Assistant 

ein Publications and Public Relations 


RoverRtT M, ZEWADSKI, M.S., Technical Editor 
as aes! Divuman, Pri poperty ori and Trust Suan, McC. es Sewn Pechnicd nae, 
parry L. Duzan, Technical Assistants era UAWRENOESS. FarLow, Technica otographer 
ROBERTO. ELLs, ‘Assistant for Operations 2. Tuovp DeMene, Technical Illustrator 
Larry D. Gross, Operations Assistant 2 Ss Tech 1 Lib: 
J. Witiiam Lusk, Mailing and Distribution Services Te echnical Library : 
Jerry McNEAR, Operations Assistant sus DORIS UBLETTE, M.S.L.S., Technical Librarian 


CONSULTANTS AND RESEARCH AFFILIATES: Systematic ENTOMOLOGY, RODERICK R. IRwIn, 
Chicago, Illinois; WILDLIFE RESEARCH, WILLARD D. KiLIMSTRA, Ph.D., Professor ‘of Zoology and Director 
of Cooperative Wildlife Research, Southern Illinois University ; PARASITOLOGY, NorMAN D. LEVINE, Ph.D., 
Professor of Veterinary Parasitology, Veterinary Research and Zoology and Director of the Center for 
Human Ecology, University of Illinois; ENTOMOLOGY, RoBERT L. METCALF, Ph.D., Professor of Biology 
and Research Professor of Entomology, University of Illinois; and GiueertT P. WALDBAUER, Ph.D., Pro- 
fessor of Entomology, University of Illinois; STATISTICS, Horace W. Norton, Ph.D., Professor of Statis- 
tical Design and Analysis, University of Illinois. 


Supporting Services 


Tegal eae 
pe 
rie COR 


Some Publications of the ILLINOIS NATURAL HISTORY SURVEY 


BULLETIN 


Volume 31, Article 3.—Nutritional Responses 
of Pheasants to Corn, with Special Refer- 
ence to High-Lysine Corn. By Ronald F. 
Labisky and William L. Anderson. July, 
1973. 26 p., index. 

Volume 31, Article 4—An Urban Epiphytotic 
of Phloem Necrosis and Dutch Elm Dis- 
ease, 1944-1972. By J. Cedric Carter and 
Lucile Rogers Carter. May, 1974. 31 p., 
index. é 

Volume 31, Article 5.—Larvae of the Seri- 
cothripini (Thysanoptera: Thripidae), with 
Reference to Other Larvae of the Tere- 
brantia, of Illinois. By Thomas C. Vance. 
August, 1974. 64 p., index. 

Volume 31, Article 6.—Root Infection of 
Woody Hosts with Verticillium albo-atrum, 
By Gerald L. Born. August, 1974. 41 p., 
index. 

Volume 31, Article 7—The Mecoptera, or 
Scorpionflies, of Illinois. By Donald W. 
Webb, Norman D. Penny, and John C. 
Marlin. August, 1975. 66 p., index. 

Volume 31, Article 8—An Electrofishing Sur- 
vey of the Illinois River, 1959-1974. By 
Richard E. Sparks and William C. Starrett. 
August, 1975. 64 p., index. 

Volume 31, Article 9.—Pesticides and En- 
vironmental Quality in Illinois. By Robert 
L. Metcalf and James R. Sanborn. August, 
1975. 56 p., index. 


BIOLOGICAL NOTES 


91.—The Distribution of Periodical’ Cicadas 
in Illinois. By Lewis J. Stannard, Jr. 
February, 1975. 12 p. 

92.—The Literature of Arthropods Associated 
with Soybeans. IV. A Bibliography of the 
Velvetbean Caterpillar Anticarsia gemma- 
talis Hubner (Lepidoptera: Noctuidae). 
By B. J. Ford, J. R. Strayer, J. Reid, and 
G. L. Godfrey. February, 1975. 15 p. 

93.—The Life History of the Stripetail 
Darter, Etheostoma kennicotti, in Big 


i 
A 


Creek, Illinois. By Lawrence M. Page. 
February, 1975. 15 p. 

94.—Illinois Pheasants: Their Distribution 
and Abundance, 1958-1973. By Ronald F. 
Labisky. February, 1975. 11 p. 

95.—The Nest Biology of the Bee Andrena 
(Ptilandrena) erigeniae Robertson (Hy-- 
menoptera: Andrenidae). By Lloyd R. 
Davis, Jr., and Wallace E. LaBerge. June, 
1975. 16 p. 

96.—Apparatus and Procedure for Extracting 
Corn Rootworm Eggs from Soil. By John 
T. Shaw, Robert O. Ellis, and _W. H. 
Luckmann. February, 1976. 4 p. 

97.—Environmental Evaluations Using Birds 
and Their Habitats. By Jean W. Graber — 
and Richard R. Graber. May, 1976. 39 p. — 

98.—Effects of Potassium on Adult Asiatic — 
Clams, Corbicula manilensis. By Kevin Bie 
Anderson, Carl M. Thompson, Richard 5-9 
Sparks, and Anthony A. Paparo. July, — 
1976. 7 p. Sem 

99.—The Life History of the Slabrock Darter. x 
Etheostoma smithi, in Ferguson Creek, Ken- 
tucky. By Lawrence M. Page and Brooks 
M. Burr. December, 1976. 12 p. it 

100.—Some Unusual Natural Areas in Illinois. — 
By Robert A. Evers and Lawrence M. Page. — 
June, 1977. 47 p. Fas 

101.—-A Bibliography of the Northern Corn — 
Rootworm and the Western Corn Rootworm: 
An Updating through 1976. By Bonnie J. rp 
Irwin. June, 1977. 8 p. 


CIRCULAR 


51.—Illinois Trees: 
“Care. By J. Cedric Carter. 

123 p. 
52.—Fertilizing and Watering Trees. By Dan 
Neely and E. B. Himelick. December, 1971. 
(Third printing.) 20 p. 
54.—Corn Rootworm Pest Management in 
Canning Sweet Corn. By W. H. Lackenaag : 
. 


poet ra ey buy ae So Rae eg on 


Selection, Planting and 
August, 1966, 


J. T. Shaw, D. E. Kuhlman, R. Randel 


and C. D. LeSar. March, 1975. 10 p. ie i 


List of available publications mailed on request 


No charge is made for publications of the ILLINoIs NaTURAL History SuRvEY. A single 


copy of most publications will be sent free to anyone requesting it until the supply becomes 
low. Costly publications, more than one copy of a publication, and publications in short supply 
are subjects for special correspondence. Such correspondence should identify the writer and 4 Bi 
explain the use to be made of the publication or publications. ca 
en 
Address orders and correspondence to the Chief, 
Illinois Natural History Survey 


Natural Resources Building, Urbana, Illinois 61801 


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UNIVERSITY OF ILLINOIS-URBANA 


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BULLETIN OF THE ILLINOIS STATE LABORATOR 


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