Medical and veterinary entomology

591 Pages·2002·48.821 MB·English

by  Gary R Mullen, Lance A Durden

SROTUBIRTNOC
Numbers in parentheses indicate the pages on which the authors" John E. Lloyd (349), Department of Entomology, University
contributions appear. of Wyoming, Laramie, Wyoming 82071, ASU
John .W McCreadie (185), Department of University Biology,
Peter H. Adler (185), Department of Entomology, Clemson of South Alabama, Mobile, Alabama 36688, ASU
,ytisrevinU Clemson, South Carolina 29634-0365, USA
Roger D. Moon (279), Department of Entomology, University
Roger D. Akre (383), Deceased, formerly Department of En- of Minnesota, .tS Paul, Minnesota 55108, USA
tomology, Washington State University, Pullman, Washing-
Gary IL Mullen (1,163,317, 425,427, 449), Department of
ton 99164-6382, ASU
Entomology and Plant Pathology, Auburn University,
Richard J. Brenner (29), USDA-ARS, Office of Technology Auburn, Alabama 36849, ASU
,refsnarT Maryland Beltsville, 20705; formerly USDA-ARS,
Bradley A. Mullens (263), Department of Entomology, Uni-
Center Agricultural Medical, for and Veterinary Entomology,
versity of California California, Riverside, 92521, ASU
,ellivseniaG Florida 32604, ASU
William L. Nicholson (517), Viral and Rickettsial Zoonoses
E. Paul Catts (317), Deceased, formerly Department of Ento-
Branch, Centers for Disease Control & Prevention, Atlanta,
mology, Washington State Pullman, University, Washington
Georgia 30333, ASU
99164-6382, ASU
Barry M. OConnor (449), University of Michigan, Museum
Lance A. Durden ( ,1 45, 103), Georgia Southern University,
of Zoology, University of Michigan, Ann Arbor, Michigan
Institute of Arthropod and Parasitology, Statesboro, Georgia
48109, ASU
30460, ASU
Hal D. Reed (383), Biology Department, Oral Roberts Uni-
Woodbridge A. Foster (203), Department of Entomology,
,ytisrev Tulsa, Oldahoma 74171, ASU
The Ohio State University, Columbus, Ohio 43210-1220,
ASU William K. Reisen (15), Arbovirus Field Station, University of
California, ,dleifsrekaB California 93312, USA
Reid R. Gerhardt (127), Department of Entomology
and Plant Pathology, University of Tennessee, Knoxville, Louis C. Rutledge (147), Mill ,yellaV California 94941-3420,
Tennessee 37901-1071, ASU ASU
Raj K. Gupta (147), Headquarters, USAMRC, Fort Detrick, Daniel E. Sonenshine (517), Department of Biological -icS
Frederick, Maryland 21702-5012, ASU ences, Old Dominion University, Norfolk, Virginia 23529-
0266, USA
Robert D. Hall (127), Department of Entomology, University
Scott A. Stockwell (411), U. .S Army, 25th Medical Detach-
of Missouri, Columbia, Missouri 65211, ASU
ment (Entomology), Fort Hood, Texas 76544, ASU
William L. Krinsky (67, 87, 303), Department Epidemiology
Robert Traub (103), Deceased, Bethesda, formerly Maryland,
and Public Health, Yale Medical School, New Haven,
Connecticut 06520-8034, ASU ASU
Edward D. Walker (203), Department of Entomology, Michi-
Robert .S Lane (517), Division of Insect Biology, University
gan State East Lansing, University, Michigan 48824, USA
of California, ,yelekreB California 94720, ASU
ECAFERP
This book si the result of the recognized need for an that the discussions of Public Health Importance and
up-to-date, entomologically oriented textbook for teach- Veterinary Importance are addressed separately. This
ing courses in medical and veterinary entomology at the should maximize the flexibility in using this sa a textbook
college and university level. It was initiated in 1993 by for courses in either medical or veterinary entomology, or
Section D (Medical and Veterinary Entomology) of the for courses in which the two disciplines are combined.
Entomological Society of America, following surveys that Literature citations generally have been kept to a mini-
were conducted to identify the scope and format of such mum throughout the text to make the book sa read-
a book that would best meet the needs for classroom able and student-friendly sa possible. The authors were
instruction. The clear sentiment was to produce a com- asked to follow a guideline of providing 45-50 ref-
prehensive textbook covering both medical and veteri- erences for each chapter, selecting only those that are
nary entomology, recognizing the close relationship particularly pertinent or serve sa recommended follow-
between these two disciplines. The individuals polled up sources for additional information relating to the
indicated a strong preference for organizing the chap- chapter subject. Exceptions were made for some of the
ters taxonomically according to the insect and related- longer chapters (e.g., Mosquitoes, Mites, and Ticks) in
arthropod groups involved, similar to that followed in which the number of references and suggested read-
.W B. Herms's original Medical and Veterinary Entomol- ings significantly exceeds the guideline, reflecting the
ogy, published in 1915. That classic work, and the seven breadth and importance of these particular arthropod
editions that followed, served sa the standard textbook groups.
for medical and veterinary entomology students for more In addition to students in the traditional sense, this
than sixty years, until its last printing in 1979. It si hoped book si targeted at a much broader audience, including:
that this book will help to fill the void that has persisted (1) entomologists in general; (2) specialists in other disci-
these past two decades in promoting the teaching of med- plines who have an interest in arthropods of medical
ical and veterinary entomology sa an important part of or veterinary importance; (3) physicians, nurses, health
college and university curricula. officials, and others in the medical field who encounter
With the exception of the first two chapters (Introduc- insects, spiders, scorpions, ticks, and other arthropods in
tion and Epidemiology of Vector-Borne Diseases), the the course of their professional activities; (4) personnel
chapters are similarly organized with the following ma- in a variety of public health- and community-related
jor subheadings: Taxonomy, Morphology, Life History, programs (e.g., mosquito control, tick control) and pro-
Behavior and Ecology, Public Health Importance, Vet- fessional pest control operators who wish to have an
erinary Importance, Prevention and Control, and Refer- informative, readable reference source on their shelf;
ences and Further Reading. This format should be helpful (5) military personnel who have responsibility for dealing
to the reader in locating specific information and to with a diversity of entomologically related problems;
teachers who wish to assign only select portions of a chap- (6) and veterinarians, wildlife personnel, zoological park
ter for their students to read. It si also for this reason officials, and other individuals in animal health-related
IIX 00 Preface
fieldswho invariably encounter insects and related arthro- (moths and butterflies). These two groups are not the first
pods in their respective lines of work. that come to mind when one thinks of medical and vet-
Traditionally, medical and veterinary entomology has erinary entomology and typically are treated only briefly,
included not only insects but also certain arachnid groups if at all, in most medical-veterinary books. However, as
that can present problems for humans and other animals. the space devoted to these groups indicates, the beetles
This book follows that tradition. However, unlike most and moths and the problems they cause deserve more at-
other books in this subject area, a greater amount of at- tention than they are generally accorded.
tention is given to these eight-legged relatives of insects We wish to acknowledge the loss of three contributors
in this work. Fully one-quarter of the text is devoted to this book who passed away after submitting, or con-
to arachnids, with separate chapters on scorpions, solpu- tributing to, the initial and one or more revised drafts
gids, spiders, mites, and ticks. In fact, the chapters on of their chapters. They are Roger D. Akre and E. Paul
mites and ticks (note: ticks are simply a subgroup of Catts of the Department of Entomology, Washington
mites) represent the first and third largest chapters State University, and the late Robert Traub of Bethesda,
in the book, respectively, reflecting their diversity and Maryland. In recognition of their contributions, the re-
medical-veterinary importance. Mosquitoes are the sub- spective chapters appropriately bear their names as au-
ject of the second largest chapter, as the most important thors, albeit posthumously. This book is dedicated to the
group of insects from a medical and public health memory of these distinguished entomologists.
perspective.
In providing appropriate balance to the insect groups
discussed, the reader may be surprised to see separate Gary R. Mullen
chapters on the Coleoptera (beetles) and Lepidoptera Lance A. Durden
STNEMGDELWONKCA
Many people have helped in various ways with the prepa- eW are grateful to the following persons for providing
ration of this textbook and we are grateful to lla of them. original photographs, slides, and other illustrations
Foremost, we must thank lla of the chapter authors who reproduced in this book, with the respective figure
wrote their assigned chapters and patiently accommo- numbers indicated in parentheses: Dr. .W .V Adams, .rJ
dated our editorial requests, comments, guidelines, and (Louisiana State University, Baton Rouge; 13.11), Dr.
also responded to comments furnished by outside review- Peter H. Adler (Clemson University, South Carolina;
ers. Obviously, without the commitment and dedication 11.1, 11.5), the late Dr. Roger D. Akre (Washington
of the chapter authors, this project could not have come State University, Pullman; 19.9, 19.14, 19.15, 19.24);
to fruition. Dr. Hans B~inziger (Chaing Mai University, Thailand;
Rebecca L. Nirns (Auburn University, Alabama and 18.18-18.23), Dr. Yehuda Braverman (Kimron
Georgia Southern University, Statesboro) saw contribut- Veterinary Institute, Israel; 10.19), Dr. Alberto .B Broce
ing art editor and expertly captured the majority of the (Kansas State University, Manhattan; 14.7), Dr. Corrie
figures sa digital images and then digitally labeled and Brown (Foreign Animal Disease Diagnostic Laboratory,
improved many of them; she also prepared original fig- Plum Island, New York and University of Georgia,
ure 23.2. Margo .A Duncan (Gainesville, Florida) saw Athens; 10.16), Dr. Jerry .F Butler (University of
commissioned to prepare original or composite pen and Florida, Gainesville; 18.13, 19.13, 23.25, 23.40, 23.43),
ink drawings for the following figures: 2.1, 2.3, 4.6, 4.8, Dr. James D. Castner (University of Florida, Gainesville;
7.3, 18.1A, and 18.4. eW also wish to thank the follow- 3.3-3.11), Dr. Ronald D. Cave (Escuela Agricola
ing individuals who contributed original artwork in the Panamericana, Zamorano, Honduras; 16.25), Valerie .J
form of line drawings, with the respective figure numbers Cervenka, (University of Minnesota, .tS Paul; 14.22),
indicated in parentheses following their institutional -fa Dr. Jack Kelly Clark (University of California, Davis;
filiations: the late Dr. .E Paul Catts (Washington State 24.16), Dr. George H. D'Andrea (Alabama State Vet-
University, Pullman; 16.1-16.9, 16.11, 16.13-16.15, erinary Diagnostic Laboratory, Auburn; 10.14), Dr.
16.17, 16.20, 16.23, 16.26, 16.31, 16.32, 16.35, 16.36, Debbie .R Folkerts (Auburn University, Alabama; 21.1),
16.39), Dr. Woodbridge .A Foster (The Ohio State Uni- Dr. Woodbridge .A Foster (The Ohio State University,
versity, Columbus; 12.14), Susan .J M. Hope (Mebane, Columbus; 12.14, 12.17, 12.19, 12.25, 12.27, 12.31),
North Carolina; 13.1), Takumasa Kondo (Auburn Uni- Dr. Ellis .G Greiner (University of Florida, Gainesville;
Alabama; versity, 8.9), Dr. William L. Krinsky elaY( Med- 11.8), Dr. Martin Hall (The Natural History Museum,
ical School, New Haven, Connecticut; 5.6), Dr. Roger London; 16.37), Dr. Robert .G Hancock (Cumberland
D. Moon (University of Minnesota, .tS Paul; 14.2), Dr. College, Williamsburg, Kentucky; 12.23), Dr. Carl
Bradley .A Mullens (University of California, Riverside; C. Hansen (National Museum of Natural History,
13.3), Dr. Blair .J Sampson (USDA-ARS, Small Fruit Washington, D.C.; 18.12), Dr. Elton .J Hansens
Research Center, Poplarville, Mississippi; 19.5), and Dr. (Asheville, North Carolina; 4.12, 4.13, 7.14, 13.6,
Lawrence .W Zettler (Illinois College, Jacksonville; 11.2). 14.12, 19.12, 19.30, 23.11), Dr. Charles M. Hendrix,
ooo
XUl
VIX stnemgdelwonkcA
(Auburn University, Alabama; 23.48, 23.60), Dr. Nancy (U.S. Livestock Insects Laboratory, USDA/ARS,
C. Hinkle (University of California, Riverside and Univer- Kerrville, Texas), Dr. Peter Landolt (USDA-ARS,
sity of GeorgAitah,e ns; 4.14, 7.7-7.10), Paul M. Horton Wapato, Washington), Dr. Donald G. Manley (Pee
(Clemson University, South Carolina; 19.28), Lacy L. Dee Research and Education Center, Florence, South
Hyche (Auburn University, Alabama; 18.15), Dr. Robert Carolina), Dr. Leonard E. Munstermann (Yale Uni-
J. Keiffer (University of California, Hopland; 24.15), versity, New Haven, Connecticut), Dr. Edward .T
Takumasa Kondo (Auburn University, Alabama; 19.23), Schmidtmann (USDA-ARS, Arthropod-Borne Animal
Peter .J Landolt (USDA-ARS, Wapato, Washington), Dr. Diseases Laboratory, Laramie, Wyoming) and Dr. Bryce
Robert .S Lane (University of California, Berkeley; 24.13, Walton (Gettysburg, Pennsylvania). Additional figure and
24.19), Lloyd L. Lauerman (Alabama State Veterinary illustration credits are provided throughout the book.
Diagnostic Laboratory, Auburn; 10.8-10.11), Dr. We are especially grateful to the following individuals
Kriangkrai Lerthudsnee (Mahidol University, Bangkok, who served sa external reviewers or provided advice for
Thailand; 12.20), Dr. John E. Lloyd (University of one or more of the chapters or chapter sections: Dr.
Wyoming, Laramie; 4.15, 17.2, 17.6, 17.10-17.12), John .R Anderson (Connecticut Agricultural Experiment
Dr. John M. MacDonald (Auburn University, Alabama; Station, New Haven), Renee Anderson (Auburn Univer-
23.45, 23.46, 23.54), the late Dr. Sturgis McKeever sity, Alabama), Dr. Arthur G. Appel (Auburn University,
(Georgia Southern University, Statesboro; 13.4, 13.13, Alabama), the late Dr. Ross H. Arnett (formerly of
16.18, 18.6, 18.8, 18.11, 18.14, 18.17, 22.13, 22.17), Gainesville, Florida), Matt Aubuchon (Auburn Uni-
Dr. Hendrik .J Meyer (North Dakota State University, versity, Alabama), Dr. Hans B~inziger (Chaing Mai
Fargo; 14.18, 14.21, 14.23), Dr. Roger D. Moon University, Thailand), Dr. Alberto .B Broce sasnaCI(
(University of Minnesota, St. Paul; 14.19), Dr. Gary .R State University, Manhattan), Dr. Richard L. Brown
Mullen (Auburn University, Alabama; 5.14, 6.9, 10.4, (Mississippi State University, Mississippi), Dr. John
10.5, 10.12, 14.20, 16.19, 18.7, 19.8, 19.19-19.22, Burger (University of New Hampshire, Durham), Dr.
19.29, 22.12, 22.19), Dr. Bradley .A Mullens (Univer- John .B Campbell (University of Nebraska, North Platte),
sity of California, Riverside; 10.2, 13.7), Dr. Harold Dr. Bruce M. Christensen (University of Wisconsin,
D. Newson (Michigan State University, East Lansing; Madison), Dr. G. .B Edwards (Florida State Collec-
12.34, 12.36), Dr. Yoshiro Ohara (Tohoku University tion of Arthropods, Gainesville), Dr. Richard G. Endris
School of Medicine, Sendai, Japan; 13.10), Dr. Jonathan (Schering Plough, Union, New Jersey), Dr. Marc Epstein
D. Patterson (Michigan State University, East Lansing; (U.S. National Museum of Natural History, Smithsonian
12.35), the late Dr. L. L. Pechuman (Cornell University, Institution, Washington D.C.), Dr. Richard D. Fell
Ithaca, New York; 13.5), Dr. Hal C. Reed (Oral Roberts (Virginia Polytechnic and State University, Blacksburg),
University, Tulsa, Oklahoma; 19.18, 19.31), Ross Ritter Dr. Howard Frank (University of Florida, Gainesville),
(Potter ,yellaV California; 24.14), Mary Elizabeth Rogers Dr. Reid .R Gerhardt (University of Tennessee,
(Waukegan, Illinois; 10.17), Dr. William .S Romoser Knoxville), Dr. M. Lee Golf (University of Hawaii
(The Ohio State University, Columbus; 12.20), Dr. at Manoa, and Chaminade University, Honolulu), Dr.
Justin O. Schmidt (Carl Hayden Bee Research Labora- Ellis C. Greiner (University of Florida, Gainesville),
tory, Tucson, Arizona; 19.4, 19.27), Dr. Philip .J Scholl Dr. William L. Grogan (Salisbury State University,
(USDA-ARS, Lincoln, Nebraska; 16.34), Dr. Scott .A Maryland), Dr. Duane .J Gubler (Centers for Disease
Stockwell (US Army, 25 ht Medical Detachment, Fort Control and Prevention, Fort Collins, Colorado), the
Hood, Texas; 20.1, 20.7, 20.8), Dr. Daniel .R Suiter late Dr. William L. Hardy (formerly at the University
(University of Georgia, Griffin; 3.2), Dr. Robert .B Tesh of California, Berkeley), Dr. Cluff E. Hopla (University
(University of Texax Medical Branch, Galveston; 5.4), .P of Oklahoma, Norman), Dr. James .A House (USDA-
Kirk Visscher (University of California, Riverside; 10.6), APHIS, Foreign Animal Disease Diagnostic Laboratory,
Dr. Laurel L. Waiters (Lieen-Follican Research, Bishop, Plum Island, New York), Dr. Lawrence .J Hribar
California; 9.7), and Dr. Ralph E. Williams (Purdue (Florida Keys Mosquito Control District, Marathon),
University, West Lafayette, Indiana; 14.17). Dr. Gregg .J Hunt (USDA/ARS, Arthropod-Borne
In addition, we express our appreciation to these indi- Animal Diseases Laboratory, Laramie, Wyoming), Lacy
viduals who assisted in the preparation of figures or in L. Hyche (Auburn University, Alabama), Dr. James
other ways provided illustrations from which the figures E. Keirans (Georgia Southern University, Statesboro),
that appear in this book were selected: Dr. Anne-Marie Dr. Robert .R Killick-Kendrick (Imperial College at
Callcott (USDA-APHIS, Gulfport, Mississippi), Dr. Silwood Park, Ascot, England), Dr. Katherine M. Kocan
Randy Davidson (Southeastern Cooperative Wildlife Dis- (Oldahoma State University, Stillwater), Dr. Elliot .S
ease Study, University of Georgia, Athens), Dr. Harold Krafsur (Iowa State University, Ames), Dr. Daniel C.
.J Harlon (Crownsville, Maryland), Dr. Sidney Kunz Kurtak (Chewelah, Washington), Dr. Phillip G. Lawyer
stnemgdelwonkcA VX
(Uniformed Services University of Health Sciences, of Natural History, New York, New York), Dr. Scott
Bethesda, Maryland), Dr. Robert E. Lewis (Ames, Iowa), Stewart (NIH/NAIAD, Hamilton, Montana), Dr.
Dr. John E. Lloyd (University of Wyoming, Laramie), Christine .A Sundermann (Auburn University, Alabama),
Dr. Timothy .J Lysyk (Agriculture Canada, Lethbridge, Dr. Sabina .F Swift (University of Hawaii at Manoa,
Alberta), Dr. Adrian G. Marshall (University of Aberdeen, Honolulu), Dr. Pete D. Teel (Texas A & M University,
Scotland), Dr. Robert Mincldey (University of Utah, College Station), Dr. Robert .B Tesh (University of Texas
Salt Lake City), Dr. Roger D. Moon (University of Medical Branch, Galveston), Dr. Stephen Torr (Natural
Minnesota, St. Paul), Dr. Charles D. Morris (Winter Resources Institute, Chatham Maritime, England), the
Haven, Florida), Dr. Bradley .A Mullens (University of late Dr. Robert Traub (formerly of Bethesda, Maryland),
California, Riverside), Dr. Roy .A Norton (State Uni- Dr. Michael .J Turell (U.S. Army Medical Research
versity of New York, Syracuse), Dr. Richard .S Patterson Institute of Infectious Diseases, Fort Detrick, Frederick,
(Gainesville, Florida), Michelle Perdue (Auburn Univer- Maryland), Dr. William .J Turner (Washington State
,ytis Alabama), Dr. Peter .V Perldns (Gainesville, Florida), University, Pullman), Dr. .S Bradleigh Vinson (Texas A
Dr. Robert .V Peterson (Monte L. Bean Life Sciences & M University, College Station), Dr. Laurel L. Waiters
Museum, Brigham Young University, Provo, Utah), Dr. (Lieen-Follican Research, Bishop, California), Amelia
Oscar J. Pung (Georgia Southern University, Statesboro), Williams (Auburn University, Alabama), Dr. Mark L.
Dr. Sarah E. Randolph (University of Oxford, England), Wilson (University of Michigan, Ann Arbor), Dr. James
Dr. William C. Reeves, .rS (University of California, C. Wright (Auburn University, Alabama), Dr. Russell E.
Berkeley), Dr. William .K Reisen (Arbovirus Field Sta- Wright (Oklahoma State University, Stillwater), and Dr.
tion, University of California, Bakersfield), Dr. Richard David .G Young (Gainesville, Florida).
G. Robbins (Defense Pest Management Information We are indebted to the following individuals for
Analysis Center, Forest Glen Section, Walter Reed their assistance in conducting library searches or other-
Army Institute of Research, Washington D.C.), Dr. wise providing the authors with relevant literature:
The Honorable Dame Miriam Rothschild (Ashton Dr. Harold .J Harlan (Crownsville, Maryland),
Wold, England), Dr. Michael .J Rust (University of Elizabeth Mason (North Dakota State University,
California, Riverside), Dr. Raymond E. Ryckman Fargo), Dr. Richard G. Robbins (Defense Pest Manage-
(Redlands, California), Dr. Justin O. Schmidt (Carl ment Information Analysis Center, Forest Glen Section,
Hayden Bee Research Center, Tucson, Arizona), Dr. Walter Reed Army Institute of Research, Washington
Edward .T Schmidtmann (USDA-ARS, Arthropod- D.C.), the late Dr. Robert Traub (formerly of Bethesda,
Borne Animal Diseases Laboratory, Laramie, Wyoming), Maryland) and Anne .R Viera (University of Tennessee,
Dr. Christopher .J Schofield (London School of Tropical I~oxville).
Medicine & Hygiene, United Kingdom), Dr. Philip J. We also would like to thank the staff with whom
Scholl (USDA-ARS, Lincoln, Nebraska), Dr. Craig .T we have worked at Academic Press/Elsevier Science in
Sheppard (University of Georgia, Coastal Experiment San Diego, especially Dr. Charles .R Crumly, Monique
Station, Tifton), Dr. Louis N. Sorkin (American Museum Larson, Molly Wofford, and Christine Vogelei.
I
N O I T C U D O R T N I
LANCE A. DURDEN AND GARY R. MULLEN
GENERAL ENTOMOLOGY 2 the study of insects and insect-related problems that
MEDICAL-VETERINARY ENTOMOLOGY affect domestic animals, particularly livestock and com-
LITERATURE 2 panion animals (dogs, cats, horses, caged birds, etc.).
In addition, veterinary entomology includes insect-
HISTORY OF MEDICAL-VETERINARY
associated problems affecting captive animals in zoologi-
ENTOMOLOGY 2
cal parks and wildlife in general. Medical-veterinary -otne
IDENTIFICATION AND SCITAMETSYS OF ygolom combines these two disciplines.
ARTHROPODS OF MEDICAL-VETERINARY Traditionally the fields of medical and veterinary en-
IMPORTANCE 3 tomology have included health-related problems involv-
SEPYT OF PROBLEMS ing arachnids (particularly mites, ticks, spiders, and scor-
pions). This broad approach encompassing insects and
CAUSED YB ARTHROPODS 4
arachnids si followed in this text. Alternatively, the study
ARTHROPOD-BORNE SESAESID 5
of health-related problems involving arachnids si called
FOOD CONTAMINANTS 6 medical-veterinary ygolonhcara or, if just mites and ticks
FEAR OF ARTHROPODS 6 are considered, medical-veterinary .ygoloraca
DELUSORY SISOTISARAP 6 Historically, both medical and veterinary entomology
TOXINS AND VENOMS 6 have played major roles in the development of human
civilization and animal husbandry. Outbreaks of insect-
HOST DEFENSES 7
borne diseases of humans have profoundly influenced hu-
FORENSIC ENTOMOLOGY 7
man history; these include such diseases sa yellow fever,
MINOR ARTHROPOD PROBLEMS plague, louse-borne typhus, malaria, African trypanoso-
OF MEDICAL-VETERINARY INTEREST 8 miasis, Chagas disease, and lymphatic filariasis. Like-
REFERENCES AND FURTHER READING 8 wise, livestock scourges such sa bovine babesiosis, bovine
theileriosis, scabies, pediculosis, and botfly infestations,
lla of which are caused or transmitted by arthropods,
have greatly influenced animal production and husbandry
Medical entomology si the study of insects, insect-borne practices. Arthropod-related disorders continue to cause
diseases, and other associated problems that affect hu- significant health problems to humans, domestic animals,
mans and public health. Veterinary ygolomotne si similarly and wildlife. At the same time, new strains of known
MEDICAL AND VETERINARY ENTOMOLOGY
Copyright 2002, Science Elsevier (USA). All rights reserved.
ecnaL A. Durden and Gary R. Mullen
pathogens, sa well sa previously unrecognized disease 1999). Publications that devote substantial sections to
agents transmitted by arthropods, are causing newly rec- arthropods associated with wildlife and the pathogens
ognized diseases (e.g., Lyme disease and human granulo- they transmit include Davis and Anderson (1971),
cytic ehrlichiosis) and the resurgence of diseases that had Davidson et al. (1981), Fowler (1986) and Davidson and
been suppressed for many years (e.g., malaria). In fact, Nettles (1997).
emerging and resurging arthropod-borne sesaesid are rec- Several journals and periodicals are devoted primar-
ognized sa a growing health concern by public health and yli to medical and/or veterinary entomology. These in-
veterinary officials (Wilson and Spielman 1994, Walker clude the Journal of Medical Entomology, published by
et al. 1996, Gubler 1998, Winch 1998, Gratz 1999). the Entomological Society of America (Lanham, MD);
Medical and Veterinary Entomology, published by the
Royal Entomological Society of London (UK); Jour-
nal of Vector Ecology, published by the Society of Vector
LARENEG YGOLOMOTNE
Ecologists (Corona, CA); Vector Borne and Zoonotic Dis-
,sesae published by Mary Ann Liebert, Inc., Larchmont,
Basic concepts of entomology, such sa morphology, tax- New York; and Review of Medical and Veterinary En-
onomy and systematics, developmental biology, and ecol- tomology, published by BAC International (Wallingford,
ogy, provide important background information for med- UK). Journals specializing in parasitology, tropical
ical and veterinary entomologists. General entomology medicine, or wildlife diseases that also include articles
books which the reader will find helpful in this regard in- on medical-veterinary entomology include Parasitology,
clude Borror et al. (1989), Gullan and Cranston (1994), published by the British Society for Parasitology; Jour-
Oillot (1995), Elzinga (1997), Chapman (1998), and nal of Parasitology, published by the American Society
Romoser and Stoffolano (1998). References that provide of Parasitologists (Lawrence, KS); Parasite-Journal de la
a more taxonomic or biodiversity-oriented approach to Societe Franfaise de Parasitologie, published by PRIN-
general entomology include works by Arnett (1993), CEPS Editions (Paris, France); Advances in Disease Vector
Richards and Davies (1994), Bosik (1997), and Daly Research, published by Springer-Verlag (New York); Bul-
et al. (1998). General insect morphology si detailed in letin of the World Health Organization, published by the
Snodgrass (1993), whereas a useful glossary of general World Health Organization (Geneva, Switzerland); Jour-
entomology si Torre-Bueno (1962). Texts on urban nal of Wildlife ,sesaesiD published by the Wildlife Disease
entomology, the study of insect pests in houses, build- Association (Lawrence, KS); Emerging Infectious ,sesaesiD
ings, and urban areas, which also has relevance to published by the Centers for Disease Control and Pre-
medical-veterinary entomology, have been prepared vention (Atlanta, GA); the American Journal of Tropical
by Ebeling (1975), Hickin (1985), MaUis (1997), and Medicine and Hygiene, published by the American Society
Robinson (1996). General texts on acarology include of Tropical Medicine and Hygiene (Northbrook, IL); and
works by Krantz (1978), Woolley (1987), Evans (1992) Memorias Do Instituto Oswaldo Cruz; published by the
and Walter and Proctor (1999). Instituto Oswaldo Cruz (Rio de Janeiro, Brazil). Various
Internet Web sites pertaining to medical-veterinary ento-
mology can also be accessed for useful information.
YRANIRETEV-LACIDEM
YGOLOMOTNE ERUTARETIL
YROTSIH FO -LACIDEM
YRANIRETEV YGOLOMOTNE
Textbooks or monographs pertaining to medical ento-
mology, veterinary entomology, or the combined disci-
pline of medical-veterinary entomology are listed under Problems caused by biting and annoying arthropods and
these headings at the end of this chapter. Most of these the pathogens they transmit have been the subject of
publications emphasize arthropod morphology, biology, writers since antiquity (Service 1978). Homer (mid-8th
systematics, and relationships, disease whereas some of the century BC), Aristophanes (ca. 448-380 BC), Aristotle
more recent texts, such sa Beaty and Marquardt (1996) (384-322 BC), Plautus (ca. 254-184 BC), Columella
and Crampton et al. (1997), emphasize molecular stcepsa (5 BC to AD 65), and Pliny (AD 23-79) lla wrote
of medical-veterinary entomology. Other works are help- about the nuisance caused by flies, mosquitoes, lice,
luf regarding common names of arthropods of medical- and/or bedbugs. However, the study of modern medical-
veterinary importance (Pittaway 1992), surveillance tech- veterinary entomology si usually recognized sa beginning
niques (Bram 1978), control measures (Drummond in the late 19th century, when blood-sucking arthropods
et al. 1988), or ectoparasites (Andrews 1977, Marshall were first proven to be vectors of human and animal
1981, miK 1985, Uilenberg 1994, Barnard and Durden pathogens.
Introduction 3
Englishman Patrick Manson ( 1844-1922 ) was the first In 1909, Frenchman Charles Nicolle (1866-1936),
to demonstrate pathogen transmission by a blood-feeding working in Tunis, showed that human body lice are
arthropod. Working in China in 1877, he showed that the vectors of Rickettsia prowazekii, the agent of louse-borne
mosquito Culex pipiensfatigans is a vector of Wuchere- (epidemic) typhus.
ria bancrofti, the causative agent of Bancroftian filaria- These important discoveries, as well as others of his-
sis. Following this landmark discovery, the role of various torical relevance to medical-veterinary entomology, are
blood-feeding arthropods in transmitting pathogens was discussed in more detail in the references listed at the end
recognized in relatively rapid succession. of this chapter. Because of the chronology of many ma-
In 1891, Americans Theobald Smith (1859-1934) jor discoveries relevant to this topic in the 50-year period
and F. L. Kilbourne (1858-1936) implicated the cattle starting in 1877, this time has been called the "golden
tick, Boophilus annulatus, as a vector of Babesia bigem- age of medical-veterinary entomology" (Philip and
ina, the causative agent of Texas cattle fever (bovine Rozeboom 1973).
babesiosis/piroplasmosis). This paved the way for a
highly successful .B annulatus-eradication program in
the United States directed by the US Department of
Agriculture. The eradication of this tick resulted in the IDENTIFICATION AND .....
projected goal: the elimination of indigenous cases of SYSTEMATICS OF ARTHROPODS OF
Texas cattle fever throughout the southern United States.
MEDICAL-VETERINARY IMPORTANCE
In 1898, Englishman Sir Ronald Ross (1857-1932),
working in India, demonstrated the role of mosquitoes
as vectors of avian malarial parasites from diseased to Table I provides a list of the eight orders of insects and
healthy sparrows. Also in 1898, the cyclical development four orders of arachnids that are of particular interest to
of malarial parasites in anopheline mosquitoes was des- medical-veterinary entomologists. Accurate identification
cribed by Italian Giovani Grassi (1854-1925). In the of these arthropods is an important first step in deter-
same year, Frenchman Paul Louis Simond (1858-1947), mining the types of problems they can cause and, subse-
working in Pakistan (then part of India), showed that fleas quently, in implementing control programs.
are vectors of the bacterium that causes plague. Although taxonomy and identification are discussed
In 1848, American physician Josiah Nott (1804- in more detail with respect to arthropod groups treated
1873) of Mobile, AL, had published circumstantial ev- in the chapters that follow, some publications provide
idence that led him to believe that mosquitoes were in- a broader perspective on the classification, taxonomy,
volved in the transmission of yellow fever virus to humans.
In 1881, Cuban-born Scottish physician Carlos Finlay
(1833-1915) presented persuasive evidence for his the- ELBAT I
ory that what we know today as the mosquito Aedes ae- Principle Orders of Insects and Arachnids of
gypti was the vector of this virus. However, it was not un- Medical-Veterinary Interest
til 1900 that American Walter Reed (1851-1902) led the Order Common names
US Yellow Fever Commission at Havanna, Cuba, which
proved A. aegypti to be the principal vector of yellow fever ssalC atcesnI
virus. Order airattalB sehcaorkcoC
Order arctparihthP eciL
In 1903, Englishman David Bruce (1855-1931)
Order aretpimeH True :sgub ,sgubdeb gnissdl ,sgub
demonstrated the ability of the tsetse fly Glossina palpalis
nissassa sgub
to transmit, during blood-feeding, the trypanosomes that Order aretpocloC selteeB
cause African trypanosomiasis. Order aretpanohpiS saelF
Other important discoveries continued well into the Order aretpiD :seilF ,seotiuqsom kcalb ,seilf ,snru-ees-on
esroh ,seilf deer ,seilf sand ,seilf estest
20th century. In 1906, American Howard Taylor Ricketts
,seilf house ,seilf elbats ,seilf horn ,seilf
(1871-1910) proved that the Rocky Mountain wood bot ,seilf blow ,seilf hself ,seilf esuol
tick, Dermacentor andersoni, is a vector of Rickettsia ,seilf ,sdek .cte
rickettsii, the causative agent of Rocky Mountain spotted Order aretpodipeL shtoM seilfrettu bdna
fever. In 1907, F. .P Mackie (1875-1944) showed that Order aretponemyH ,spsaW horncts, tevlev ants, ants, seeb
human body lice are vectors of Borrelia recurrentis, the ssalC adinhcarA
Order adinoiprocS snoiprocS
spirochete that causes louse-borne (epidemic) relapsing
Order adiguploS ,sdiguploS nus ,sredips lemac ,sredips
fever. In 1908, Brazilian Carlos Chagas (1879-1934)
lerrab sredips
demonstrated transmission of the agent that causes Order iracA ,setiM skcit
American trypanosomiasis, later named Chagas disease in Order caenarA sredipS
his honor, by the cone-nose bug Panstrongylus megistus.