Advances in the Biology and Management of Modern Bed Bugs ebook

Table of Contents

Cover

Title Page

List of Contributors

Foreword

Reference

Acknowledgments

Introduction

References

Part I: Bed Bugs in Society

1 Bed Bugs Through History

1.1 Introduction

1.2 Origins and Spread

1.3 Early Extermination Methods

1.4 Propagation Within Cities (1880s–1950s)

1.5 Determination – and a Silver Bullet

1.6 Past is Present

References

2 Bed Bugs in Popular Culture

2.1 Introduction

2.2 Bed Bugs in Poetry

2.3 Bed Bugs in the Figurative Arts

2.4 Bed Bugs in Theatre

2.5 Bed Bugs in Literature

2.6 Bed Bugs in Music

2.7 Bed Bugs in Television

2.8 Bed Bugs in Linguistics

2.9 Bed Bugs in Erotica

2.10 The Use of Bed Bugs in Popular Culture Through Time

References

Part II: The Global Bed Bug Resurgence

3 The Bed Bug Resurgence in North America

3.1 Introduction

3.2 Bed Bug Resurgence in the United States

3.3 Bed Bug Resurgence in Canada

3.4 Summary

References

4 The Bed Bug Resurgence in Latin America

4.1 Introduction

4.2 Bed Bugs in Brazil

4.3 The Resurgence of Bed Bugs in Brazil

4.4 Elsewhere in Latin America

References

5 The Bed Bug Resurgence in Europe and Russia

5.1 Introduction

5.2 History of Bed Bugs in the Region

5.3 Historical Laws to Control Bed Bugs Pre‐resurgence

5.4 Documented Evidence for the Bed Bug Decline

5.5 Early Evidence for the Resurgence

5.6 The Degree of the Resurgence

5.7 Region‐specific Factors in the Resurgence

5.8 Strategies to Combat the Resurgence

References

6 The Bed Bug Resurgence in Asia

6.1 Introduction

6.2 History of Bed Bugs in Asia

6.3 Laws, Regulations and Policies for Bed Bug Control Prior to the Resurgence

6.4 Modern Resurgence of Bed Bugs in Asia

6.5 Specific Factors Related to the Bed Bug Resurgence in Asia

6.6 Bed Bug Management Strategies in Asia

References

7 The Bed Bug Resurgence in Australia

7.1 Introduction

7.2 History of Bed Bugs in Australia

7.3 Documented Evidence for the Bed Bug Decline

7.4 The Modern Resurgence

7.5 Strategies to Combat the Bed Bug Resurgence

References

8 The Bed Bug Resurgence in Africa

8.1 Introduction

8.2 History of Bed Bugs in Africa

8.3 Laws, Regulations, and Policies for Bed Bug Control Pre‐resurgence

8.4 Documented Evidence for the Bed Bug Decline

8.5 Early Evidence for the Resurgence

8.6 The Extent of the Resurgence

8.7 Region‐ and Country‐specific Reasons for the Resurgence

8.8 Strategies to Combat the Resurgence and Relative Success

References

9 The Bed Bug Resurgence in the Indian Subcontinent

9.1 Introduction

9.2 History of Bed Bugs in the Indian Subcontinent

9.3 Laws to Control Bed Bugs Pre‐resurgence

9.4 Documented Evidence for the Bed Bug Decline

9.5 The Modern Resurgence

9.6 Strategies to Combat the Bed Bug Resurgence

References

10 The Bed Bug Resurgence in the Middle East

10.1 Introduction

10.2 History of Bed Bugs in The Middle East

10.3 Regional Reports of the Bed Bug Resurgence

10.4 Impact of the Bed Bug Resurgence

10.5 Pest Management Professionals Close Encounters with Cimicids

References

Part III: Bed Bug Impacts

11 Dermatology and Immunology

11.1 Introduction

11.2 Bed Bug Saliva

11.3 Cutaneous Reactions

11.4 Dermatological Complications from Bed Bug Bites

11.5 Systemic Reactions

11.6 Immunological and Pathogenic Mechanisms

11.7 Dermatopathology

11.8 Differential Diagnoses of Bed Bug Bites

11.9 Clinical Management

11.10 Conclusion

References

12 Bed Bugs and Infectious Diseases

12.1 Introduction

12.2 Vectors and Transmission Pathways

12.3 Bed Bugs and Infectious Diseases: an Overview

12.4 Why do Bed Bugs not Transmit Infectious Diseases?

12.5 The Future Hunt for Pathogens: A Cautionary Note

12.6 Conclusion

References

13 Mental Health Impacts

13.1 Introduction

13.2 Methods

13.3 Main Findings

13.4 What Can Be Inferred from the Current State of the Literature?

13.5 Limitations and Future Research

13.6 Conclusion

References

14 Miscellaneous Health Impacts

14.1 Introduction

14.2 Respiratory Issues

14.3 Blood Loss

14.4 Sleep Loss

14.5 Chemical Exposure

14.6 Miscellaneous Health Impacts

References

15 Fiscal Impacts

15.1 Introduction

15.2 Types of Cost

15.3 Costs to the Multi‐Unit Housing Industry

15.4 Cost to the Hospitality and Travel Industry

15.5 Cost to the Retail Industry

15.6 Brand Damage in the Housing, Hospitality, and Retail Industries

15.7 Legal Expenses

15.8 Cost to Pest Management Companies

15.9 Bed Bug Management Revenues

15.10 Conclusions

References

Part IV: Bed Bug Biology

16 Bed Bug Biology

16.1 Introduction

16.2 Hematophagy

16.3 Anachoresis

16.4 Flightlessness

16.5 Reproduction

16.6 Egg Laying

16.7 Host‐seeking Behavior

16.8 Harborage Seeking Behavior and Aggregation

16.9 Dispersal

References

17 Chemical Ecology

17.1 Introduction

17.2 Olfaction and Contact Chemoreception

17.3 Pheromones

17.4 Host Seeking

References

18 Population Genetics

18.1 Introduction

18.2 The Evolution of Modern Bed Bugs

18.3 Genetic Variation Within Populations

18.4 Genetic Variation Among Populations

18.5 Mitochondrial Heteroplasmy

18.6 Future Directions in Bed Bug Population Genetics

References

19 Physiology

19.1 Introduction

19.2 Stress Tolerance and Starvation Resistance

19.3 Blood Feeding

19.4 Reproduction and Development

19.5 Summary and Future Directions

References

20 Symbionts

20.1 Introduction

20.2 Identity of Endosymbionts

20.3 Impact of Symbionts on Bed Bug Biology

20.4 Transmission of Symbionts

20.5 Symbionts and Bed Bug Management

References

21 Bed Bug Laboratory Maintenance

21.1 Introduction

21.2 General Colony Maintenance

21.3 Feeding Techniques

21.4 Need for Plasma

21.5 Development of an Artificial Blood Source

References

Part V: Bed Bug Management

22 Bed Bug Industry Standards: Australia

22.1 Introduction

22.2 Why was the Code Required?

22.3 The History and Aims of the Code

22.4 The Key Elements of the Code

22.5 The Benefits of the Code

References

23 Bed Bug Industry Standards: Europe

23.1 Introduction

23.2 Why was the Code Required?

23.3 The History and Aims of the Code

23.4 The Benefits of the Code

References

24 Bed Bug Industry Standards: USA

24.1 Introduction

24.2 History and Development of the NPMA Best Management Practices for Bed Bugs

24.3 Target Audience

24.4 Key Elements of the NPMA Best Management Practices

24.5 Marketing and Adoption of the NPMA Best Management Practices

24.6 Acceptance of the BMP

References

25 A Pest Control Company Perspective

25.1 Introduction

25.2 The Resurgence

25.3 Responsibility and Liability

25.4 Inspection and Control Methods

25.5 Conclusion

References

26 Prevention

26.1 Introduction

26.2 Education

26.3 Monitors

26.4 Mattress Encasements

26.5 Desiccant Dusts

26.6 Heat

26.7 Bed Bug Management Policy

References

27 Detection and Monitoring

27.1 Importance of Detection and Monitoring

27.2 Detection of Bed Bugs

27.3 Field Comparison of Detection Methods

27.4 Bed Bug Inspections

Conflict of Interest Statement

References

28 Non‐chemical Control

28.1 Introduction

28.2 Excluding Bed Bugs

28.3 Physically Removing Bed Bugs

28.4 Creating Adverse Environmental Conditions against Bed Bugs

28.5 Biological Agents Tested Against Bed Bugs

28.6 Other Non‐Chemical Control Methods

28.7 Conclusion

References

29 Insecticide Resistance

29.1 Introduction

29.2 Insecticides and Insecticide Resistance in Bed Bugs

29.3 Metabolic Resistance

29.4 Reduced Penetration Resistance

29.5 Target‐site Resistance

29.6 Evolution of Resistance and Fitness Costs

29.7 Conclusions

References

30 Chemical Control

30.1 Introduction

30.2 Insecticide Classes used Against Bed Bugs

30.3 Insecticide Formulations for Bed Bug Management

30.4 Factors Affecting Insecticide Efficacy

References

31 Limitations of Bed Bug Management Technologies

31.1 Introduction

31.2 Bed Bug Detection

31.3 Bed Bug Control

References

32 Bed Bug Education

32.1 Introduction

32.2 Strategies and Successes in Bed Bug Education

32.3 Educational Programs Focusing on Bed Bugs

32.4 The Media

32.5 The Effect of Social Media on Bed Bug Education

32.6 Identifying the Target Audience

32.7 Effective Adult Education

32.8 Measuring the Impacts of Bed Bug Education

32.9 Conclusion

References

Part VI: Bed Bug Control in Specific Situations

33 Low‐income Housing

33.1 Introduction

33.2 Management of Bed Bugs in Low‐income Housing

33.3 Components of a Successful Building‐ or Complex‐wide IPM Program

33.4 The Future

References

34 Multi‐Unit Housing

34.1 Introduction

34.2 Challenges Unique to the Human Living Environment

34.3 Obstructions to Control Success

34.4 Future Prospects for Success

References

35 Shelters

35.1 Introduction

35.2 Challenges in Shelters

35.3 Obstacles to Successful Control

35.4 Methods of Control in Shelters

35.5 Key Elements to Successful Control

References

36 Hotels

36.1 Introduction

36.2 Bed Bugs in Hotels: The Challenges

36.3 Successful Bed Bug Management in Hotels

References

37 Healthcare Facilities

37.1 Introduction

37.2 The Challenges Bed Bugs Pose to Healthcare Facilities

37.3 The History of Bed Bugs in Healthcare Facilities

37.4 Bed Bugs in Healthcare Facilities with the Modern Resurgence

37.5 Bed Bug Management in Healthcare Facilities

References

38 Aircraft

38.1 Introduction

38.2 Aviation Entomology – a Brief History

38.3 Bed Bug Management on Aircraft: The Challenges

38.4 Bed Bug Management on Aircraft

38.5 Improving the Pest Management Protocol

References

39 Cruise Ships and Trains

39.1 Introduction

39.2 Cruise Ships and Ferries

39.3 Trains

39.4 Conclusion

References

40 Poultry Industry

40.1 History – Cimicids and Poultry

40.2 Mexican Chicken Bug,

Haematosiphon inodorus

40.3 Brazilian Chicken Bug,

Ornithocoris toledoi

40.4 Tropical Bed Bug,

Cimex hemipterus

40.5 The Common Bed Bug,

Cimex lectularius

40.6 Biology and Impact of

Cimex lectularius

on Poultry

40.7 Dispersal

40.8 Bed Bug Control in Poultry Facilities

40.9 Insecticide Assays

References

Part VII: Legal Issues

41 Bed Bugs and the Law in the USA

41.1 Introduction

41.2 Registration of Pesticides

41.3 Legal Requirements Regarding Who Can Apply Pesticides in the USA

41.4 Legal Requirements for PMPs Regarding the Standard of Care

41.5 Public Health Acts Regarding Bed Bugs

41.6 Bed Bug‐related Statutes, Laws, and Ordinances

41.7 Laws Addressing Bed Bug Remediation

41.8 Tenants and Public Housing

41.9 Legal Standing Clients Encountering Bed Bugs in Temporary Occupancies

41.10 Bed Bug Lawsuit Landscape

41.11 Conclusion

References

Court Cases

Restatements of Law

Statutes

42 Bed Bugs and the Law in the United Kingdom

42.1 Introduction

42.2 Training of Pest Management Professionals

42.3 Bed Bug Pesticide Approval

42.4 Tenants, Guests and Bed Bugs

42.5 Local Authority Duties and Powers Regarding Bed Bugs

References

43 Bed Bugs and the Law in Australia

43.1 Introduction

43.2 Registration and Use of Bed Bug Management Products

43.3 Legal Requirements of Pest Management Professionals

43.4 Public Health Laws Regarding Bed Bugs

43.5 Tenancy and Public Housing

43.6 Bed Bug Legal Cases

References

Legal Cases

44 Bed Bugs and the Law in Asia

44.1 Introduction

44.2 Registration and Use of Bed Bug Management Products

44.3 Legal Requirements for Pest Management Professionals

44.4 Bed Bug Legal Cases

44.6 Future

References

45 On Being an Expert Witness

45.1 Introduction

45.2 What is an Expert Witness?

45.3 The Expert’s Role

45.4 Providing Expert Testimony

45.5 Bed Bugs in a Court Case

45.6 Summary

References

Part VIII: Bed Bugs: the Future

46 Bed Bugs: the Future

Summary

References

Index

End User License Agreement

List of Tables

Chapter 06

Table 6.1 The number of bed bug treatments in Singapore from 2006 to 2011.

Table 6.2 Importance and difficulty of control of bed bugs in southeast Asian countries compared to other major urban pests.

Table 6.3 The ranking of bed bug related services offered by pest management professionals in southeast Asia.

Chapter 08

Table 8.1 Summary of bed bug surveys conducted in Africa where bed bug prevalence was specifically expressed as percentage of infested sampling localities.

Table 8.2 Summary of studies conducted on bed bugs in Africa where occurrence is reported on, but not necessarily in terms of prevalence.

Table 8.3 Percentage of enquiries received per category for years indicated, as obtained from a survey completed by 32 pest management companies or individuals in South Africa and Namibia during June 2016.

Table 8.4 Specific bed bug recommendations proposed for African countries.

Chapter 13

Table 13.1 Synthesis of the main findings of the seven studies with original data on mental health and bed bug infestations.

Chapter 18

Table 18.1 Within and among

Cimex lectularius

infestation population genetic summary statistics.

Table 18.2 Among

Cimex lectularius

infestation measures of genetic differentiation.

Chapter 19

Table 19.1 Stress tolerance in the Common bed bug,

Cimex lectularius

.

Chapter 21

Table 21.1 Blood sources used for

in‐vivo

and

in‐vitro

feeding of

C. lectularius

and

C. hemipterus

.

Chapter 29

Table 29.1 Reports of resistance to different insecticide classes in bed bugs (

Cimex

spp.), by year, and in different countries.

Chapter 30

Table 30.1 Insecticide classes and their compounds evaluated for the control of

C. lectularius

and

C. hemipterus

during 1990–2016.

Table 30.2 Insecticide formulations evaluated for the control of C. lectularius and C. hemipterus during 1990–2016.

Chapter 36

Table 36.1 A comparison of known number of rooms infested with bed bugs in three London hotels, pre‐ and post‐inspection, and rate of new infestations.

Table 36.2 Challenges to achieve bed bug eradication in hotels.

List of Illustrations

Chapter 01

Figure 1.1 Bed bugs depicted in

Hortus Sanitatis

(Anonymous, 1536). This was the first ever encyclopedic compilation of natural history, originally published in Germany in 1485.

Figure 1.2 “Summer Amusement, Bugg Hunting” (Cruikshank, 1782). People hand‐picked bed bugs long before the use of insecticides.

Figure 1.3 Title and facing page from John Southhall’s “

A Treatise of Buggs

” (London, 1730).

Figure 1.4 Bed bug inspections used to be important in maintaining a clean and healthful home.

Figure 1.5 Mercury chloride, popularly known as “bed bug poison,” was a common remedy for bed bugs. Many people died from accidentally or intentionally ingesting the poison.

Figure 1.6 Promotions for bed bug products were common and often entertaining. The cartoonist for this 1928 advertisement was Theodore Geisel (Dr Suess).

Figure 1.7 In 1945, suppliers began advertising the availability of DDT for civilian (non‐military) uses, including control of bed bugs.

Figure 1.8 When controlling bed bugs with DDT, treatment of the entire bed was recommended, including the entire mattress.

Figure 1.9 Early patent for a bed bug steamer, published in 1873.

Chapter 02

Figure 2.1 Bed bug postcards through history. (A) Stereogram dated 1897 from Missouri, USA. (B) “A hunting we will go”, 1910 from Maine, USA. (C) Also “A hunting we will go”, postmarked 1907 (clearly the same illustrator as B). (D) “I’ve seen all the big bugs of this place”, postmarked 1907. (E) “A one night stand”, 1908, New York, USA. (F) “It’s the little things that count! Drop me a line”, c1910. (G) “Bed bug Islands (where poets dream), Lake Hopatcong, N.J.”, c1940. From the private collection of David Cain, except (C) from the private collection of Stephen Doggett.

Figure 2.2 More postcards through history. (A) “It’s very lively”, undated. (B) A postcard of an early photograph depicting a couple searching for bed bugs, undated. (C) “Bed‐bugs outwitted!”, c. 1900. (D–I). A series of late‐Victorian postcards, c. 1900; see text for details.

Figure 2.3 Bed bugs as practical jokes across the ages. (A) Imitation Bed Bugs, 1920s. (B) Fake Bed Bugs, 2016. (C) A close up of the “bed bugs”, which are surprisingly similar despite almost a century difference in age, ~5 mm in length. Both originate from the US.

Figure 2.4 Bed bug jewellery. (A&B) A small vintage 14 K charm; folding back the bed base reveals a bed bug (date unknown), approx. 1 cm in length; from the private collection of Stephen Doggett. (C) A modern necklace of a replica bed bug poison bottle, c. 2007;

Chapter 04

Figure 4.1 Overall distribution of responses to resident questionnaires in Brazil. The numbers represent the number of counties represented in the survey. The percentages represent the percentage of counties responding in relation to the total number of counties in each state within the geographical regions of Brazil. Data compiled from September 2012 to March 2016.

Chapter 06

Figure 6.1 The number of bed bug enquiries to the government of Tokyo, 1995–2014

Figure 6.2 The number of bed bug treatments per year reported by two different leading pest control companies in Japan from 2004 to 2014 (Motokazu Hirao, unpublished results).

Chapter 16

Figure 16.1 An infested bed frame showing the distinctive periodic and equidistant distribution of bed bug “clusters”, despite the homogeneity of the refuge

Chapter 17

Figure 17.1 Following a recent blood meal, replete

Cimex lectularius

aggregate in folds of bed linen.

Figure 17.2 Exuviae and feces (dark spots on filter paper) of

Cimex lectularius

, sources of the aggregation pheromone.

Figure 17.3 Types of

Cimex lectularius

pheromones: (a) alarm pheromone produced by nymphs and adults in a state of alarm causing dispersal; (b) aggregation pheromone mediating aggregation of all instar nymphs as well as male and female adults; (c) anti‐mating pheromone released by males mounted by males; and (d) anti‐aphrodisiac pheromone produced by nymphs mounted by males. Notes: (1) some of the same components re‐occur in context‐specific functions; (2) in

Cimex hemipterus

, whole‐body extracts of nymphs [containing the components in (d)] – but not of males or females – repel nymphs as well as male and female adults.

Chapter 18

Figure 18.1 Genetic diversity in

Cimex lectularius

populations: (a) Within and among infestation mean number of alleles per locus; (b) Comparison of heterozygosities between the Common bed bug (

C. lectularius

) and the German cockroach (

B. germanica

). Hashed bar, expected heterozygosity; open bar, observed heterozygosity;

A

within‐infestation,

B

within city,

C

bat‐associated lineage,

D

human‐associated lineage.

E

within USA,

F

within Eurasia.

Figure 18.2

kdr

haplotype distribution in

Cimex lectularius

across the USA. Letter (A–D) refers to haplotype. A: 419 susceptible, 925 susceptible, B: 419 susceptible, 925 resistant, C: 419 resistant, 925 resistant, D: 419 resistant, 925 susceptible. Number below pie is total number of infestations screened. Data combined from Zhu

et al

., (2008); Vargo

et al

., (2011); Holleman and Booth (unpublished results).

Chapter 21

Figure 21.1 Mason jar and components used to house bed bugs. Unassembled and assembled cap with mesh screening, and folded, filter paper harborage.

Figure 21.2 Commercially‐available glass feeders used for feeding bed bugs

in vitro

.

Figure 21.3 Hemotek membrane feeding system for

in‐vitro

feeding of bed bugs: (a) heating unit; (b) Parafilm, left; blood reservoir, center; and O‐ring to attach Parafilm to blood reservoir, right; (c) assembled blood reservoir; (d) reverse side of blood reservoir (after attachment of Parafilm and O‐ring) showing the two fill holes, left and right, and stoppers. Middle opening accepts the screw from heating unit. To fill the unit, after tilting the reservoir about 45°, approximately 3.5 ml blood is introduced through the bottom hole; the unit is then tilted 45° in the opposite direction and the left hole is plugged; after tilting 45° in the original direction, the right hole is plugged, resulting in a filled blood reservoir; (e) filled blood reservoir; (f) blood reservoir screwed into the heating unit; and (g) wooden stand to support the heating units and bed bug colonies (inverted for feeding).

Chapter 25

Figure 25.1 A cluttered apartment, which is infested with bed bugs

Chapter 27

Figure 27.1 Examples of passive and active monitors currently in the marketplace. (a) passive harborage‐style monitor (brand name: PackTite™ Passive Bed Bug Monitor, Bed Bugs Limited, London, UK, image courtesy David Cain, Bed Bugs Limited, London); (b) passive pitfall‐style monitor designed to be placed under legs of beds (brand name: BlackOut® BedBug Central, Lawrenceville, NJ); (c) passive pitfall‐style monitors designed to be placed under legs of beds (brand name: ClimbUp® Insect Interceptor, Susan McKnight, Inc., Memphis, TN); (d) passive monitor designed to be placed near or away from beds (brand name: SenSci Volcano®, BedBug Central, Lawrenceville, NJ); (e) active monitors baited with carbon dioxide, heat, and chemical lure (brand name: NightWatch® Bed Bug Monitor, BioSensory Inc., Putnam, CT); (f) active monitor baited with chemical lure (brand name: SenSci Volcano® with SenSci Activ® Bed Bug Lure, BedBug Central, Lawrenceville, NJ, image courtesy of Susannah Reese, Cornell University, NY).

Chapter 28

Figure 28.1 Bed bugs and fecal spotting inside a wall void in multi‐family housing. (S.A. Kells, University of Minnesota).

Figure 28.2 Use of an infrared thermometer to verify surface temperatures immediately after steam application. (S.A. Kells, University of Minnesota).

Figure 28.3 Shipping container fitted with a heater for larger containerized heat treatments.

Figure 28.4 Stacking furniture in a portable container for heat treatment. (S.A. Kells, University of Minnesota).

Figure 28.5 Whole‐room heat treatment with electric furnaces.

Chapter 36

Figure 36.1 The broad processes required to manage bed bugs in hotels (David Cain, unpublished results and after various sources, see also www.bed‐bugs.co.uk/hotels).

Guide

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Caption: “War on the bed bug”. Postcard c. 1916. Clearly humanity’s dislike of the bed bug has not changed through the years!

Advances in the Biology and Management of Modern Bed Bugs

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Cover Design: WileyCover Image: © John‐Reynolds/Gettyimages

Dedicated to the works of luminaries such as J.R. Busvine, A.A. Girault, A. Hase, C.G. Johnson, H. Kemper, K. Mellanby, K. Newberry, N. Omori, and R.L. Usinger. We are merely dwarfs standing of the shoulders of giants.

List of Contributors

Mohammad AkhoundiService de Parasitologie‐Mycologie, Centre Hospitalier Universitaire de Nice – Hôpital de l'Archet, Nice, France

Ondřej BalvínCzech University of Life Sciences, Prague, Czech Republic

Paul J. BelloPJB Pest Management Consulting, Alpharetta, Georgia, USA

Joshua B. BenoitDepartment of Biological Sciences, University of Cincinnati, Cincinnati, Ohio, USA

Clive BoaseThe Pest Management Consultancy, Haverhill, United Kingdom

Warren BoothDepartment of Biological Science, The University of Tulsa, Tulsa, Oklahoma, USA

David CainBed Bugs Limited, London, United Kingdom

Toni CainsSydney Local Health District Public Health Unit, Camperdown, Australia

Ana Eugênia de Carvalho CamposUnidade Laboratorial de Referênciaem Pragas Urbanas, Instituto Biológico, São Paulo, Brasil

Richard CooperCooper Pest Solutions and Bedbug Central, Lawrenceville, New Jersey, USA

Dionne CraffordClinvet International (Pty) Ltd, Bloemfontein, South Africa

Pascal DelaunayService de Parasitologie‐Mycologie, Centre Hospitalier Universitaire de Nice – Hôpital de l'Archet, Nice, France

Stephen L. DoggettDepartment of Medical Entomology, NSW Health Pathology, Westmead Hospital, Westmead, Australia

Odelon Del Mundo ReyesEcovar, Dubai, United Arab Emirates

Sophie E.F. EvisonDepartment of Animal and Plant Sciences, University of Sheffield, Sheffield, United Kingdom

Mark F. FeldlauferUSDA‐ARS, Invasive Insect Biocontrol and Behavior Laboratory, Beltsville, Maryland, USA

Pablo Fernandez‐PeñasDermatology, Sydney Medical School, The University of Sydney, Westmead, Australia

Josephus FourieClinvet International (Pty) Ltd, Bloemfontein, South Africa

Jim FredericksNational Pest Management Association (USA), Fairfax, Virginia, USA

Jody Gangloff‐KaufmannNew York State Integrated Pest Management Program, Cornell University, Geneva, New York, USA

Mark GoodmanVarment Guard Environmental Services Inc., Columbus, Ohio, USA

Gerhard GriesDepartment of Biological Sciences, Simon Fraser University, Burnaby, Canada

Geneviève HamelinPublic Health Department of Montreal, Montreal, Canada

Harold J. HarlanUS Army Medical Entomologist (retired), USA

William T. HentleyDepartment of Animal and Plant Sciences, University of Sheffield, Sheffield, United Kingdom

Motokazu HiraoHirao Biological Institute, Shizuoka, Japan

Andrew Ho‐OharaEden Law Corporation, Singapore

Shelley Ji Eun HwangDepartment of Dermatology, Westmead Hospital, Westmead, Australia

Garry JonesConsultant Pest Management Professional, Buff Point, Australia

Adam JusonMerlin Environmental Solutions Ltd, Carshalton, United Kingdom

Catherine JusonMerlin Environmental Solutions Ltd, Carshalton, United Kingdom

João Justi (Jr.)Unidade Laboratorial de Referênciaem Pragas Urbanas, Instituto Biológico, São Paulo, Brasil

David KaiserPublic Health Department of Montreal, Montreal, Canada

Stephen A. KellsDepartment of Entomology, University of Minnesota, St. Paul, Minnesota, USA

Márcio R. LageFaculdade de SaúdePública, Universidade de São Paulo, São Paulo, Brasil

Chow‐Yang LeeSchool of Biological Sciences, Universiti Sains Malaysia, Penang, Malaysia

David G. LillyEcolab Pest Elimination, Macquarie Park, Australia

Jeffrey LipmanLipman Law Firm, West Des Moines, Iowa, USA

Dini M. MillerDepartment of Entomology, Virginia Tech, Blacksburg, Virginia, USA

Richard NaylorThe Bed Bug Foundation, Chepstow, United Kingdom

Linda‐Lou O’ConnorAirRx Antimicrobial Sciences Inc., Baltimore, Maryland, USA

Joelle F. OlsonPest Elimination Division, Ecolab Research Center, Eagan, Minnesota, USA

Roberto M. PereiraEntomology and Nematology Department, University of Florida, Gainesville, Florida, USA

Stéphane PerronPublic Health Department of Montreal, Montreal, Canada

Michael F. PotterDepartment of Entomology, University of Kentucky, Lexington, Kentucky, USA

Anil S. RaoPest Control (India) Private Ltd, Mumbai, India

Joshua A. RaoPest Control (India) Private Ltd, Mumbai, India

Alvaro RomeroDepartment of Entomology, Plant Pathology and Weed Science, New Mexico State University, Las Cruces, New Mexico, USA

Coby SchalDepartment of Entomology, North Carolina State University, Raleigh, North Carolina, USA

Mike T. Siva‐JothyDepartment of Animal and Plant Sciences, University of Sheffield, Sheffield, United Kingdom

Allen SzalanskiDepartment of Entomology, University of Arkansas, Fayetteville, Arkansas, USA

Allison Taisey AllenNational Pest Management Association, Fairfax, Virginia, USA

Kevin R. UlrichUSDA‐ARS, Invasive Insect Biocontrol and Behavior Laboratory, Beltsville, Maryland, USA

Karen VailDepartment of Entomology and Plant Pathology, The University of Tennessee, Knoxville, Tennessee, USA

Edward L. VargoDepartment of Entomology, Texas A&M University, College Station, Texas, USA

Changlu WangDepartment of Entomology, Rutgers University, New Brunswick, New Jersey, USA

Mark W. WilliamsEcolab Pest Elimination, Northwich, United Kingdom

Molly S. WilsonDepartment of Biochemistry, Virginia Tech, Blacksburg, Virginia, USA

Rebecca WilsonDepartment of Animal and Plant Sciences, University of Sheffield, Sheffield, United Kingdom

Yijuan XuDepartment of Entomology, South China Agricultural University, Guangzhou, China.

Foreword

Harold J. Harlan

Bed bugs were a serious pest of human communities long before recorded history. However, for millenia, information about bed bugs, and advice on how to deal with them, has been in evidence across many lineages of cultural lore. This fact is most obvious when you consider that bed bugs are known to have at least 71 common names originating from 36 different languages across the world (Usinger, 1966). The Monograph of Cimicidae (Usinger 1966) was the first comprehensive publication on bed bugs that combined extensive worldwide information from historic, cultural, scientific, pest management, and general public resources.

Advances in the Biology and Management of Modern Bed Bugs (ABMMBB) updates and expands much of Usinger’s (1966) information, with emphasis on the worldwide resurgence of both the Common bed bug, Cimex lectularius L., and the Tropical bed bug, Cimex hemipterus (F.). ABMMBB incorporates extensive new information from a wide range of basic and applied research, as well as the recently observed medical, legal and regulatory impacts of bed bugs.

Today there are many new, extremely precise technologies, and laboratory tools that could not have been imagined in 1966. Recent innovations, especially in molecular biology and genetics, offer a fascinating range of potential applications. In addition, we have new systems for information gathering, processing, and sharing with international colleagues. These technologies have opened up whole new fields of scientific investigation over the past 20 years.

Stephen L. Doggett, Dini M. Miller, and Chow‐Yang Lee have done a terrific job of assembling and coordinating more than 60 contributing authors who are highly experienced and widely recognized as experts in their topic areas. The contributing authors offer new information on basic science and advice on using applied management strategies and bed bug bioassay techniques. The authors also present cutting‐edge information on the major impacts that bed bugs have had on the medical, legal, housing, and hotel industries across the world, as well as their impacts on public health.

ABMMBB is the most comprehensive compilation yet produced about these bugs that includes historic, technical, and practical information. It will certainly be the most thorough single reference on bed bugs for many decades to come. I believe that ABMMBB will be an essential reference for anyone who is engaged in managing bed bugs, be it in an academic, basic or applied scientific setting, or in a public outreach or pest management role, worldwide. I am very honored, and humbled, to have been asked to provide this foreword. I can hardly wait to buy my own personal copy.

Reference

Usinger, R.L. (1966)

Monograph of Cimicidae (Hemiptera – Heteroptera)

, Entomological Society of America, College Park.

Acknowledgments

The editors, Stephen L. Doggett, Dini M. Miller and Chow‐Yang Lee, would like to express gratitude to all the authors and reviewers that contributed towards the development of Advances in the Biology and Management of Modern Bed Bugs (ABMMBB). We would especially like to acknowledge Harold Harlan for writing the foreword, and for contributing so much to so many. The editors also wish to acknowledge Wiley‐Blackwell and staff (notably Sonali Melwani, Ramya Raghaven, Bella Talbot, David McDade, Emma Strickland, Gunalan Lakshmipathy, and Ward Cooper) for their assistance, advice, editorial, and production efforts. A special thank you to Junichiro Katayama from Semco Co. Ltd. in Japan, who kindly hosted the editors of ABMMBB in mid‐October 2016 so that we could finalize much of the text (thank you also for all the wonderful meals!).

The following are acknowledged for particular chapters:

Chapter 3: The Global Resurgence in North America

The author wishes to acknowledge the assistance of Dennis Monk (Director, Bedbug Solutions) and Sean Rollo (Regional Manager, Orkin Canada) for providing information on the current bed bug situation in Canada.

Chapter 5: The Global Resurgence in Europe and Russia

The authors are especially grateful to the contribution of Václav Rupeš, former head (retired) of the Department of Desinfection, Desinsection and Deratization, State Health Institute, Prague.

Chapter 8: The Global Resurgence in Africa

The authors would like to thank Henda Pretorius and Reinier Zwiegers [Clintest (Pty) Ltd], and Ashleigh Caddick (PES Africa) for their assistance in running the online survey cited in the chapter. The authors would also like to thank Carmen Neethling [Clinvet (Pty) Ltd] for assistance with literature searches.

Chapter 9: The Global Resurgence in India and the Subcontinent

The editors greatly appreciate the assistance of K.P. Jayanth (PCI India) in the development of this chapter.

Chapter 17: Chemical Ecology of Bed Bugs

The author wishes to thank John Borden for comments, Sharon Oliver for word processing and comments, Michael Hrabar for photographs, and Stephen DeMuth for graphical illustration. The Chapter is dedicated to Regine Gries who was the lead investigator in the research resulting in the identification of the bed bug aggregation pheromone and who in the process endured >180 000 bed bug bites for the maintenance of a thriving bed bug laboratory colony.

Chapter 18: Population Genetics

The published and unpublished population genetics studies mentioned in the chapter would not have been possible without the generous cooperation of many colleagues, too many to mention individually, who provided bed bug samples. Ron Harrison is especially acknowledged who facilitated the collections of samples from Orkin and Orkin International, and Rick Santangelo who managed the collections.

Chapter 19: Bed Bug Physiology

The author wishes to acknowledge that funding while developing this chapter was provided by the University of Cincinnati.

Chapter 21: Bed Bug Laboratory Maintenance

The authors wish to thank; A. Aak, J.F. Anderson, O. Balvin, B. Campbell, S.L. Doggett, C.Y. Lee, D.G. Lilly, J. Olson, R. Naylor, A. Romero, K. Reinhardt, A. Vander Pan, and C. Wang, for completing the rearing survey. Those individuals are acknowledged that contribute to the rearing efforts in all of the laboratories currently maintaining bed bugs. Members of the Armed Forces Pest Management Board (Silver Spring, Maryland USA) are also thanked, along with the personnel at the Walter Reed National Military Medical Center (Bethesda, Maryland USA) for their assistance in obtaining blood products for the senior author.

Chapter 41: Bed Bugs and the Law in the USA

The authors wish to express gratitude to Kolby Warren, Drake University Law School, Des Moines, Iowa, USA, who assisted in the production of the chapter.

Chapter 43: Bed Bugs and the Law in Australia

The authors wish to acknowledge the following individuals for providing important information pertaining to environmental health laws pertaining to bed bugs, from the different states; Keith Rogers (Australian Capital Territory, Health), Rebecca Feldman (Department of Health & Human Services, Victoria), Michaela Hobby (Department of Health, South Australia), Nicola Slavin (Department of Health, Northern Territory), Rebecca Richardson (Department of Health, Queensland), and Mike Lindsay and Donald Howell (Department of Health, Western Australia). Keith Farrow (Rapids Solutions) generously provided information pertaining to the licensing of professional pest managers. Nicholas Cowdery AM QC (University of Sydney, Australia) kindly reviewed the chapter for legal accuracy.

Chapter 44: Bed Bugs and the Law in Asia

The authors wish to thank the following individuals for providing important information pertaining pesticide registration and the licensing of pest management professionals in their respective countries: Erh‐Lieh Hsu (National Taiwan University), Pascal Cai (Chinese Pest Control Association, China), Suchart Leelayouthyotin (King Service Center, Bangkok, Thailand), Sulaeman Yusuf (Indonesia Institute of Science), and Motokazu Hirao (Japan Pest Control Association).

The following individuals have generously supplied images for use in ABMMBB or critically reviewed manuscripts:

Zachary Adelman (Texas A&M University, College Station, Texas, USA)

Joshua Benoit (University of Cincinnati, Cincinnati, Ohio, USA)

Clive Boase (the Pest Management Consultancy, United Kingdom)

Kaci Buhl (National Pesticide Information Center, Oregon State University, Oregon, USA)

David Cain (Bed Bugs Limited, United Kingdom)

Richard Cooper (Cooper Pest Solutions and Bedbug Central, Lawrenceville, New Jersey, USA)

Richard deShazo, (University of Mississippi Medical Center, Mississippi, USA)

Keith Farrow (Rapid Training, Australia)

Mark Feldlaufer (USDA‐ARS, Invasive Insect Biocontrol and Behavior Laboratory, Maryland, USA)

Toby Fountain (Uppsala University, Sweden)

Adam Juson (Merlin Environmental Solutions Ltd, United Kingdom)

David Lilly (Ecolab Pest Elimination, Macquarie Park, Australia)

Oliver Madge (Bed Bug Foundation, United Kingdom)

Frank Meek (Orkin LLC, USA)

Mike Merchant (Texas A&M, AgriLife Extension Service, Texas, USA)

Kosta Y. Mumcuoglu (Hebrew University, Israel)

Richard Naylor (Bed Bug Foundation, United Kingdom)

Faith Oi (Entomology and Nematology Department, University of Florida, Florida, USA)

Christopher Orton (PiMACs, Australia)

Lawrence Pinto (Pinto & Associates, Mechanicsville, Maryland, USA)

Alvaro Romero (New Mexico State University, New Mexico, USA)

Bob Rosenberg (National Pest Management Association, Virginia, USA)

Veera Singham (Centre for Chemical Biology, Universiti Sains Malaysia, Malaysia)

Coby Schal (North Carolina State University, North Carolina, USA)

Allison Taisey Allen (National Pest Management Association, Virginia, USA)

Changlu Wang (Department of Entomology, Rutgers School of Environmental and Biological Sciences, New Jersey, USA)

Jeff White (Bed Bug Central, New Jersey, USA)

Introduction

Stephen L. Doggett, Dini M. Miller and Chow-Yang Lee

“…misery acquaints a man with strange bedfellows…”

William Shakespeare, The Tempest, 1610

“…intellects vast and cool and unsympathetic, regarded this earth with envious eyes, and slowly and surely drew their plans against us…”

H.G. Wells, The War of the Worlds, 1898

The quotation above from one of William Shakespeare’s greatest works could easily read as an allegory for the personal suffering one experiences with a bed bug infestation. However, Shakespeare’s play was written in 1610, only a short time after the UK produced their first reliable bed bug record in 1583 (Mouffet, 1634). Thus it was quite possible that Shakespeare himself never acquainted himself with the misery of bed bugs. Yet it was not long before the insect became so common in the country that companies appeared which specialized in bed bug extermination, such as the famous Tiffin & Son, founded in 1690 (Potter, 2011).

In fact, bed bugs have a long history of inflicting their misery upon humanity. The remains of bed bugs have been found in Egyptian settlements dating back some 3565 years (Paragiotakopulu and Buckland, 1999). With the discovery in 1939 that dichloro‐diphenyl trichloroethane (better known as DDT) had a powerful insecticidal action, suddenly the world had the magical solution that could rid humanity of bed bugs forever. Subsequently DDT (and other organochlorines), and the organophosphates where widely employed to control bed bugs, and infestations became rare in the developed world for many years after World War II (Usinger, 1966). Yet forever was not to be. The late 1990s saw a worldwide re‐emergence of both the Tropical bed bug, Cimex hemipterus (F.), and the Common bed bug, Cimex lectularius L. Not unlike the Martians in H.G. Wells’ classic novel, suddenly nowhere on earth was exempt from bed bugs.

In the early days of the modern resurgence, infestations mainly occurred in the hospitality sector and bed bugs were more limited to premises with high guest turnover. Then people started to take the insect with them wherever they went and, in the process, spread bed bugs into the wider community. Infestations began to appear in such diverse locations as in private homes, on public transport, within the retail sector, in cinemas, at the office, in schools and universities, and even in healthcare facilities. Thus wherever a person went, they could be potentially exposed to bed bugs and take them elsewhere. The greatest concern however, has been the proliferation of bed bugs amongst the socially disadvantaged, a group that often does not have the economic resources to pay for control. As a result, infestations can go uncontrolled and spread throughout a building complex. With a lack of public and government support to ensure that infestations are successfully eradicated, it is unfortunate that such groups have become bed bug reservoirs for the wider society. Thus support should be provided for bed bug eradication programs in low income housing…even if it is only to selfishly protect ourselves from future infestations.

There have been a number of reasons postulated for the modern bed bug resurgence (Pinto et al., 2007; Doggett et al., 2012). This includes globalization and the ease with which people move around the world, thereby enabling the spread of modern bed bug strains. Changes in pest management have meant that hotel rooms are no longer routinely treated for pests. The insecticides available today do not have the same residual life as the organochlorines and are simply not as efficacious. Poor knowledge of bed bugs, in particular how to control modern strains, has clearly been a major factor in the degree of the resurgence, as treatment failures often result in the infestation spreading to other apartments in multiple occupancy dwellings. This particular phenomenon has been aided by the ever increasing concentration of people into high density living due to the uncontrolled growth in the world’s population. These days it is much easier for bed bugs to spread from an infestation to invade other premises. A variety of other reasons have been given for the resurgence, but it appears that the key contributing factor to the modern bed bug resurgence is the development of insecticide resistance within the insect.

In many ways, the bed bugs that our grandparents experienced are very different to those that we are exposed to now. The contemporary bugs have developed multiple defences against the insecticides we use against them. They have thicker skins, which slows down the penetration of insecticides into the body. They produce a range of complex enzymes that can break down insecticides, and they possess mutations that prevent the insecticides from acting at the target sites. Thus the modern bed bug is truly the superbug of the 21st century. The challenge for modern scientists is to find ways to circumvent this range of extraordinary adaptations.

One aspect that has been the subject of debate is the geographical origin of the modern (and resistant) bed bug. Some US based researchers have suggested that resistant strains of C. lectularius arrived from Eastern Europe, although somewhat amusingly, locals in Eastern Europe have the opposing view of blaming American tourists (Borel, 2015). In spite of the contrary opinions, neither belief can explain the origin and simultaneous rise of C. hemipterus. This species has a more tropical distribution, has never been reported from Eastern Europe, and only recently reappeared in the USA (Campbell et al., 2016). Arguably a more simple answer to the origin of resistance in both species is that resistant strains originated from a region where the two species are sympatric and where insecticide selection pressures were equally applied. This would help to explain the synchronous revival of the two species. The most logical locations would be from areas of Africa where infestations of C. hemipterus and C. lectularius coexisted even when infestations elsewhere in the world had become uncommon. In KwaZulu, South Africa, during the late 1980s it was observed that bed bug infestations (of both species) were more common in human dwellings that were sprayed annually with DDT to combat malaria than unsprayed dwellings (Newberry et al., 1987, 1990; Newberry, 1991). The increase in nuisance biting from bed bugs meant that householders often refused chemical treatments despite the potential risk of death from malaria as they believed that spraying may have contributed to greater bed bug numbers! Even some 20 years earlier, a report from the early 1970s noted an increase in bed bug infestations occurred in spite of regular treatments with the organochlorines (Rafatjah, 1971). In both cases, the development of insecticide resistance was the suspected cause of these increases. In fact, by the early 1970s, insecticide resistance had been long known in South African bed bug populations.

The first report of resistance to DDT in C. lectularius was from Hawaii in 1947. This was only three years after the pesticide was first employed (Johnson and Hill, 1948). Within a relatively short time thereafter, reports of resistance to both bed bug species had become widespread (Busvine, 1957). Thus it appears that resistance evolved rapidly, but this may not have been the case. Natural pyrethrins were used for bed bug control after the mid‐1800s, and resistance to this class of insecticides confers cross resistance to the organochlorines. Perhaps some degree of resistance in bed bugs had developed long before the late 1940s but was simply not identified. The organochlorines being more efficacious than pyrethrins may have helped to rapidly eliminate the non‐resistant and less resistant insects. Furthermore, presumably the bed bugs that disappeared in developed nations post World War II were all susceptible strains (or had low‐level resistance). Yet in pockets of the world, high levels of resistance had evolved (and presumably continued to evolve). These “superbugs” just required the means of escape in order to spread elsewhere. The means were provided by humans, with our modern tendency to move about the world. Widespread global travel is a relatively recent behavior in human history, yet rapidly growing. The World Bank estimated that in 1970 there were some 310 million people movements globally, and by 2015 this number had risen to 3.4 billion (World Bank, 2016). Thus insecticide‐resistant bed bugs had ample means to spread from their original source to a new location. The constant rise in human population combined with the increased movement of people across the globe, the recent changes in pesticides and pest management practices mentioned above, and the presence of resistant bed bugs in certain parts of the world, makes it seem that, perhaps in hindsight, the global bed bug resurgence was inevitable and should have been anticipated.

As a consequence of the modern resurgence, there has been a renewed interest in bed bug research. In recent years, a plethora of publications relating to bed bugs have appeared. For example, a search on Pubmed (www.ncbi.nlm.nih.gov/pubmed/) using the term “Cimex”, reveals 259 peer reviewed publications between 1912 and 1999. That number rose to 492 between 2000 to the end of 2016. If industry publications and newspaper reports were included, the increase in bed bug publications would be even more dramatic. Interestingly, the number of scientific publications from 2010 onwards have plateaued at around 50 per year, which probably reflects how minimally funded bed bug research is today. While Usinger’s (1966) seminal manuscript, the Monograph of Cimicidae (Hemiptera – Heteroptera), continues to be the key reference for taxonomy of the Cimicidae, other areas of research are now much further advanced. Thus there is a need for the distillation of all the contemporary information into a modern academic text and hence the birth of Advances in the Biology and Management of Modern Bed Bugs (ABMMBB).

ABMMBB is a synthesis of bed bug information from the past to the present. It aims to serve as a reference book for academic researchers and students alike. It is a valuable text for those in the hospitality sector and accommodation managers, who are tasked with the job of minimizing the risk of bed bugs in their facility, or who organize the eradication of active infestations. With the growth in bed bug litigation, both the litigant and defendant legal teams will find ABMMBB an essential source of contemporary information. Finally, ABMMBB provides up‐to‐date information for the pest management professional on bed bug biology and management. In recent years, most bed bug research has focused on C. lectularius, as this species has impacted the more economically advantaged nations of Europe and North America. However, ABMMBB is aiming to be global in context, and where possible, both bed bug species are discussed and key differences highlighted. With the aim of having an international appeal, ABMMBB has over 60 contributing authors, spanning some 14 nations.

There are seven main parts within ABMMBB. These cover Bed Bugs in Society, the Global Resurgence, Bed Bug Impacts, Biology, Management, Control in Specific Situations, and Legal Issues. Finally, ABMMBB ends with a discussion on the future of bed bugs in society and research needs.

The first part, Bed Bugs in Society, contains two chapters. Chapter 1, “Bed Bugs Through History”, reviews the early methods of extermination. By examining past control methodologies when synthetic insecticides had yet to be discovered, perhaps insights can be gained in how to successfully eradicate insecticide‐resistant strains now; thus the importance of including this work in ABMMBB. The chapter also covers the origins and spread of bed bugs throughout the world. While the bed bug spread was documented historically to a reasonable extent, few bothered to determine which species was involved. It can be surmised (based on past records and current distributions) that it was mainly C. lectularius that was introduced and established in Europe, North America, and initially into Australia. However, few records exist regarding the global spread of C. hemipterus. Thus it is necessary to review the early taxonomic descriptions for this species, as these texts contain information on the site of collection (reviewed in Usinger, 1966). Fabricius first described C. hemipterus, which was captured from South America houses in 1803 (Fabricius, 1803), while other early records include “Ile Bourbon” (1852) [now known as Réunion], “Ost‐Indien” (1861) [East India], Colombia (1854), and Sokotra (1899) [also spelled Socrota, which is part of Yemen]. These references indicate that C. hemipterus had spread around the world by the mid‐19th century as had C. lectularius.

Chapter 2, “Bed Bugs in Popular Culture”, highlights the intimate relationship that humans had with bed bugs throughout history via the depiction of the insect in various forms of media. Bed bugs have appeared over the years in poetry, art, the theatre, literature, music, and more recently, in television.

Part 2, on the global bed bug resurgence, contains contributions from all the major regions across the world, including chapters from the Americas, Europe and Russia, Asia, Australia, Africa, the Indian subcontinent, and the Middle East. Each chapter discusses the history of bed bugs in their respective region, the resurgence, and strategies employed to combat the return of the bed bug.

Bed Bug Impacts (Part 3) reviews the dermatological, mental health, and miscellaneous heath impacts associated with bed bugs. The potential for bed bugs to transmit infectious diseases is also considered. It is difficult to argue the fact that the health‐related impacts of bed bugs are relatively minor compared with insects, such as mosquitoes, that are known to transmit vector‐borne diseases. However, the fiscal impacts of bed bugs are significant and the resurgence has probably cost the world economy billions of dollars. The monetary effects of bed bugs are many and varied, and detailed in the final chapter of this section.

The basis of integrated pest management (IPM) is a comprehensive understanding of the biology of the pest as it relates to effective control strategies, hence the need to include such issues within ABMMBB. Topics covered within Part 4 include host‐seeking and blood‐feeding behaviors, harbourage selection and aggregation, dispersal, chemical ecology, population genetics, physiology, symbionts, and laboratory maintenance. The bizarre reproductive behaviour of bed bugs known as “traumatic insemination” is also discussed.

Part 5 focuses on bed bug management. The first three chapters of this section review the industry standards on bed bug control that have been developed in recent years in Australia, Europe, and the USA. These standards are followed by a chapter on how a large multinational pest management firm approaches bed bug management to ensure consistency of treatments and a positive outcome for the clients. The following chapters focus on prevention (in terms of minimizing the risk of bed bug establishment and spread); detection and monitoring; non‐chemical management; insecticide resistance; and chemical control. In recent years with the modern bed bug resurgence, many technologies have appeared on the market, but only few are truly efficacious. Thus Chapter 31 reviews the inherent limitations in bed bug management technology. The bed bug management coverage finishes with a focus on education. This involves sending the correct message to the community on how to minimize the risk of bed bugs, and how an infestation should be properly managed.

Part 6, on bed bug control in specific situations, reviews the experiences of industry leaders who are responsible for bed bug management in particular environments. These include low‐income housing, multi‐unit housing, shelters, hotels, healthcare facilities, aircraft, trains and cruise ships, and within the poultry industry. In many cases, little has been published in these areas, so the authors often had to recount their own personal experiences and the challenges involved in achieving a successful outcome.

In recent years, bed bugs have been the cause of legal action, especially in the USA, where some cases involve settlements of several million dollars. Thus Part 7 deals with bed bugs and the law, covering legal aspects from the USA, the UK, Asia, and Australia. There is also a chapter on the challenges of being an expert witness involved in bed bug litigation.

The final chapter of ABMMBB undertakes some crystal‐ball gazing to imagine what the future will look like in terms of bed bug and human interaction, and reviews the strategies and research required to reverse the resurgence. Perhaps one of the greatest challenges ahead is to make control affordable for all people. However, only considerable technological and methodological advancements will make this happen. Ultimately the real question is, can we defeat bed bugs again, or are bed bugs set to plague human society forever? Only time will tell.

References