Bed Bug Population Dynamics: Genetic Variation , Geographic Diversity, etc.


Akhoundi, M., D. Sereno, R. Durand, A. Mirzaei, C. Bruel, P. Delaunay, P. Marty, and A. Izri. 2020. Bed Bugs (Hemiptera, Cimicidae): Overview of Classification, Evolution and Dispersion. Int. J. Environ. Res. Public Health 17(12):4576. doi: 10.3390/ijerph17124576

The bed bugs (Cimex lectularius and C. hemipterus) have undergone a significant resurgence worldwide since the 1990s. A compilation of findings from a database, including 2650 scientific publications from seven major medical databases, allowed us to document main evolutionary events, from fossil evidence, dating from 11,000 years ago, until the present that has led to the current worldwide expansion of Cimicid species. We present the hypotheses on the possible dispersion pathways of bed bugs in light of the major historical and evolutionary events. A detailed classification of the Cimicidae family and finally, an illustrative map displaying the current distribution of known Cimex species in each geographical ecozone of Asia, Europe, Africa, the Americas, and Australia are presented.”



Masran, S., S. N. Ain, A. Majid, and A. Hafiz2018. Isolation and characterization of novel polymorphic microsatellite markers for Cimex hemipterus F. (Hemiptera: Cimicidae). Journal of Medical Entomology. 55(3): 760-765. doi: 10.1093/jme/tjy008

“Due to the growing public health and tourism awareness, Cimex hemipterus Fabricius (Hemiptera: Cimicidae) has gained a great interest in increasing reported infestation cases in tropical regions of the world, including Malaysia. Since the information on the molecular ecology and population biology of this species are tremendously lacking, the isolation and development of molecular markers can be used to determine its genetic structure. In this study, novel microsatellite primers isolated from enriched genomic libraries of C. hemipterus were developed using 454 Roche shotgun sequencing. Seven validated polymorphic microsatellite primers were consistently amplified and characterized from 70 tropical bed bugs collected from seven locations throughout Malaysia. The number of alleles per locus identified ranged from 6 to 14. Comparison of loci for overall and between population were done with mean observed and expected heterozygosity were determined at 0.320 and 0.814, 0.320 and 0.727, respectively. Polymorphic information criteria (PIC) valued the markers as highly informative as PIC >0.5. Overall population, they are possibly in Hardy–Weinberg equilibrium with loci Ch_09ttn, Ch_01dn, and Ch_13dn showing signs of a null allele. There were no scoring errors caused by stutter peaks, no large allele dropout was detected for all loci and showed no evidence of linkage disequilibrium. In conclusion, all seven molecular microsatellite markers identified can be beneficially used to gain more information on the population genetic structure and breeding patterns of C. hemipterus as well as the relationship of dispersal and infestation.”


Balvín, O., T. Bartonička, K. Pilařová, Z. DeVries, and C. Schal2017. Discrimination between lineage-specific shelters by bat- and human-associated bed bugs does not constitute a stable reproductive barrier. Parasitology Research. 116: 237–242. doi: 10.1007/s00436-016-5284-y

“The common bed bug Cimex lectularius, has been recently shown to constitute two host races, which are likely in the course of incipient speciation. The human-associated lineage splits from the ancestral bat-associated species deep in the history of modern humans, likely even prior to the Neolithic Period and establishment of the first permanent human settlements. Hybridization experiments between these two lineages show that post-mating reproductive barriers are incomplete due to local variation. As mating takes place in off-host refugia marked by aggregation semiochemicals, the present investigation tested the hypothesis that bed bugs use these semiochemicals to differentiate between refugia marked by bat- and human-associated bed bugs; this would constitute a pre-copulation isolation mechanism. The preference for lineage-specific odors was tested using artificial shelters conditioned by a group of either male or female bed bugs. Adult males were assayed individually in four-choice assays that included two clean unconditioned control shelters. In most assays, bed bugs preferred to rest in conditioned shelters, with no apparent fidelity to shelters conditioned by their specific lineage. However, 51 % of the bat-associated males preferred unconditioned shelters over female-conditioned shelters of either lineage. Thus, bed bugs show no preferences for lineage-specific shelters, strongly suggesting that semiochemicals associated with shelters alone do not function in reproductive isolation.”

Masran, S., S. N. Ain, A. Majid, and A. Hafiz2017. Genetic diversity and phylogenetic relationships of cytochrome C oxidase subunit I in Cimex hemipterus (Hemiptera: Cimicidae) populations in Malaysia. Journal of Medical Entomology. 54: 974–979. doi: 10.1093/jme/tjw227

“The tropical bed bug is scientifically recognized as a significant public health problem. While there is an increased awareness about their resurgence by medical and life science committees, efficient bed bug management still remains unresolved. The solution may soon arise, as information about bed bugs’ infestation dynamics and systematics are becoming more distinguishable. Recent developments in studies about bed bugs are based on molecular intervention by determining their genetic variation and phylogeography. The aim of this study is to assess the phylogenetic relationships and genetic diversity among the populations of tropical bed bugs inhabiting Malaysia. A molecular genotyping study was conducted with 22 tropical bed bug populations composed of three individuals per population. The mitochondrial (COI) gene was used as a marker. The data obtained were analyzed using the T-Coffee, ClustalX, MEGA 6.0, and PAUP software. The results showed one main monophyletic clade that consisted of two groups: Ch01 and Ch02. Ch02 consists of samples from the Bandar Hilir population, differing from the other populations studied by one singleton base. However, as there were no changes in the amino acid, this singleton genetic variation was considered to have no effect on genetic differentiation. Ch01 shows similarity with some sequence of Cimex hemipterus (F.) from Thailand, suggesting an international diversity connection. The disparity index apparently suggests that all isolates are homogeneous populations and are supported by the low value of the mean pairwise distance between isolates. This study will increase the knowledge about phylogeographic diversity of tropical bed bug in Malaysia.”

Talbot, B., M. Vonhof, H. G. Broders, M. B. Fenton, and N. Keyghobadi2017. Population structure in two geographically sympatric and congeneric ectoparasites: Cimex adjunctus and Cimex lectularius, in the North American Great Lakes region. Canadian Journal of Zoology. doi: 10.1139/cjz-2017-0014

“Subdivided populations can be described by different models of population structure that reflect population organisation, dynamics and connectivity. We used genetic data to investigate population structure in two geographically sympatric, congeneric species of generalist ectoparasites of warm-blooded animals. We characterized the spatial genetic structure of Cimex adjunctus Barber, 1939, an understudied and fairly abundant species, using microsatellite markers at a spatial scale representing contemporary dispersal of the species. We found seven genetic clusters, global GST of 0.2, 33% of genetic variation among sites, and non-significant isolation-by-distance. We compared these results with Cimex lectularius L., 1758, a closely related but conversely well-known species, in the same geographic area. We found stronger genetic structuring in C. lectularius than in C. adjunctus, with eleven genetic clusters, GST of 0.7, 57% of genetic variation and significant but weak isolation-by-distance (R2 = 0.09). These results suggest that while both species can be described as having classic metapopulation structure, C. adjunctus leans more towards a patchy population and C. lectularius leans more towards a non-equilibrium metapopulation. The difference in population structure between these species may be attributable to differences in movement potential and extinction-colonization dynamics.”


Akhoundi, M., P. Kengne, A. Cannet, C. Brengues, J.-M. Berenger, A. Izri, P. Marty, F. Simard, D. Fontenille, and P. Delaunay2015. Spatial genetic structure and restricted gene flow in bed bugs (Cimex lectularius) populations in France. Infection, Genetics and Evolution. 34: 236–243. doi: 10.1016/j.meegid.2015.06.028

“Bed bugs (Cimex lectularius) are resurgent blood-sucking ectoparasites that are currently increasing at a rapid rate, particularly in industrialized countries, such as France. Despite the rapid spread of bed bugs, there is a lack of knowledge concerning the population structure and gene flow among C. lectularius populations in France. To fill this gap, a genetic study was conducted using 183 C. lectularius from 14 populations of bed bugs collected in a hotel and in individual apartments in the French Riviera and in the Saint Ouen suburb of Paris. The samples were genotyped using an isolated set of six polymorphic microsatellite loci, including five new loci which were newly isolated and chosen based on prior successful amplification, and one previously described loci (bb15b). The low genetic diversity observed in the samples (of one to five alleles) suggested that most of prospected populations were established by only a few individuals, possibly from a single mated female. The overall genetic differentiation was high and statistically significant (FST=0.556, p<0.0001). Pairwise analysis of the populations indicated significant genetic differentiation for 24 out of the 45 (53%) population pairs associated with FST, ranging from 0.0042 to 0.862. No obvious relationship between the level of genetic differentiation and the geographic distance was observed when considering all samples. Analysis with Structure software identified nine distinct genetic clusters within the dataset. These preliminary results help to elucidate the genetic structure and gene flow of C. lectularius populations in France; however, the available information should be expanded in further studies.”

Booth, W., O. Balvín, E. L. Vargo, J. Vilímová, and C. Schal2015. Host association drives genetic divergence in the bed bug, Cimex lectularius. Molecular Ecology. 24: 980–992. doi: 10.1111/mec.13086

“Genetic differentiation may exist among sympatric populations of a species due to long‐term associations with alternative hosts (i.e. host‐associated differentiation). While host‐associated differentiation has been documented in several phytophagus insects, there are far fewer cases known in animal parasites. The bed bug, Cimex lectularius, a wingless insect, represents a potential model organism for elucidating the processes involved in host‐associated differentiation in animal parasites with relatively limited mobility. In conjunction with the expansion of modern humans from Africa into Eurasia, it has been speculated that bed bugs extended their host range from bats to humans in their shared cave domiciles throughout Eurasia. C. lectularius that associate with humans have a cosmopolitan distribution, whereas those associated with bats occur across Europe, often in human‐built structures. We assessed genetic structure and gene flow within and among populations collected in association with each host using mtDNA, microsatellite loci and knock‐down resistance gene variants. Both nuclear and mitochondrial data support a lack of significant contemporary gene flow between host‐specific populations. Within locations human‐associated bed bug populations exhibit limited genetic diversity and elevated levels of inbreeding, likely due to human‐mediated movement, infrequent additional introduction events per infestation, and pest control. In contrast, populations within bat roosts exhibit higher genetic diversity and lower levels of relatedness, suggesting populations are stable with temporal fluctuations due to host dispersal and bug mortality. In concert with previously published evidence of morphological and behavioural differentiation, the genetic data presented here suggest C. lectulariusis currently undergoing lineage divergence through host association.”

Fountain, T., R. K. Butlin, K. Reinhardt, and O. Otti2015. Outbreeding effects in an inbreeding insect, Cimex lectularius. Ecology and Evolution. 5: 409–418. doi: 10.1002/ece3.1373

“In some species, populations with few founding individuals can be resilient to extreme inbreeding. Inbreeding seems to be the norm in the common bed bug, Cimex lectularius, a flightless insect that, nevertheless, can reach large deme sizes and persist successfully. However, bed bugs can also be dispersed passively by humans, exposing inbred populations to gene flow from genetically distant populations. The introduction of genetic variation through this outbreeding could lead to increased fitness (heterosis) or be costly by causing a loss of local adaptation or exposing genetic incompatibility between populations (outbreeding depression). Here, we addressed how inbreeding within demes and outbreeding between distant populations impact fitness over two generations in this re-emerging public health pest. We compared fitness traits of families that were inbred (mimicking reproduction following a founder event) or outbred (mimicking reproduction following a gene flow event). We found that outbreeding led to increased starvation resistance compared to inbred families, but this benefit was lost after two generations of outbreeding. No other fitness benefits of outbreeding were observed in either generation, including no differences in fecundity between the two treatments. Resilience to inbreeding is likely to result from the history of small founder events in the bed bug. Outbreeding benefits may only be detectable under stress and when heterozygosity is maximized without disruption of coadaptation. We discuss the consequences of these results both in terms of inbreeding and outbreeding in populations with genetic and spatial structuring, as well as for the recent resurgence of bed bug populations.”

Narain, R. B., S. Lalithambika, and S. T. Kamble2015. Genetic variability and geographic diversity of the common bed bug (Hemiptera: Cimicidae) populations from the Midwest using microsatellite markers. Journal of Medical Entomology. 52: 566–572. doi: 10.1093/jme/tjv061

“With the recent global resurgence of the bed bugs (Cimex lectularius L.), there is a need to better understand its biology, ecology, and ability to establish populations. Bed bugs are domestic pests that feed mainly on mammalian blood. Although bed bugs have not been implicated as vectors of pathogens, their biting activity inflicts severe insomnia and allergic reactions. Moreover, they have recently developed resistance to various insecticides, which requires further molecular research to determine genetic variation and appropriate interventions. Population dynamics, including genetic differentiation and genetic distance of 10 populations from the Midwest were analyzed in this study. The bed bug samples collected by pest control companies were genotyped using eight species-specific microsatellite markers. Results showed all eight markers were polymorphic, with 8-16 alleles per locus, suggesting high genetic diversity. The FST values were >0.25, signifying pronounced genetic differentiation. The G-test results also indicated high genetic differentiation among populations. The frequency of the most common allele across all eight loci was 0.42. The coefficient of relatedness between each of the populations was >0.5, indicative of sibling or parent-offspring relationships, while the FIS and its confidence interval values were statistically insignificant within the populations tested. The populations departed from Hardy-Weinberg equilibrium, possibly because of high heterozygosity. The genetic distance analysis using a neighbor-joining tree showed that the populations from Kansas City, MO, were genetically separate from most of those from Nebraska, indicating a geographic pattern of genetic structure. Our study demonstrated the effectiveness of using microsatellite markers to study bed bugs population structure, thereby improving our understanding of bed bug population dynamics in the Midwest. Overall, this study showed a high genetic diversity and identified several new alleles in the bed bug populations in the Midwest.”

Robinson, G. A., O. Balvin, C. Schal, E. L. Vargo, and W. Booth. 2015. Extensive mitochondrial heteroplasmy in natural populations of a resurging human pest, the bed bug (Hemiptera: Cimicidae). Journal of Medical Entomology. 52(4): 734-738. doi: 10.1093/jme/tjv055

“Homoplasmy, the occurrence of a single mitochondrial DNA haplotype within an individual, has been the accepted condition across most organisms in the animal kingdom. In recent years, a number of exceptions to this rule have been reported, largely due to the ease with which single nucleotide polymorphisms can be detected. Evidence of heteroplasmy—two or more mitochondrial variants within a single individual—has now been documented in a number of invertebrates; however, when present, heteroplasmy usually occurs at low frequencies both within individuals and within populations. The implications of heteroplasmy may be far reaching, both to the individual in relation to its health and fitness, and when considering the evolutionary dynamics of populations. We present novel evidence for frequent mtDNA heteroplasmy in the bed bug, Cimex lectularius L. (Hemiptera: Cimicidae). Our findings show that heteroplasmy is common, with 5 of 29 (17%) populations screened exhibiting two mitochondrial variants in a 1:2 ratio within each individual. We hypothesize that the mechanism underlying heteroplasmy in bed bugs is paternal leakage because some haplotypes were shared among unrelated populations and no evidence for nuclear mitochondrial DNA sequences was detected.”


Fountain, T., L. Duvaux, G. Horsburgh, K. Reinhardt, and R. K. Butlin2014. Human-facilitated metapopulation dynamics in an emerging pest species, Cimex lectularius. Molecular Ecology. 23: 1071–1084. doi: 10.1111/mec.12673

“The number and demographic history of colonists can have dramatic consequences for the way in which genetic diversity is distributed and maintained in a metapopulation. The bed bug (Cimex lectularius) is a re-emerging pest species whose close association with humans has led to frequent local extinction and colonization, that is, to metapopulation dynamics. Pest control limits the lifespan of subpopulations, causing frequent local extinctions, and human-facilitated dispersal allows the colonization of empty patches. Founder events often result in drastic reductions in diversity and an increased influence of genetic drift. Coupled with restricted migration, this can lead to rapid population differentiation. We therefore predicted strong population structuring. Here, using 21 newly characterized microsatellite markers and approximate Bayesian computation (ABC), we investigate simplified versions of two classical models of metapopulation dynamics, in a coalescent framework, to estimate the number and genetic composition of founders in the common bed bug. We found very limited diversity within infestations but high degrees of structuring across the city of London, with extreme levels of genetic differentiation between infestations (FST = 0.59). ABC results suggest a common origin of all founders of a given subpopulation and that the numbers of colonists were low, implying that even a single mated female is enough to found a new infestation successfully. These patterns of colonization are close to the predictions of the propagule pool model, where all founders originate from the same parental infestation. These results show that aspects of metapopulation dynamics can be captured in simple models and provide insights that are valuable for the future targeted control of bed bug infestations.”


Pereira, R. M., A. S. Taylor, M. P. Lehnert, and P. G. Koehler2013. Potential population growth and harmful effects on humans from bed bug populations exposed to different feeding regimes. Medical and Veterinary Entomology. 27: 148–155. doi: 10.1111/j.1365-2915.2012.01057.x.

“Effects of host availability and feeding period on bed bugs, Cimex lectularius (L.) (Hemiptera: Cimicidae), were measured. Population growth and the potential harmful effect of bed bug populations on human hosts were modelled. Bloodmeal sizes were affected by both feeding length and frequency, with >2-fold difference between insects fed daily or weekly. Blood consumption increased >2-fold between bed bugs fed occasionally and often, and 1.5-fold between occasional and daily feeding. Bed bugs fed more often than once a week, potentially every 2-4 days. Egg production was associated with nutrition, being strongly correlated with blood consumption in the previous week. Bed bug populations can grow under different feeding regimes and are hard to control with <80% mortality. Bed bugs can survive and grow even in locations with a limited blood supply, where bed bug persistence may be important for the continual spread of populations. Persistence in non-traditional locations and a potential association with human pathogens increase the health risks of bed bugs. Potential blood loss as a result of a bed bug can have serious consequences because uncontrolled populations can reach harmful levels in 3-8 months. The reproduction potential of bed bug populations suggests serious consequences to human health and the need for efficacious control measures.”


Saenz, V. L., W. Booth, C. Schal, and E. L. Vargo2012. Genetic analysis of bed bug populations reveals small propagule size within individual infestations but high genetic diversity across infestations from the Eastern United States. Journal of Medical Entomology. 49: 865–875. doi:

“Bed bugs (Cimex lectularius L.) are a resurgent pest worldwide and infestations within the United States are increasing at a rapid rate. Because of the physical and psychological discomfort inflicted by their blood feeding habits, and allergies and secondary infections associated with bites, bed bugs are recognized as a significant public health problem. Although bed bug infestations are spreading and becoming more prevalent, we have a poor understanding of their dispersal patterns and sources of infestation. To help fill this gap, we conducted a genetic study of 21 bed bug infestations from the Eastern United States, nearly all of which came from single rooms within residences. We genotyped samples comprised of 8-10 individuals per infestation at nine polymorphic microsatellite loci. Despite high genetic diversity across all infestations, with 5-17 alleles per locus (mean = 10.3 alleles per locus), we found low genetic diversity (1-4 alleles per locus) within all but one of the infestations. These results suggest that nearly all the studied infestations were started by a small propagule possibly consisting of a singly mated female and/or her progeny, or a female mated with multiple males that were highly related to her. All infestations were strongly genetically differentiated from each other (mean pairwise F(ST) between populations = 0.68) and we did not find strong evidence of a geographic pattern of genetic structure, indicating infestations located in closer proximity to each other were nearly as genetically differentiated as those located hundreds of kilometers away. The high level of genetic diversity across infestations from the eastern United States together with the lack of geographically organized structure is consistent with multiple introductions into the United States from foreign sources.”

Schaafsma, E. J., S. D. Hapke, and M. G. Banfield2012. Bed bug (Cimex lectularius L.) population composition as determined by baited traps. Insects. 3: 442–452. doi: 10.3390/insects3020442

“Two established field populations of bed bugs were sampled using host-mimicking traps baited with a combination of CO2, heat and a synthetic kairomone. The proportion of first instar nymphs (between 52% and 78% of all captured insects) was significantly higher than reported in previous studies, which had employed different sampling methods. The proportion of adults was correspondingly much lower than previously reported, between 5% and 7% of total capture. As many as 120 bed bugs were captured in a single trap in one night; the variation in catches between sampling locations within the same room and between days at the same location indicates that multiple nights of trapping may be required to obtain an accurate representation of population structure.”


Polanco, A. M., C. C. Brewster, and D. M. Miller2011. Population growth potential of the bed bug, Cimex lectularius L.: a life table analysis. Insects. 2: 173–185. doi: 10.3390/insects2020173

“Experimental life tables were constructed and analyzed for three strains of the common bed bug: a pyrethroid-susceptible laboratory strain (HS), a highly resistant field strain (RR), and a field strain with a declining level of resistance (KR). Egg to adult survival in the RR strain was 94% compared with 79% and 69% in the HS and KR strains, respectively. The RR strain also developed significantly faster from egg to adult (~35 days) than the other two strains (~40 days). Analysis of a survivorship and fecundity life table for the RR strain produced the following results. The average life expectancy for a newly laid egg was ~143 days, and that of a newly molted adult was ~127 days. Females produced an average of 0.64 daughter eggs/day with the highest weekly production during the fifth week of adult life. Analysis of daily reproductive parity showed that females produced 1–3 and 4–6 eggs on 79 and 21% of the days, respectively, when egg laying occurred. The net reproductive rate (R0) of the RR strain was ~35, which represents a 35-fold increase in the population per generation (~92 days). The intrinsic rate of increase, r, was 0.054 indicating that the population multiplies 1.1 times/female/day (λ) and doubles in size every 13 days. The stable age distribution (cx) was dominated by nymphs (54%), followed by eggs (34%) and adults (12%). Reproductive values (vx) for the strain increased from egg to the adult stage.”