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Introduction
My research direction centers around the behavioral, ecological, and physiological adaptations of urban insect pests, especially understanding how these adaptations help them to thrive in the urban environment and their biological trade-offs. I am also interested in the roles of human activities and propagule pressure in invasion history of urban insect pests. Using the research findings obtained, my students and I design, evaluate, and integrate multiple management tactics to provide a system-level approach towards urban pest management. Our recent research activities focus on morphological and biological traits, insecticide resistance and its underlying mechanisms, endosymbiont roles, phylogenetics, population genetics, and environmental physiology of urban pests.
Additional affiliations
April 2002 - December 2002
April 2015 - August 2015
July 2011 - November 2011
Publications
Publications (384)
The management of subterranean termite pests remains a major challenge in Southeast Asia, where these pests cause significant structural and economic damage. Termite baiting has emerged as an effective option to conventional soil termiticides, offering a safer pest management approach with reduced chemical input into the
environment. In this paper,...
The global resurgence of bed bugs (Cimex lectularius L. and Cimex hemipterus [F.]) over the past 25 years has presented significant challenges to the pest management industry, with insecticide resistance as a leading cause of control failures. This review provides a synthesis of bed bug insecticide resistance research from 2018 to the present, high...
Three chitin synthesis inhibitors (CSIs), bistrifluron, chlorfluazuron, and noviflumuron, were evaluated for their toxicity and horizontal transfer against the western drywood termite, Incisitermes minor (Hagen), when used to treat wood. In a no-choice bioassay, bistrifluron provided significantly faster kill than chlorfluazuron or noviflumuron tre...
This study investigated how management strategies influence resistance profiles in German cockroach (Blattella germanica (L.)) populations and their impact on the performance of commercial gel baits containing fipronil, imidacloprid, and indoxacarb. Field populations from premises managed under 3 different strategies: Baiting, random insecticide (R...
This is the first report on the use of live cockroaches as feed for farmed arowanas. Firstly, the nymphal development period, reproductive potential, longevity and nutrient profiles of the various development stages of the lobster cockroach, Nauphoeta cinerea, were determined. The fecundity of each female cockroach averaged 82.9 nymphs. An increase...
The higher classification of termites requires substantial revision as the Neoisoptera, the most diverse termite lineage, comprise many paraphyletic and polyphyletic higher taxa. Here, we produce an updated termite classification using genomic-scale analyses. We reconstruct phylogenies under diverse substitution models with ultraconserved elements...
Argentine ants are a major pest in California. In this study, a biodegradable calcium alginate hydrogel with an aqueous boric acid bait was tested against Argentine ant populations in a citrus orchard and a vineyard. A new continuous method was developed to produce large quantities of hydrogel bait for the field test. Foraging activity levels of an...
The western drywood termite, Incisitermes minor (Hagen), causes significant economic damage to wood structures in the United States of America, especially California. When infestation is not widespread, localized insecticide injections may be useful for remedial control. However, the extensive gallery structure of drywood termites and their tendenc...
Isocycloseram is a new insecticide in the isoxazoline class that targets insect GABA-gated chloride channels. In this study, we evaluated a cockroach gel bait formulation containing 1% isocycloseram against a susceptible strain (UCR) and 5 field-collected strains (WM, RG386, Ryan, CDR, and SY) of the German cockroach, Blattella germanica (L.) (Blat...
We investigated the expression of 4 cytochrome P450 genes (CYP4G19, CYP6J1, CYP6K1, and CYP4C21) in 4 field-collected strains (WM, RG386, CDR, and Ryan) of the German cockroach, Blattella germanica (L.), collected from California. The UCR susceptible strain was used as a comparison. Topical assays using a diagnostic dose (3× LD95) of deltamethrin r...
Artificial or non-nutritive sweeteners are indigestible by most animals. Some sweeteners are orally toxic to insects and have received recent interest as potential safe insecticides due to their low mammalian toxicity. In this study, we investigated the oral toxicity of sucralose on insecticide-susceptible and resistant German cockroaches, Blattell...
Urban entomology is the study of arthropod and other pests of the urban environment. It has gained worldwide recognition as a distinct discipline. Its origin is associated with Walter Ebeling's publication Urban Entomology in 1975. Urbanization, invasive pests, increased demand for pest management services, and changes in legislation collided in th...
Insecticide resistance is an evolved ability to survive insecticide exposure. Compared to nonsocial insects, eusocial insects have lower numbers of documented cases of resistance. Eusocial insects include beneficial and pest species that can be incidentally or purposely targeted with insecticides. The goal of this review is to explore factors that...
This book is the first comprehensive resource drawing on all the literature on Coptotermes formosanus since Tokuichi Shiraki first described the species in 1909. The book covers the worldwide distribution of this species, its biogeography, and how it has dispersed from its native range in Southern China and Taiwan to different parts of the world. I...
Liquid termiticides remain the most popular option for subterranean termite management worldwide. This chapter describes the historical and present perspectives on liquid termiticide use against subterranean termites, particularly the Formosan subterranean termite (FST). We also discuss the types of termiticides (repellent and non-repellent) and th...
The Asian subterranean termite, Coptotermes gestroi (Wasmann), is a successful urban exploiter endemic to Southeast Asia. Over the past century, anthropogenic activity has spread this species far beyond its native range. Besides the Formosan subterranean termite, Coptotermes formosanus Shiraki, C. gestroi is the most economically important and inva...
Many animals take advantage of the shaded, humid, and protected environments in subcortical spaces, i.e., thin spaces under the loosened bark of dead trees. Permanent inhabitants of subcortical spaces often show specialized morphologies, such as a miniaturized or dorsoventrally flattened body. However, the evolutionary consequences of these special...
Subterranean termites in the genus Reticulitermes Holmgren 1913 are among the most economically important wood-destroying pests in the western United States. Yet, there remains uncertainty regarding the tax-onomy and biology of the species in this genus. The 2 species described as having distributions in this region are the western subterranean ter...
Histamine is a biogenic amine that regulates multiple physiological functions in diverse organisms, specifically playing a central role in the mammalian immune response. The common bed bug, Cimex lectularius L. (Hemiptera: Cimicidae), excretes histamine in large amounts in its feces as a component of its aggregation pheromone, which contaminates ho...
Throughout history, bed bugs have been persistent pests that have plagued humans for millennia, causing discomfort and sleepless nights. This two-part series delves into the fascinating journey of bed bug control methods employed by our ancestors and explores ancient remedies, cultural practices, and technological advancements that aimed to both pr...
We investigated insecticide resistance profiles of field populations of the German cockroach, Blattella germanica (L.), collected from central regions of Thailand. Seven strains (PW, RB, MTH, MTS, TL, AY, and SP) were evaluated with diagnostic doses (DD; 3 × LD 95 generated from a susceptible strain) of deltamethrin, fipronil, and imidacloprid usin...
Selenium, a naturally occurring metalloid, is an essential trace element for many higher organisms, including humans. Humans primarily become exposed to selenium by ingesting food products containing trace amounts of selenium compounds. Although essential in these small amounts, selenium exhibits toxic effects at higher doses. Previous studies inve...
Simple Summary
The western yellowjacket, Vespula pensylvanica, is an important seasonal pest of recreational and outdoor venues in the western United States. Effective control strategies are limited, and the objective of this study was to determine the potential of the isoxazoline fluralaner as a bait toxicant. A minimum of 27 colonies were detecte...
Due to their mutualistic relationship with plant pests, the Argentine ant is considered a major pest in subtropical fruit orchards and vineyards. Besides insecticide sprays, liquid baiting has been demonstrated as an effective method to suppress the Argentine ant populations. To improve the economic feasibility of liquid baiting, hydrogel materials...
Euborellia (Anisolabididae: Anisolabidinae) is one of the most speciose genera of earwigs (Dermaptera), and its species-level classification is difficult. To settle the classification of brachypterous species with abbreviated tegmina recorded from East and Southeast Asia, we examined the morphology and reproductive isolation of three tentative Eubo...
The longlegged ant Anoplolepis gracilipes (Smith) is a highly invasive tramp ant species known for its deleterious effects on native ecosystems. While tramp ants are associated with human activity, information on how different intensities of human activity affect their distribution is limited. This study investigated how anthropogenic activities af...
Katydids produce sound for signaling and communication by stridulation of the tegmina. Unlike crickets, most katydids are known to sing at ultrasonic frequencies. This has drawn interest in the investigation of the biophysics of ultrasonic sound production, detection, evolution, and ecology (including predator–prey interactions) of these katydids....
Clonal reproduction can provide an advantage for invasive species to establish as it can circumvent inbreeding depression which often plagues introduced populations. The world's most widespread invasive ant, Paratrechina longicornis, was previously found to display a double‐clonal reproduction system, whereby both males and queens are produced clon...
The DDT, deltamethrin, and malathion susceptibilities of five tropical bed bug (Cimex hemipterus) strains, and their resistance mechanisms were investigated using synergism studies and detection of kdr mutations. All C. hemipterus strains showed high resistance to DDT and deltamethrin and moderate to high resistance to malathion. Synergism studies...
Simple Summary
Based on both the morphological and molecular data of the earwig samples collected from a bakery in Beni-Suef, Egypt, we identified the species as Marava arachidis (Spongiphoridae). The predation capability of M. arachidis against the eggs and larvae (in-ground stages) of the Rhipicephalus annulatus tick under laboratory conditions w...
Bed bugs (Hemiptera: Cimicidae) are an important group of obligate hematophagous urban insect pests. The global resurgence of bed bugs, involving the common bed bug, Cimex lectularius L., and the tropical bed bug, Cimex hemipterus (F.), over the past two decades is believed to be primarily due to the development of insecticide resistance, along wit...
The yellow crazy ant, Anoplolepis gracilipes is a widespread invasive ant that poses significant threats to local biodiversity. Yet, compared to other global invasive ant species such as the red imported fire ant (Solenopsis invicta) or the Argentine ant (Linepithema humile), little is known about the diversity of RNA viruses in the yellow crazy an...
Bed bugs are an important group of medical and urban insect pests. They are obligate blood-feeders. Their bites may cause skin irritation and allergic reactions and, under some circumstances, may lead to mental and other health issues. Despite numerous discoveries on the biology of these obnoxious pests and progress in control strategies over the l...
New infestations of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae), were discovered in southern California, namely in Rancho Santa Fe and La Mesa (San Diego County) and Highland Park (Los Angeles County) in 2021. We investigated whether these new infestations were related to the previous infestations...
Some diopsid flies have sexually dimorphic eye stalks that are assumed to require considerable nutrition for growth but are advantageous in competition and courtship. According to the handicap theory, the eye span in some dimorphic species serves as a reliable signal of individual quality to an opponent. However, it is not well understood how well...
Modern bed bugs are resistant to multiple insecticide classes, particularly the pyrethroids. The efficacy of pyrethroid-impregnated mattress liners marketed for bed bug management has been variable. This study evaluated the efficacy of a permethrin-impregnated mattress liner, ActiveGuard, against 24 bed bug strains, consisting of both Cimex hemipte...
The residual performance of two pyrethroid-neonicotinoid mixture formulations: Temprid SC (10.5% beta- cyfluthrin and 21% imidacloprid) and Tandem (3.5% lambda-cyhalothrin and 11.6% thiamethoxam) on two substrates (glass and filter paper) against eight pyrethroid-resistant strains (BM-MY, BP-MY, CH-MY, GL-MY, KL-MY, SAJ-MY,TT-MY, and QLD-AU) of the...
Despite insecticide resistance issues, pyrethroids and fipronil have continued to be used extensively to control the German cockroach, Blattella germanica (L.) (Blattodea: Ectobiidae) for more than two decades. We evaluated the physiological insecticide resistance in five German cockroach populations collected from 2018 to 2020 and measured the ext...
Reproduction mode represents a key determinant for success of biological invasion as it influences the genetic variation and evolutionary potential of introduced populations. The world’s most widespread invasive ant, Paratrechina longicornis , was found to display an unusual double-clonal reproduction system, whereby both males and queens were prod...
The longlegged ant ( Anoplolepis gracilipes) is one of the most damaging invasive tramp ants globally. It is generally found between latitudes 27°N and 27°S in Asia, although it has been introduced to other continents. Its native range remains debatable, but it is believed to be in Southeast Asia. Anoplolepis gracilipes invasion has many serious ec...
Gel bait insecticides have been extensively used to manage the German cockroach, Blattella germanica (L.) (Blattodea: Ectobiidae), but issues with reduced effectiveness of such formulations are becoming increasingly common. We collected five field strains of German cockroaches in California and evaluated them against five commercial bait products [...
The modern resurgence of the common (Cimex lectularius L.) and tropical bed bugs (C. hemipterus [F.]) is thought to be primarily due to insecticide resistance. While there are many reports on insecticide resistance mechanisms in C. lectularius, such information in C. hemipterus is limited. We examined dichloro-diphenyl-trichloroethane (DDT), malath...
Over the last few years, there have been several exciting discoveries related to termite biology and management. This article reviews a few of the exciting findings and examines their potential in future termite management strategies.
The behavioral responses of two bed bug species, Cimex lectularius L. and C. hemipterus (F.), to conspecific or heterospecific nymphal aldehyde blends were examined using a two-choice olfactometer. Volatile cues from exuviae or a synthetic blend containing (E)-2-hexenal, 4-oxo-(E)-2-hexenal, (E)-2-octenal, and 4-oxo-(E)-2-octenal were tested. In bo...
Overexpression of a cytochrome P450 gene, CYP4G19, is known to associate with pyrethroid resistance in the German cockroach, Blattella germanica (L.) (Blattodea: Ectobiidae). In this study, we investigated the CYP4G19 expression level in 20 field-collected strains of B. germanica in Taiwan. We also examined the level of adult male susceptibility to...
The Formosan subterranean termite was first discovered in California in La Mesa, San Diego County in 1992. The original colony was baited, but newer infestations were found and were also baited, fumigated, or underwent corrective soil treatment. In 2018, a pest management professional found another infestation of the Formosan subterranean termite i...
As a species, the German cockroach is one of the most widespread indoor urban pests worldwide. While numerous products have been developed to control their spread, German cockroaches continue to contaminate food, transmit disease and cause significant, long-term economic expense to homes, restaurants, hospitals and more.
Biology and Management of...
A new infestation of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae), was discovered in Canyon Lake, Riverside County, California. We used three mitochondrial DNA (COI, COII, and 16S) and seven polymorphic microsatellite markers to characterize the genetic relationship of the colony with two other colo...
We unveiled the penile penetration mechanics of two earwig species, Echinosoma horridum, whose intromittent organ, termed virga, is extraordinarily long, and E. denticulatum, whose virga is conversely short. We characterised configuration, geometry, material and bending stiffness for both virga and spermatheca. The short virga of E. denticulatum ha...
Spillover of honey bee viruses have posed a significant threat to pollination services, triggering substantial effort in determining the host range of the viruses as an attempt to understand the transmission dynamics. Previous studies have reported infection of honey bee viruses in ants, raising the concern of ants serving as a reservoir host. Most...
Artificial sweeteners are sweet-tasting additives found in consumable products as substitutes for naturally occurring sugars. They are derived from plant extracts or manufactured by chemical synthesis. Ingestion of sweeteners by insects can lead to significant physiological effects, such as mortality, decreased fecundity, and behavioral change. Due...
Reproductive biology, including the mating behavior, ootheca rotation and deposition, and the embryonic development of a nocticolid cockroach Nocticola sp. were examined and described, and the phylogenetic issues regarding Nocticolidae and Blattodea were discussed. The abdominal terga II to IV are specialized in males, relating to the tergal glands...
Simple Summary
The common (Cimex lectularius) and tropical (C. hemipterus) bed bugs are widespread blood-feeding human pests. To better understand their chemical ecology, the current study quantified the four most abundant bed bug aldehydes in the bed bugs’ immature stages by extracting their freshly shed exuviae. We also quantified the two most ab...
A 2012-13 survey on Penang Island, Malaysia, revealed the existence of both Drosophila ananassae and Drosophila parapallidosa, the latter of which carries chromosomes Y and 4 from D. ananassae and thus is of hybrid origin. We collected the flies again from the same location in 2018. The hybrid population remained present, which suggests that the D....
Simple Summary
Bed bugs cause health and economic impacts due to their hematophagous behavior. The tropical bed bug, Cimex hemipterus (F.) is predominant in tropical and subtropical regions and modern strains demonstrate high levels of insecticide resistance making them a major challenge to control. In this present study, we investigated the effect...
While Wolbachia, an intracellular bacterial symbiont, is primarily transmitted maternally in arthropods, horizontal transmission between species has been commonly documented. We examined kleptoparasitism as a potential mechanism for Wolbachia horizontal transmission, using ant crickets and their host ants as the model system. We compared prevalence...
This paper explores the influence of COVID-19 regarding its effect on the pest management industry.
The yellow crazy ant, Anoplolepis gracilipes (Smith, 1857), is a widespread invasive ant in tropical and subtropical regions. In our study, we surveyed ant cricket species (Myrmecophilinae) associated with A. gracilipes in the Indo-Pacific region and provided a taxonomic revision using an integrative approach by combining morphological and molecula...
Insecticide resistance in the German cockroach, Blattella germanica (L.), is a significant challenge to the pest management professionals worldwide. We collected 24 field populations of B. germanica from different localities in Taiwan island, reared them for one to two generations, and evaluated them for their resistance to deltamethrin, propoxur,...
The recent discovery of multiple viruses in ants, along with the widespread infection of their hosts across geographic ranges, provides an excellent opportunity to test whether viral prevalence in the field is associated with the complexity of social interactions in the ant population. In this study, we examined whether the association exists betwe...
...
This study examined the presence of insecticide resistance in different developmental stages (adults, first instars, and eggs) of the tropical bed bug, Cimex hemipterus (F.) using several insecticide formulations. Adults and first instars of five strains (Queensland, Kuala Lumpur, Bukit Mertajam, Saujana, and Krystal Point) were evaluated using the...
Understanding the phylogeographic history of an invasive species may facilitate reconstructing the history and routes of its invasion. The longhorn crazy ant, Paratrechina longicornis, is a ubiquitous agricultural and household pest throughout much of the tropics and subtropics, but little is known about the history of its spread. Here, we examine...
The cavities of bamboos (Poaceae) are used by various animals. Most of the animals access these cavities either by existing cracks or by excavating bamboos with soft walls or small, thin-walled bamboos. Only a few animals excavate into the cavities of large and thick- and hard-walled internodes of mature bamboos. We studied two lizard beetle specie...
The longhorn crazy ant, Paratrechina longicornis (Latreille), is a ubiquitous agricultural and urban pest that has invaded most tropical and subtropical regions. Although P. longicornis has been found worldwide for more than a century, the genetic struc- ture, origin, and invasion history of this species have not yet been extensively studied, parti...
The behavioural ecology of ultrasonic-singing katydids is not well understood, and the general bioacoustics, barely known for a few Neotropical Meconematinae, tends to be overlooked for species from Southeast Asia. These include Asiatic species of Phlugidini, commonly known as crystal predatory katydids. One of its genera, Asiophlugis consists of 1...
Many insect species display daily variation of sensitivity to insecticides when they are exposed to the same concentration at different times during the day. To date, this has not been investigated in bed bugs. To address this, we explored circadian rhythms in insecticide susceptibility, xenobiotic metabolizing (XM) gene expressions, and metabolic...
Rattan is an important climbing palm taxon in Malaysian tropical rain forests. Many rattan species have unique structures directly associated with certain ant species. In this study, four rattan species (Daemonorops lewisiana, Calamus castaneus, Daemonorops geniculata and Korthalsia scortechinii) were inspected and documented in a field survey conc...
The evolution of laterality, i.e., the biased use of laterally paired, morphologically symmetrical organs, has attracted the interest of researchers from a variety of disciplines. It is, however, difficult to quantify the fitness benefits of laterality because many organs, such as human hands, possess multimodal functions. Males of the earwig Labid...
The insect male accessory gland (MAG) is an internal reproductive organ responsible for the synthesis and secretion of seminal fluid components, which play a pivotal role in the male reproductive strategy. In many species of insects, the effective ejaculation of the MAG products is essential for male reproduction. For this purpose, the fruit fly Dr...
Several taxa of Nemobiinae and Trigonidiinae from some islands and coasts of Pacific Ocean and Andaman Sea (Indian Ocean) are considered. The tribes Marinemobiini Gorochov, 1985 and Burcini Gorochov, 1986 are briefly redescribed, and the tribes Nemobiini Saussure, 1877 and Pteronemobiini Vickery, 1973 are also discussed. Some genera of Marinemobiin...
At the start of this book it was mentioned that Advances in the Biology and Management of Modern Bed Bugs (ABMMBB) is the first academic text since Robert L. Usinger’s seminal 1966 publication, The Monograph of Cimicidae. However, there are a number of key differences between the two texts. For example, Advances has over 60 different authors and fo...
The first comprehensive scholarly treatment of bed bugs since 1966
This book updates and expands on existing material on bed bugs with an emphasis on the worldwide resurgence of both the common bed bug, Cimex lectularius L., and the tropical bed bug, Cimex hemipterus (F.). It incorporates extensive new data from a wide range of basic and applied r...
“…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...
In a survey of pest management professionals, bed bugs were ranked as the most difficult of all urban pests to control, largely due to issues of insecticide resistance. Successful bed bug management relies on integrated pest management, encompassing non‐chemical means of control, as well as the judicious use of insecticides. Despite the resistance...
Many researchers are trying to develop fungus-based biological control methods against insect pests, including termites. This study explored the termite−fungus relationship using Globitermes sulphureus (Haviland) as a model species. Fungal species in termite mounds of G. sulphureus were isolated, purified, and identified. These fungal species were...
Dispersal of soil-dwelling organisms via the repeatedly exposed Sunda shelf through much of the Pleistocene in Southeast Asia has not been studied extensively, especially for invertebrates. Here we investigated the phylogeography of an endemic termite species, Macrotermes gilvus (Hagen), to elucidate the spatiotemporal dynamics of dispersal routes...
Additional M. gilvus samples used in microsatellite analysis.
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Partitioning schemes for the concatenated dataset based on GARLI and MrBayes optimization process.
Numbers in bracket represents 1st, 2nd and 3rd codon positions of COII gene respectively. Partitioning scheme with best AIC score is highlighted in bold.
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Bayes factor analysis.
Best partitioning scheme for Bayesian inference is shown in bold. Italicized values indicate number of parameters used to calculate the bayes factor.
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Maximum likelihood values and statistics for the alternative phylogenetic hypotheses calculated in GARLI and CONSEL.
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PCR conditions and primers used for the mtDNA sequencing.
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Permission to reprint copyrighted figure.
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Sampling details of M. gilvus across SE Asia.
Positive PCR amplifications are indicated with x symbol.
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Best-fit nucleotide substitution model based on hierarchical likelihood ratio test (hLRT) and Akaike’s information criteria (AIC).
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Pairwise ΦST values (lower diagonal) and geographical Euclidean distance (shortest distance between pairs of populations, km) used in isolation-by-distance analysis (upper diagonal).
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Allelic richness (AR), number of alleles (NA), number of private alleles (in parenthesis) and its percentages (PA) for each locus and population.
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