The Emergence and Sustainability of Urban Entomology.

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 the 1970s to create a need for research and extension activities worldwide. This resulted in urban entomology as a discipline and, within two decades, its national and international recognition. In this review, we present the factors that led to the development of urban entomology and how they have shaped its current meaning. As urbanization intensifies and the global economy increases, the demands for urban pest management will continue to grow. We discuss how these future challenges may shape and alter the discipline.

Historical and Contemporary Control Options Against Bed Bugs, Cimex spp.

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 with global travel and poor pest management, which have contributed to their spread. This review examines and synthesizes the literature on bed bug origins and their global spread and the literature on historical and contemporary control options. This includes bed bug prevention, detection and monitoring, nonchemical and chemical control methodologies (and their limitations), and potential future control options. Future research needs are highlighted, especially the factors behind the modern resurgence, the necessity of identifying differences between the two bed bug species relevant to control, and the need to improve insecticide test protocols and management strategies.

Genetic analysis reveals the putative native range and widespread double-clonal reproduction in the invasive longhorn crazy ant.

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 clonally, resulting in separate male and queen lineages, while workers are produced sexually. Under this unusual reproduction mode, inbreeding is avoided in workers as they carry hybrid interlineage genomes. Despite the ubiquitous distribution of P. longicornis, the significance of this reproductive system for the ant's remarkable success remains unclear, as its prevalence is still unknown. Further investigation into the controversial native origin of P. longicornis is also required to reconstruct the evolutionary histories of double-clonal lineages. Here, we examine genetic variation and characterize the reproduction mode of P. longicornis populations sampled worldwide using microsatellites and mitochondrial DNA sequences to infer the ant's putative native range and the distribution of the double-clonal reproductive system. Analyses of global genetic variations indicate that the Indian subcontinent is a genetic diversity hotspot of this species, suggesting that P. longicornis probably originates from this geographical area. Our analyses revealed that both the inferred native and introduced populations exhibit double-clonal reproduction, with queens and males around the globe belonging to two separate, nonrecombining clonal lineages. By contrast, workers are highly heterozygous because they are first-generation interlineage hybrids. Overall, these data indicate a worldwide prevalence of double clonality in P. longicornis and support the prediction that the unusual genetic system may have pre-adapted this ant for global colonization by maintaining heterozygosity in the worker force and alleviating genetic bottlenecks.

Biology, Ecology, and Management of the Invasive Longlegged Ant, Anoplolepis gracilipes.

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 ecological consequences, especially for native invertebrate, vertebrate, and plant communities, altering ecosystem dynamics and functions. We examine and synthesize the literature about this species' origin and distribution, impacts on biodiversity and ecosystems, biology and ecology, chemical control, and potential biocontrol agents. We highlight emerging research needs on the origin and invasion history of this species, its reproductive mode, its relationship with myrmecophiles, and its host-microbial interactions, and we discuss future research directions.

Combined metabolic and target-site resistance mechanisms confer fipronil and deltamethrin resistance in field-collected German cockroaches (Blattodea: Ectobiidae).

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 extent of metabolic detoxification and target-site insensitivity resistance mechanisms. Topically applied doses of the 3 x LD95 of deltamethrin, fipronil, DDT, or dieldrin of a susceptible strain (UCR, Diagnostic Dose) failed to cause >23% mortality, and the 10 x LD95 of deltamethrin or fipronil failed to cause >53% mortality. All field-collected strains possessed a combination of metabolic and target-site insensitivity mechanisms that cause reduced susceptibility. Elevated activities of esterase and glutathione S-transferase were measured, and the synergists piperonyl butoxide or S,S,S-tributyl phosphorotrithioate increased topical mortality up to 100% for deltamethrin and 93% for fipronil 10 x LD95. The target-site mutations L993F of the para-homologous sodium channel and A302S of the GABA-gated chloride channel associated with pyrethroid and fipronil resistance, respectively, were found at ~80-100% frequency in field populations. Pyrethroid and fipronil spray formulations also were ineffective in a choice box assay against field-collected strains suggesting that these treatments would fail to control cockroaches under field conditions.

Fitness advantages of the biased use of paired laterally symmetrical penises in an insect.

The evolution of laterality, that is 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 Labidura riparia (Insecta: Dermaptera: Labiduridae) have morphologically similar laterally paired penises, only one of which is used for inseminating the female during a single copulation bout, and thus provide a rare opportunity to address how selection pressure may shape the evolution of population-level laterality. Our population studies revealed that in 10 populations, located at 2.23-43.3° north, the right penis is predominantly used for copulating (88.6%). A damaged penis was found in 23% of rare left-handers, suggesting that the left penis can function as a spare when the right one is damaged. By pairing L. riparia females with surgically manipulated males, we found that males forced to use the right penis outperformed left-handed males in copulation (the probability of establishing genital coupling during the 1-hr observation period: odds ratio [OR] of 3.50) and insemination (probability of transferring a detectable amount of sperm: OR of 2.94). This right-handed advantage may be due to the coiled morphology of the sperm storage organ with a right-facing opening. Thus, female genital morphology may play a significant role in the evolution of handedness and may have acted as a driving force to reduce penis number in related taxa.

Heads or tails: exaggerated morphologies in relation to the use of large bamboo internodes in two lizard beetles, Doubledaya ruficollis and Oxylanguria acutipennis (Coleoptera: Erotylidae: Languriinae).

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 species (Coleoptera: Erotylidae: Languriinae), Doubledaya ruficollis and Oxylanguria acutipennis, that excavate into large internode cavities of recently dead mature bamboos and have morphological modifications. We observed that females of D. ruficollis used their mandibles to bore oviposition holes on Schizostachyum sp. (mean wall thickness = 3.00 mm) and O. acutipennis did so on Dendrocalamus sp. (3.37 mm) bamboos. Previous studies suggested that the markedly asymmetrical mandibles and needle-like ovipositors of females in the genus Doubledaya are adaptive traits for excavating hard-walled bamboos for oviposition. Therefore, we measured their mandibular lengths and ovipositor lengths. D. ruficollis females had greater asymmetry in the mandibles and shorter and less-sclerotized ovipositors than females of congeners using small bamboos. In contrast, O. acutipennis females had slightly asymmetrical mandibles and elongated, well-sclerotized ovipositors. Oviposition holes of D. ruficollis were cone-shaped (evenly tapering), whereas those of O. acutipennis were funnel-shaped (tube-like at the internal apex). This suggests that D. ruficollis females excavate oviposition holes using the mandibles only, and O. acutipennis females use both the mandibles and ovipositors. These differences suggest different oviposition-associated morphological specialization for using large bamboos: the extremely asymmetrical mandibles in D. ruficollis and elongated, needle-like ovipositors in O. acutipennis.

Worker reproduction of the invasive yellow crazy ant Anoplolepis gracilipes.

BACKGROUND: Reproductive division of labor is one of the key features of social insects. Queens are adapted for reproduction while workers are adapted for foraging and colony maintenance. In many species, however, workers retain functional ovaries and can lay unfertilized male eggs or trophic eggs. Here we report for the first time on the occurrence of physogastric workers and apparent worker reproduction in the invasive yellow crazy ant Anoplolepis gracilipes (Fr. Smith). We further examined the reproductive potential and nutritional role of physogastric workers through multidisciplinary approaches including morphological characterization, laboratory manipulation, genetic analysis and behavioral observation. RESULTS: Egg production with two types of eggs, namely reproductive and trophic eggs, by physogastric workers was found. The reproductive egg was confirmed to be haploid and male-destined, suggesting that the workers produced males via arrhenotokous parthenogenesis as no spermatheca was discovered. Detailed observations suggested that larvae were mainly fed with trophic eggs. Along with consumption of trophic eggs by queens and other castes as part of their diet, the vital role of physogastric workers as "trophic specialist" is confirmed. CONCLUSION: We propose that adaptive advantages derived from worker reproduction for A. gracilipes may include 1) trophic eggs provisioned by physogastric workers likely assist colonies of A. gracilipes in overcoming unfavorable conditions such as paucity of food during critical founding stage; 2) worker-produced males are fertile and thus might offer an inclusive fitness advantage for the doomed orphaned colony.

A Natural Population Derived from Species Hybridizationin the Drosophila ananassae Species Complexon Penang Island, Malaysia.

We surveyed natural population of the Drosophila ananassae species complex on Penang Island, Malaysia. Analyses of phenotypic traits, chromosome arrangements, molecular markers, and reproductive isolation suggest the existence of two species: D. ananassae and D. cf. parapallidosa. Molecular marker analysis indicates that D. cf. parapallidosa carries chromosome Y and 4 introgressions from D. ananassae. Thus, D. cf. parapallidosa seems to be a hybrid descendant that recently originated from a natural D. parapallidosaâ™€× D. ananassae♂ cross. Furthermore, D. cf. parapallidosa behaves differently from authentic D. parapallidosa with respect to its reproductive isolation from D. ananassae. Premating isolation is usually seen in only the D. ananassaeâ™€× D. parapallidosa♂ cross, but we observed it in crosses of both directions between D. ananassae and D. cf. parapallidosa. In addition, hybrid males from the D. ananassaeâ™€× D. parapallidosa♂ cross are usually sterile, but they were fertile when D. ananassae♀ were mated with D. cf. parapallidosa ♂. We attempted an artificial reconstruction of the hybrid species to simulate the evolutionary process(es) that produced D. cf. parapallidosa. This is a rare case of natural hybrid population in Drosophila and may be a useful system for elucidating speciation with gene flow.

Paederus Outbreaks in Human Settings: A Review of Current Knowledge.

Although rove beetles (Paederus spp.) play a beneficial role as biological control agents to manage crop pests in agro-ecosystems, their high prevalence in human settings has elevated them to pest status in urban areas. Rove beetles neither bite nor sting, but accidental crushing on human skin causes them to release the toxin paederin, which causes dermatitis linearis. This review integrates currently available knowledge about the issues pertaining to Paederus infestation. For instance, the results of life history studies conducted under different food and temperature regimes are summarized, as they indicate how large a population can be in a habitat to cause massive and widespread infestation and illustrate the physiological traits required to maintain the population at the maximum level even under adverse conditions. In contrast to what is generally reported, we speculate that climatic factors do not necessarily result in Paederus dispersal in temperate regions; instead, habitat disturbance and site unsuitability may be the main factors that lead to massive dispersal to human settings. Factors such as whether dispersers are adaptable to xeric conditions in human settings, the probability that dispersed Paederus mate with the opposite sex, and whether dispersers have adequate nutrient intake to reproduce are considered to evaluate their potential to reproduce in human settings. Finally, the effectiveness of current commercial insecticides, challenges faced in managing infestations, and sustainable management practices are discussed to provide information for long-term control programs.

Induced Disturbances Cause Monomorium pharaonis (Hymenoptera: Formicidae) Nest Relocation.

Budding and relocation of nests are important characteristics of the Pharaoh ant, Monomorium pharaonis (L.), an important pest of artificial structures. Pharaoh ant colony movements induced by several types of disturbances were evaluated in the laboratory. The percentages of workers and brood in the source and new nest sites were determined at Days 0, 1, 3, and 5 following physical disturbance (temporal removal of nestmates), chemical disturbance (application of pyrethroid insecticide), invasion by heterospecific ants, food depletion, and moisture depletion in the laboratory. All disturbances were performed in the source nest, which was connected to an empty new nest site. Almost all workers moved and carried the entire brood to the new nest site when subjected to physical disturbance, chemical disturbance, and ant invasion on Day 1, whereas only <5% of workers were present in the new nest site in the undisturbed control. After these disturbances, the brood was never relocated back to the original nest site in this 5-d study. When subjected to food depletion, ∼60% of the brood were found in the new nest site and ∼40% of the brood remained in the original nest on Day 5, resulting in a polydomous population. In contrast, moisture depletion did not show any significant effect on colony movement. These results provide useful information about the causes of Pharaoh ant colony budding and guidance about how to develop effective control and prevention strategies.

A Laboratory Maintenance Regime for a Fungus-Growing Termite Macrotermes gilvus (Blattodea: Termitidae).

The optimum maintenance conditions of the fungus-growing termite, Macrotermes gilvus (Hagen) (Blattodea: Termitidae), in the laboratory were studied. Termites were kept on a matrix of moist sand and with fungus comb as food. The survival of groups of termites was measured when maintained at different population densities by changing group size and container volume. Larger groups (≥0.6 g) were more vigorous and had significant higher survival rates than smaller groups (≤0.3 g). The population density for optimal survival of M. gilvus is 0.0025 g per container volume (ml) or 0.0169 g per matrix volume (cm(3)), i.e., 1.2 g of termites kept in a 480-ml container filled with 71 cm3 of sand. In termite groups of smaller size (i.e., 0.3 g) or groups maintained in smaller container (i.e., 100 ml) the fungus comb was overgrown with Xylaria spp., and subsequently all termites died within the study period. The insufficient number of workers for regulating the growth of unwanted fungi other than Termitomyces spp. in the fungus comb is the most likely reason. Unlike some other mound-building termite species, M. gilvus showed satisfactory survival when maintained in non-nutritious matrix (i.e., sand). There was no significant difference in the survival rate between different colonies of M. gilvus (n=5), with survival in the range of 78.5-84.4% after 4 wk. Advances in the maintenance of Macrotermes will enable researchers to study with more biological relevance many aspects of the biology, behavior, and management of this species.

Investigation of Age Polyethism in Food Processing of the Fungus-Growing Termite Odontotermes formosanus (Blattodea: Termitidae) Using a Laboratory Artificial Rearing System.

Laboratory rearing systems are useful models for studying Rhinotermitid behavior. Information on the biology of fungus-growing termites, however, is limited because of the difficulty of rearing colonies in the laboratory settings. The physical structure of termite nests makes it impossible to photograph or to observe colonies in the field. In this study, an artificial rearing system for field-collected colonies of the fungus-growing termite Odontotermes formosanus (Shiraki) was developed to facilitate observation in the laboratory. We recorded colony activity within the artificial rearing system and documented a variety of social behaviors that occurred throughout the food processing of the colony. This complex miniature ecosystem was cooperatively organized via division of labor in the foraging and processing of plant materials, and the observed patterns largely resembled the caste and age-based principles present in Macrotermes colonies. This work extends our insights into polyethism in the subfamily Macrotermitinae.

Effect of diet quality on survival and reproduction of adult Paederus fuscipes (Coleoptera: Staphylinidae).

The survival and reproduction of the beetle Paederus fuscipes Curtis on diets that differed in macronutrient composition were investigated in the laboratory. The sex organs of females fed each test diet were dissected to evaluate the ovarian development over time. Adults fed on a carbohydrate-rich diet lived longer than adults fed the other diets, but this diet provided insufficient nutrients for reproduction. Females fed on a protein-rich diet had high fecundity; however, the mean longevity of reproductive adult P. fuscipes significantly was shortened by 20-30 d compared with longevity of 60-70 d for adults fed a carbohydrate-rich diet. In contrast, adults that were provided lipid-rich diet had low survival and fecundity. Overall, the development ofprevitellogenic follicles was significantly affected by diet regime and days since starting a particular diet. The follicle size of females given protein-rich diet increased 0.5-fold 2 wk after beginning the diet. In contrast, the development of the follicles was slow in females given the carbohydrate-rich diet and the follicles degenerated in females given the lipid-rich diet. In terms of ovarian maturation, females fed on a protein-rich diet contained mostly vitellogenic and chorionated follicles. In contrast, the ovarioles of females on the carbohydrate-rich diet were largely occupied with previtellogenic and vitellogenic follicles, whereas for a lipid-rich diet, the follicles remained at the previtellogenic stage throughout the experiment.

Pyrethroid resistance in Aedes aegypti larvae (Diptera: Culicidae) from Singapore.

We report the first comprehensive insecticide susceptibility status ofAedes aegypti (L.) larvae from Singapore. The study indicated that Ae. aegypti is susceptible to temephos, although resistance (RR50 = 1.29-4.43-fold) couldbe developing. Of high concern is the detection of moderate to high resistance to permethrin (RR50 = 29-47-fold) and etofenprox (RR50 = 14-34-fold). Biolarvicide Bacillus thuringiensis israelensis (Bti) remains effective. The insecticide susceptibility profile of Ae. aegypti larvae was found to be homogenous among the different sites studied across the island city. The addition of synergists piperonyl butoxide, S,S,S,-tributyl phosphorotrithioate, and triphenyl phosphate generally failed to enhance the toxicity of the insecticides investigated, suggesting an insignificant role of metabolic-based resistance, and a possible involvement of target site resistance. Further biochemical investigation of specific metabolic enzyme activities suggested that detoxifying enzymes, mono-oxygenases, esterases, glutathione S-transferases, and altered acetylcholinesterases, generally did not contribute to the resistance observed. This study clearly demonstrated that pyrethroid resistance is widespread among Ae. aegypti population and lowered susceptibility to organophosphates is developing.

Influences of pyriproxyfen on fecundity and reproduction of the Pharaoh ant (Hymenoptera: Formicidae).

The effects of the juvenile hormone analog pyriproxyfen (at concentrations of 0.1, 0.5, and 1%) on egg production, number of ovarioles, and length of oocytes were examined in queens of the Pharaoh ant Monomorium pharaonis (L.). Pyriproxyfen significantly reduced egg production in queens from week 3 onwards. Queens that were exposed to 1% pyriproxyfen stopped producing eggs at week 8. After 8 wk, ovaries were dissected from all queens, and the number of ovarioles and the length of the largest oocytes were recorded. The ovaries of queens in treated colonies were smaller than those in untreated queens, and the number of ovarioles in the ovaries was significantly lower in all pyriproxyfen-treated queens. Queens treated with the highest concentrations of pyriproxyfen tended to have significantly shorter oocytes than untreated queens. Histological studies of the ovaries revealed that pyriproxyfen caused vacuolation in the ovarioles, thickening of the tunica propria, development of small eggs, and underdevelopment of nurse cells and the follicular epithelium. Exposure to pyriproxyfen reduced egg production and induced severe morphological changes in the ovaries of queens, and the effects increased with increased concentration of pyriproxyfen.

Colony Size Affects the Efficacy of Bait Containing Chlorfluazuron Against the Fungus-Growing Termite Macrotermes gilvus (Blattodea: Termitidae).

The efficacy of chitin synthesis inhibitors (CSIs) against fungus-growing termites is known to vary. In this study, 0.1% chlorfluazuron (CFZ) cellulose bait was tested against medium and large field colonies of Macrotermes gilvus (Hagen). The termite mounds were dissected to determine the health of the colony. Individual termites (i.e., workers and larvae) and fungus combs were subjected to gas chromatography-mass spectrometry (GC-MS) analysis to detect the presence of CFZ. In this study, 540.0 ± 25.8 g (or equivalent to 540.0 ± 25.8 mg active ingredient) and 680.0 ± 49.0 g (680.0 ± 49.0 mg active ingredient) of bait matrix were removed by the medium- and large-sized colonies, respectively, after baiting. All treated medium-sized colonies were moribund. The dead termites were scattered in the mound, larvae were absent, population size had decreased by 90%, and the queens appeared unhealthy. In contrast, no or limited effects were found in large-sized colonies. Only trace amounts of CFZ were detected in workers, larvae, and fungus combs, and the population of large-sized colonies had declined by only up to 40%. This might be owing to the presence of large amount of basidiomycete fungus and a drastic decrease of CFZ content per unit fungus comb (a main food source of larvae) in the large-sized colonies, and hence reduced the toxic effect and longer time is required to accumulate the lethal dose in larvae. Nevertheless, we do not deny the possibility of CSI bait eliminating or suppressing the higher termite if the test colonies could pick up adequate lethal dose by installing more bait stations and prolonging the baiting period.

Mortality patterns in Coptotermes gestroi (Blattodea: Rhinotermitidae) following horizontal transfer of nonrepellent and repellent insecticides: effects of donor:recipient ratio and exposure time.

The donor: recipient ratio and the time of donor exposure to termiticide required for maximal toxicant transfer among termites are crucial information for the development of termite management plans. Most of the available information on termiticide toxicity came from temperate zonal termite species, whereas little is known about tropical Asian species. In this study, mortality patterns of recipient termites, Coptotermes gestroi (Wasmann) subjected to seven formulated insecticide exposures under different donor exposure times and donor: recipient ratios were examined. For fipronil, lethal transfer was not affected by donor exposure time but was affected by the mixing ratio. The moderate-to-less toxic termiticides (imidacloprid, indoxacarb, bifenthrin, chlorfenapyr, and chlorantraniliprole) required long exposure time and a high mixing ratio to ensure maximal uptake by recipient workers compared with fipronil. For chlorantraniliprole and chlorfenapyr, donors must constitute > 30% of the donor-recipient mixture to achieve 100% mortality of the recipient workers. Among the termiticides tested, cyantraniliprole was the most fast-killing insecticide against C. gestroi. The potential of lethal transfer among recipient termites does not necessarily require both high donor exposure time and a high mixing ratio, but the toxicity of a given termiticide against termites must be factored in to achieve colony elimination.

Insecticide resistance in social insects: assumptions, realities, and possibilities.

Insecticide resistance is an evolved ability to survive insecticide exposure. Compared with 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 central goal of this review is to explore factors that either limit resistance or the ability to detect it in eusocial insects. We surveyed the literature and found that resistance has been documented in bees, but in other pest groups such as ants and termites, the evidence is more sparse. We suggest the path forward for better understanding eusocial resistance should include more tractable experimental models, comprehensive geographic sampling, and targeted testing of the impacts of social, symbiont, genetic, and ecological factors.

Oral toxicity of an artificial sweetener sucralose on the German cockroach (Blattodea: Ectobiidae) and its impact on water balance and gut microbiome.

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, Blattella germanica (L.). In a nonchoice test, we evaluated 5, 10, and 20% sucralose solutions. Depending on the cockroach strains, mean mortality ranged from 62.5 to 92.5%, 15 to 55%, and 2.5 to 27.5% for 20, 10, and 5% sucralose, respectively. Next, we measured the impact of a 20% sucralose treatment on water loss rates in the cockroach strains. All strains lost 23.0-30.29% of body water by 6 d. Dehydrated cockroaches were more prone to be killed by sucralose than nondehydrated ones. Lastly, we evaluated the effect of 20% sucralose treatment on gut bacterial composition and found the diversity of gut bacteria in treated cockroaches was significantly reduced after 3 days, implicating a rapid change in the alimentary environment.