Inter-clonal competition over queen succession imposes a cost of parthenogenesis on termite colonies.

In social insect colonies, selfish behaviour due to intracolonial conflict among members can result in colony-level costs despite close relatedness. In certain termite species, queens use asexual reproduction for within-colony queen succession but rely on sexual reproduction for worker and alate production, resulting in multiple half-clones of a single primary queen competing for personal reproduction. Our study demonstrates that competition over asexual queen succession among different clone types leads to the overproduction of parthenogenetic offspring, resulting in the production of dysfunctional parthenogenetic alates. By genotyping the queens of 23 field colonies of Reticulitermes speratus, we found that clone variation in the queen population reduces as colonies develop. Field sampling of alates and primary reproductives of incipient colonies showed that overproduced parthenogenetic offspring develop into alates that have significantly smaller body sizes and much lower survivorship than sexually produced alates. Our results indicate that while the production of earlier and more parthenogenetic eggs is advantageous for winning the competition for personal reproduction, it comes at a great cost to the colony. Thus, this study highlights the evolutionary interplay between individual-level and colony-level selection on parthenogenesis by queens.

Dynamics of dimorphic workers of Constrictotermes cyphergaster (Blattodea: Termitidae) during nest repair.

Termite nest repairs are considered a defensive conduct as they reduce the colony's exposure to the external environment. Repair activities are carried out by worker castes that can be polymorphic, representing a relationship between polymorphism and divisions of functions that can enhance task completion. Repairs are influenced by the extent of damage, nest volume, and the population dynamics of the building species, which regulate the recruitment of individuals for this activity. Our objective was to verify the performances (recruitment for repair) of dimorphic workers of Constrictotermes cyphergaster (Silvestri, 1901) during the damage repair activities performed on the external walls of termite nests of different sizes. We found a significant difference in the presence of dimorphic workers that performed repairs, with greater recruitment of the small morphotype, and observed an alternation of morphotypes between initial and final repair activities, with no influence of morphotype on the replacement pattern. Our results also showed that the total number of recruited workers decreased with increasing nest volume. These results help to better understand the social organization of a Nasutitermitinae termite species and the strategies adopted to protect its colonies.

Economic framework to assess the impact of banning pesticides, with application to sulfuryl fluoride for drywood termites (Blattodea: Kalotermitidae) in California.

Sulfuryl fluoride (SF) is a fumigant used to eliminate drywood termites (DWT: Kalotermitidae; Froggatt) and other structural pests. Because of its global warming potential, it has been suggested that SF be restricted as a greenhouse gas (GHG). We present an economic model to assess the net social cost of restricting SF. We consider 3 approaches to address DWT control- no treatment, allowing SF fumigation and localized treatments, and only local treatment. Each approach generates private and public benefits and costs. We estimate that the annual damage and home equity loss by DWT in California is US$4.5-16.8 billion without treatment. If fumigation is used on 20% of the houses and local treatments on the others, the combined social cost of treatment, damage, and GHG emissions are between US$1-US$2 billion annually. The annual cost of local treatments only would be between US$3.2 and US$4.9 billion. If the application of SF is severely restricted or banned, the social costs will increase between US$1.43 and US$4.31 billion annually. The implied cost per ton of CO2 eliminated is between US$624 and US$1,465, much above the price range of CO2 in other applications. The restriction/ban has significant equity and environmental effects, impacting low-income individuals living in rented properties and replacing damaged wood in housing will increase GHG emissions. We further recommend the continued use of SF until a comparable whole-structure alternative is developed that fits the parameters of our model.

Economic impact disharmony in global biological invasions.

A dominant syndrome of the Anthropocene is the rapid worldwide spread of invasive species with devastating environmental and socio-economic impacts. However, the dynamics underlying the impacts of biological invasions remain contested. A hypothesis posits that the richness of impactful invasive species increases proportionally with the richness of non-native species more generally. A competing hypothesis suggests that certain species features disproportionately enhance the chances of non-native species becoming impactful, causing invasive species to arise disproportionately relative to the numbers of non-native species. We test whether invasive species with reported monetary costs reflect global numbers of established non-native species among phyla, classes, and families. Our results reveal that numbers of invasive species with economic costs largely reflect non-native species richness among taxa (i.e., in 96 % of families). However, a few costly taxa were over- and under-represented, and their composition differed among environments and regions. Chordates, nematodes, and pathogenic groups tended to be the most over-represented phyla with reported monetary costs, with mammals, insects, fungi, roundworms, and medically-important microorganisms being over-represented classes. Numbers of costly invasive species increased significantly with non-native richness per taxon, while monetary cost magnitudes at the family level were also significantly related to costly invasive species richness. Costs were biased towards a few 'hyper-costly' taxa (such as termites, mosquitoes, cats, weevils, rodents, ants, and asters). Ordination analysis revealed significant dissimilarity between non-native and costly invasive taxon assemblages. These results highlight taxonomic groups which harbour disproportionately high numbers of costly invasive species and monetary cost magnitudes. Collectively, our findings support prevention of arrival and containment of spread of non-native species as a whole through effective strategies for mitigation of the rapidly amplifying impacts of invasive species. Yet, the hyper- costly taxa identified here should receive greater focus from managers to reduce impacts of current invasive species.

Can wood-feeding termites solve the environmental bottleneck caused by plastics? A critical state-of-the-art review.

The abundance of synthetic polymers has become an ever-increasing environmental threat in the world. The excessive utilization of plastics leads to the accumulation of such recalcitrant pollutants in the environment. For example, during the COVID-19 pandemic, unprecedented demand for personal protective equipment (PPE) kits, face masks, and gloves made up of single-use items has resulted in the massive generation of plastic biomedical waste. As secondary pollutants, microplastic particles (<5 mm) are derived from pellet loss and degradation of macroplastics. Therefore, urgent intervention is required for the management of these hazardous materials. Physicochemical approaches have been employed to degrade synthetic polymers, but these approaches have limited efficiency and cause the release of hazardous metabolites or by-products into the environment. Therefore, bioremediation is a proper option as it is both cost-efficient and environmentally friendly. On the other hand, plants evolved lignocellulose to be resistant to destruction, whereas insects, such as wood-feeding termites, possess diverse microorganisms in their guts, which confer physiological and ecological benefits to their host. Plastic and lignocellulose polymers share a number of physical and chemical properties, despite their structural and recalcitrance differences. Among these similarities are a hydrophobic nature, a carbon skeleton, and amorphous/crystalline regions. Compared with herbivorous mammals, lignocellulose digestion in termites is accomplished at ordinary temperatures. This unique characteristic has been of great interest for the development of a plastic biodegradation approach by termites and their gut symbionts. Therefore, transferring knowledge from research on lignocellulosic degradation by termites and their gut symbionts to that on synthetic polymers has become a new research hotspot and technological development direction to solve the environmental bottleneck caused by synthetic plastic polymers.

Termite diversity and damage assessment in urban trees from different parks of Lahore, Punjab, Pakistan.

Termite infestation is one of the fundamental problems associated with the loss of urban trees and ecological services. However, no such study has been performed in Pakistan to investigate the termite occurrence and assess such damages to urban trees caused by termites. For Lahore, research and comparable data on urban tree damages are rare or missing. This study surveyed six different microhabitats, including Bagh-e-Jinnah, canal vegetation, Model Town Park, Jallo Forestry, Race-Course Park, and FC College Vegetation employing the three belt transects (100 × 5 m) method. We geo-referenced termite-infested trees to investigate the termite occurrence on living and dead standing trees, termite diversity, and the assessment of tree damage by termites' attack. We recorded four termite species (Odontotermes obesus Rambur, Coptotermes heimi Wasmann, Heterotermes indicola Wasmann, and Microtermes obesi Holmgren) representing two families (Rhinotermitidae and Termitidae). However, the diversity indices revealed that O. obesus (higher termite) and C. heimi (lower termite) were dominant with 46.60 and 36% of occurrence among observed trees, respectively. Kernel density function indicated that the Lahore Canal and Bagh-e-Jinnah plantation shared all four termites' infestation evenly compared to other study sites. We observed the maximum number of damaged trees by termites in canal vegetation with the most damaged exotic tree species Populus euramericana along the canal green belt. Additionally, we observed significant (P < 0.05) termite-tree interactions with exotic, living, and dead standing tree species and found termite colony size positively (R = 0.985) correlated with the tree trunk diameter up to breast height (DBH). The average population of termites per unit volume of deadwood log was (0.39/cm3) within all plantation sites. In conclusion, this study provides simple, reckless, and inexpensive knowledge about the assessment of termite damage to trees, which may give a better idea in making decisions on tree selection and management in urban ecosystems.

Short and long-term costs of inbreeding in the lifelong-partnership in a termite.

Social life and lifelong partner commitments are expected to favor thorough partner choice, as an ill-suited partnership may have long-term consequences, adversely affecting the parents and spanning several cohorts of offspring. Here, we used ~1400 termite incipient colonies to estimate the short- and long-term costs of inbreeding upon the survival of the parents over a 15-month period, their productivity, and the resistance of their offspring toward pathogen pressure. We observed that foundation success was not influenced by the relatedness of partners, but by their levels of microbial load. We showed faster growth in inbred colonies with low levels of microbial load, revealing a potential tradeoff between pathogen defense and offspring production. Yet, inbreeding takes its toll later in colony development when offspring from incipient colonies face pathogen pressure. Although the success of a lifetime partnership is initially determined by the partner's health, the cost of inbreeding in incipient colonies favors outbred colonies reaching maturity.

Influences of the bark economics spectrum and positive termite feedback on bark and xylem decomposition.

The plant economics spectrum integrates trade-offs and covariation in resource economic traits of different plant organs and their consequences for pivotal ecosystem processes, such as decomposition. However, in this concept stems are often considered as one unit ignoring the important functional differences between wood (xylem) and bark. These differences may not only affect the performance of woody plants during their lifetime, but may also have important "afterlife effects." Specifically, bark quality may strongly affect deadwood decomposition of different woody species. We hypothesized that (1) bark quality strongly influences bark decomposability to microbial decomposers, and possibly amplifies the interspecific variation in decomposition by invertebrate consumption, especially termites; and (2) bark decomposition has secondary effects on xylem mass loss by providing access to decomposers including invertebrates such as termites. We tested these hypotheses across 34 subtropical woody species representing five common plant functional types, by conducting an in situ deadwood decomposition experiment over 12-month in two sites in subtropical evergreen broad-leaved forest in China. We employed visual examination and surface density measurement to quantify termite consumption to both bark and the underlying xylem, respectively. Using principal component analysis, we synthesized seven bark traits to provide the first empirical evidence for a bark economics spectrum (BES), with high BES values (i.e., bark thickness, nitrogen, phosphorus, and cellulose contents) indicating a resource acquisitive strategy and low BES values (i.e., carbon, lignin, and dry matter contents) indicating a resource conservative strategy. The BES affected interspecific variation in bark mass loss and this relationship was strongly amplified by termites. The BES also explained nearly half of the interspecific variation in termite consumption to xylem, making it an important contributor to deadwood decomposition overall. Moreover, the above across-species relationships manifested also within plant functional types, highlighting the value of using continuous variation in bark traits rather than categorical plant functional types in carbon cycle modeling. Our findings demonstrate the potent role of the BES in influencing deadwood decomposition including positive invertebrate feedback thereon in warm-climate forests, with implications for the role of bark quality in carbon cycling in other woody biomes.

In vitro assessment of two novel Cellulases from Trabulsiella odontotermitis for agricultural waste utilization.

BACKGROUND: The production of agricultural wastes still growing as a consequence of the population growing. However, the majority of these residues are under-utilized due their chemical composition, which is mainly composed by cellulose. Actually, the search of cellulases with high efficiency to degrade this carbohydrate remains as the challenge. In the present experiment, two genes encoding an endoglucanase (EC 3.2.1.4) and ß-glucosidase (EC 3.2.1.21) were overexpressed in Escherichia coli and their recombinant enzymes (egl-FZYE and cel-FZYE, respectively) characterized. Those genes were found in Trabulsiella odontermitis which was isolated from the gut of termite Heterotermes sp. Additionally, the capability to release sugars from agricultural wastes was evaluated in both enzymes, alone and in combination. RESULTS: The results have shown that optimal pH was 6.0 and 6.5, reaching an activity of 1051.65 ± 47.78 and 607.80 ± 10.19 U/mg at 39 °C, for egl-FZYE and cel-FZYE, respectively. The Km and Vmax for egl-FZYE using CMC as substrate were 11.25 mg/mL and 3921.57 U/mg, respectively, whereas using Avicel were 15.39 mg/mL and 2314.81 U/mg, respectively. The Km and Vmax for cel-FZYE using Avicel as substrate were 11.49 mg/mL and 2105.26 U/mg, respectively, whereas using CMC the enzyme did not had activity. Both enzymes had effect on agricultural wastes, and their effect was improved when they were combined reaching an activity of 955.1 ± 116.1, 4016.8 ± 332 and 1124.2 ± 241 U/mg on corn stover, sorghum stover and pine sawdust, respectively. CONCLUSIONS: Both enzymes were capable of degrading agricultural wastes, and their effectiveness was improved up to 60% of glucose released when combined. In summary, the results of the study demonstrate that the recombinant enzymes exhibit characteristics that indicate their value as potential feed additives and that the enzymes could be used to enhance the degradation of cellulose in the poor-quality forage generally used in ruminant feedstuffs.

How disease constrains the evolution of social systems.

Animal populations are occasionally shocked by epidemics of contagious diseases. The ability of social systems to withstand epidemic shocks and mitigate disruptions could shape the evolution of complex animal societies. We present a mathematical model to explore the potential impact of disease on the evolutionary fitness of different organizational strategies for populations of social species whose survival depends on collaborative efficiency. We show that infectious diseases select for a specific feature in the organization of collaborative roles-cohort stability-and that this feature is costly, and therefore unlikely to be maintained in environments where infection risks are absent. Our study provides evidence for an often-stated (but rarely supported) claim that pathogens have been the dominant force shaping the complexity of division of labour in eusocial societies of honeybees and termites and establishes a general theoretical approach for assessing evolutionary constraints on social organization from disease risk in other collaborative taxa.

Moving beyond the distinction between the bright and dark sides of termites to achieve sustainable development goals.

Termites are amongst the main macroinvertebrate decomposers in tropical ecosystems and they exert additional impacts through the creation of biostructures (mounds, galleries, sheetings, etc.) with different soil physical and chemical properties, thereby impacting positively on numerous ecosystem services for humankind. Unfortunately, this positive or 'bright' role of termites is often overshadowed by their 'dark' side, that is, their status as pests threatening agriculture and constructions. This article assesses advances in our knowledge of the impact of termites on several sustainable development goals (SDGs 1 'no poverty', 2 'zero hunger', 3 'good health', 9 'innovation', 11 'sustainable cities', 13 'climate action' and 15 'life on land'). Finally, using the Indian myth of Valmiki as a parable, we illustrate that a reconciliation between the termite's dark and bright sides is needed if we want to reduce our dramatic impact on biodiversity and more generally achieve SDGs.

Taxonomic assessment of the termite genus Neotermes (Isoptera: Kalotermitidae) in the Ryukyu-Taiwan Island arc, with description of a new species.

The Ryukyu Archipelago of Japan is a recognized global biodiversity hotspot with many endemic species. However, our knowledge of the termite fauna of the Ryukyu Archipelago is insufficient. Here, we report a new species of endemic drywood termite (Kalotermitidae) from the Ryukyu Archipelago. Our systematic study of the genus Neotermes (Isoptera: Kalotermitidae) from the Ryukyu-Taiwan Island arc using molecular and morphological methods found that N. koshunensis had also been composed of a second cryptic species, N. sugioi sp. nov. These two species are distributed allopatrically in the Ryukyu-Taiwan Island arc, with the former only in Taiwan and the latter only in the Ryukyu Archipelago. Our discovery of a new drywood termite species from the Ryukyu Archipelago suggests that both morphological and molecular assessments of the species now considered to be distributed widely in the Ryukyu-Taiwan Island arc are needed to clarify the termite fauna of the Ryukyu Archipelago. [Species Zoobank registration: http://zoobank.org/urn:lsid:zoobank.org:pub:E8C9693A-E24F-445D-8445-320564565964].

Use of chemically activated termite feces a low-cost adsorbent for the adsorption of norfloxacin from aqueous solution.

Antibiotics, as emerging contaminants, are of global concern due to the development of antibiotic resistant microorganisms. Current wastewater treatment technology cannot efficiently remove sewage antibiotics and therefore new low-cost technologies are needed. Adsorption is a widely used process for removal of substances, and the search for efficient, low-cost adsorbents is ongoing. In this work, termite feces treated with H2SO4 (FT/H2SO4) were used as a low-cost adsorbent for removal of norfloxacin (NOR) present in aqueous medium. Termite feces were treated with H2SO4 at a ratio of 1:1 for 24 h, at 100 °C. The parameters contact time, initial NOR concentration, medium pH and temperature were evaluated. The optimum adsorption pH was 8.0. The pseudo-second-order model was found to best represent the kinetics of NOR adsorption. The maximum adsorption capacity, calculated from the Sips isotherm model, was 104.4 mg/g at 55 °C. The positive values of ΔH0 (change in enthalpy) confirm the endothermic nature of the adsorption. The results show that FT/H2SO4 is an efficient adsorbent for removal of NOR present in aqueous medium. The adsorption capacity is higher than those reported in the literature for other low-cost adsorbents.

Juvenile hormone and hemimetabolan eusociality: a comparison of cockroaches with termites.

Termites are social Dictyoptera that evolved eusociality independently from social Hymenoptera. They are characterized by unique developmental plasticity that is the basis of caste differentiation and social organization. As developmental plasticity is a result of endocrine regulation, in order to understand the evolution of termite sociality it is helpful to compare the endocrine underpinning of development between termites and cockroaches. Nijhout and Wheeler (1982) proposed that varying JH titers determine caste differentiation in termites. Based on current results, we extend this model by adding the importance of social interactions. High JH titers in the presence of soldiers lead to regressive development (decrease in body size, apparent regression in development), while an absence of soldiers induces (pre-)soldier differentiation. On the opposite side, low JH titers in colonies headed by reproductives result in progressive molts toward adults, while an absence of reproductives induces development of replacement reproductives. In cockroaches, transcription factors involved in JH signaling, including the adult specifier E93 (the co-called MEKRE93 pathway) regulate the morphogenetic transition between the nymph and the adult. In termites, we speculate that castes might be determined by social effects playing a modulatory action of JH in the MEKRE93 pathway.

Cultural significance of termites in sub-Saharan Africa.

BACKGROUND: The number of termite species in the world is more than 2500, and Africa with more than 1000 species has the richest intercontinental diversity. The family Termitidae contains builders of great mounds up to 5 m high. Colonies are composed of casts: a queen, a king, soldiers and workers. Some species of termite cultivate specialised fungi to digest cellulose. Termites constitute 10% of all animal biomass in the tropics. The purpose of the study was to make an overview of how termites are utilized, perceived and experienced in daily life across sub-Saharan Africa. METHOD: Ethno-entomological information on termites (Isoptera) in sub-Saharan Africa was collected by: (1) interviews with more than 300 people from about 120 ethnic groups from 27 countries in the region; (2) library studies in Africa, London, Paris and Leiden. RESULTS: Vernacular names relate to mounds, insects as food, the swarming, and the behaviour of termites. Swarming reproductive, soldiers and queens are collected as food. There are many different ways to harvest them. Termites can also be used as feed for poultry or as bait to catch birds and fish. The mushrooms that grow each year from the fungus gardens on the termite mounds are eaten. The soldiers, the fungus gardens and the soil of termite mounds are used for multiple medicinal purposes. Mounds and soil of termites have numerous functions: for geochemical prospecting, making bricks, plastering houses, making pots, and for storage. Termite soil is often used as fertilizer. The act of eating soil (geophagy) among women, especially those that are pregnant, is practised all over Africa. The mounds can serve as burying places and are often associated with the spiritual world, especially containing the spirits of ancestors. Termites also play a role as oracle, in superstitious beliefs, in art and literature. CONCLUSION: The following characteristics make termites so appealing: the dominance in the landscape, the social organization, the destructive power, and the provision of food. The study shows that termites play a major role in peoples' lives, in physical as well as spiritual aspects.

Isolation and assessment of gut bacteria from the Formosan subterranean termite, Coptotermes formosanus (Isoptera: Rhinotermitidae), for paratransgenesis research and application.

Paratransgenesis targeting the gut protozoa is being developed as an alternative method for the control of the Formosan subterranean termite (FST). This method involves killing the cellulose-digesting gut protozoa using a previously developed antiprotozoal peptide consisting of a target specific ligand coupled to an antimicrobial peptide (Hecate). In the future, we intend to genetically engineer termite gut bacteria as "Trojan Horses" to express and spread ligand-Hecate in the termite colony. The aim of this study was to assess the usefulness of bacteria strains isolated from the gut of FST as "Trojan Horses." We isolated 135 bacteria from the guts of workers from 3 termite colonies. Sequencing of the 16S rRNA gene identified 20 species. We tested 5 bacteria species that were previously described as part of the termite gut community for their tolerance against Hecate and ligand-Hecate. Results showed that the minimum concentration required to inhibit bacteria growth was always higher than the concentration required to kill the gut protozoa. Out of the 5 bacteria tested, we engineered Trabulsiella odontotermitis, a termite specific bacterium, to express green fluorescent protein as a proof of concept that the bacteria can be engineered to express foreign proteins. Engineered T. odontotermitis was fed to FST to study if the bacteria are ingested. This feeding experiment confirmed that engineered T. odontotermitis is ingested by termites and can survive in the gut for at least 48 h. Here we report that T. odontotermitis is a suitable delivery and expression system for paratransgenesis in a termite species.

Massive yet grossly underestimated global costs of invasive insects.

Insects have presented human society with some of its greatest development challenges by spreading diseases, consuming crops and damaging infrastructure. Despite the massive human and financial toll of invasive insects, cost estimates of their impacts remain sporadic, spatially incomplete and of questionable quality. Here we compile a comprehensive database of economic costs of invasive insects. Taking all reported goods and service estimates, invasive insects cost a minimum of US$70.0 billion per year globally, while associated health costs exceed US$6.9 billion per year. Total costs rise as the number of estimate increases, although many of the worst costs have already been estimated (especially those related to human health). A lack of dedicated studies, especially for reproducible goods and service estimates, implies gross underestimation of global costs. Global warming as a consequence of climate change, rising human population densities and intensifying international trade will allow these costly insects to spread into new areas, but substantial savings could be achieved by increasing surveillance, containment and public awareness.

Assessment of genetically engineered Trabulsiella odontotermitis as a 'Trojan Horse' for paratransgenesis in termites.

BACKGROUND: The Formosan subterranean termite, Coptotermes formosanus is an invasive urban pest in the Southeastern USA. Paratransgenesis using a microbe expressed lytic peptide that targets the termite gut protozoa is currently being developed for the control of Formosan subterranean termites. In this study, we evaluated Trabulsiella odontotermitis, a termite-specific bacterium, for its potential to serve as a 'Trojan Horse' for expression of gene products in termite colonies. RESULTS: We engineered two strains of T. odontotermitis, one transformed with a constitutively expressed GFP plasmid and the other engineered at the chromosome with a Kanamycin resistant gene using a non- disruptive Tn7 transposon. Both strains were fed to termites from three different colonies. Fluorescent microscopy confirmed that T. odontotermitis expressed GFP in the gut and formed a biofilm in the termite hindgut. However, GFP producing bacteria could not be isolated from the termite gut after 2 weeks. The feeding experiment with the chromosomally engineered strain demonstrated that T. odontotermitis was maintained in the termite gut for at least 21 days, irrespective of the termite colony. The bacteria persisted in two termite colonies for at least 36 days post feeding. The experiment also confirmed the horizontal transfer of T. odontotermitis amongst nest mates. CONCLUSION: Overall, we conclude that T. odontotermitis can serve as a 'Trojan Horse' for spreading gene products in termite colonies. This study provided proof of concept and laid the foundation for the future development of genetically engineered termite gut bacteria for paratransgenesis based termite control.

Temporal Assessment of Molting in Workers of Formosan Subterranean Termites (Isoptera: Rhinotermitidae).

Molt frequency of workers in laboratory-reared juvenile colonies and foraging population from field colonies of Coptotermes formosanus Shiraki was determined using planar arenas in laboratory. Given that, chitin synthesis inhibitor (CSI)-incorporated baits disrupt the molting process of workers that comprises the major population of a termite colony, temporal assessment of molting frequency in workers can give insights into potential methods of reducing the time to eliminate a CSI-baited colony. In our study the 10-d observation of juvenile colonies of C. formosanus suggested average daily molting incidence of workers in a colony is 1.7 ± 0.3% (mean ± SD). The results from a time lapse study on foraging population of workers showed that on average there is a 44-d intermolt period for second-instar workers molting to third instar and 45 d for third-instar workers molting to fourth instar. At low temperature (21 °C), molting frequency of workers (0.6% per day) was significantly lower than that of workers at 27 °C (2.2% per day). Information from this study suggests that time to molt is an important component of total time for eliminating colonies treated with CSI baits and reduction in time lapse between two consecutive molts may reduce the time required for colony elimination.