Effects of Menadione on Survival, Feeding, and Tunneling Activity of the Formosan Subterranean Termite

The Formosan subterranean termite, Coptotermes formosanus Shiraki, is a highly destructive pest and a cosmopolitan invasive species. Sustainable termite management methods have been improving with the search for novel insecticides that are effective, safe, and cost efficient. Menadione, also known as vitamin K3, is a synthetic analogue and biosynthetic precursor of vitamin K with low mammalian toxicity. Menadione has shown insecticidal activity in several insects, presumably due to interference with mitochondrial oxidative phosphorylation. However, little is known about its effectiveness against termites. In this study, we evaluated the toxicity and repellency of menadione in C. formosanus. Our results showed that menadione affected the survival and feeding activity of termites both in filter paper and substrate (sand) treatments, and menadione influenced termite tunneling activity in treated sand. In a no-choice assay, ≥90% mortality after seven days and minimal or no food consumption were recorded when sand was treated with menadione at 6 to 600 ppm. In a two-choice assay with a combination of treated and untreated sand, termites were deterred by menadione at 6 to 600 ppm and exhibited low mortality (≤30%) over seven days, while tunneling activity was prevented with 60 to 600 ppm of menadione treatment. Overall, our study demonstrated dose-dependent toxicity and repellency of menadione in C. formosanus. The potential use of menadione as an alternative termite control agent is discussed.

Differential time allocation of foraging workers in the subterranean termite

Background Foraging in group living animals such as social insects, is collectively performed by individuals. However, our understanding on foraging behavior of subterranean termites is extremely limited, as the process of foraging in the field is mostly concealed. Because of this limitation, foraging behaviors of subterranean termites were indirectly investigated in the laboratory through tunnel geometry analysis and observations on tunneling behaviors. In this study, we tracked subsets of foraging workers from juvenile colonies of Coptotermes formosanus (2-yr-old) to describe general foraging behavioral sequences and to find how foraging workers allocate time between the foraging site (food acquisition or processing) and non-foraging site (food transportation). Results Once workers entered into the foraging site, they spent, on average, a significantly longer time at the foraging site than the non-foraging site. Our clustering analysis revealed two different types of foraging workers in the subterranean termite based on the duration of time they spent at the foraging site and their foraging frequency. After entering the foraging site, some workers (cluster 1) immediately initiated masticating wood fragments, which they transferred as food boluses to recipient workers at the foraging site. Conversely, the recipient workers (cluster 2) moved around after entering the foraging site and received food from donating workers. Conclusions This study provides evidence of task specialization within foraging cohorts in subterranean termites.

GC/TOF-MS-Based Metabolomics Reveals Altered Metabolic Profiles in Wood-Feeding Termite Coptotermes formosanus Shiraki Digesting the Weed Mikania micrantha Kunth

Effective approaches to exploiting the biomass of the abundant invasive weed Mikania micrantha Kunth are limited. Termites have been a focus of significant attention as mediators of biomass-processing owing to their ability to digest lignocellulose. Here, the GC/TOF-MS approach was employed to assess the effects of a diet composed of M. micrantha leaves on Coptotermes formosanus workers, with the growth performance of these workers also being assessed. The workers increased their dietary intake when fed M. micrantha leaves, with a concomitant gradual increase in mortality rate. A total of 62 differentially abundant metabolites and nine significantly affected pathways were found when comparing termites fed M. micrantha leaves to pinewood. Key metabolites, including carbohydrates, polyols, 4-hydroxyphenylacetic acid, and their related metabolic pathways, suggested that termites can digest and utilize M. micrantha-derived lignocellulose. However, changes in the tryptophan metabolism, tyrosine metabolism, and sphingolipid metabolism suggest an adverse effect of M. micrantha leaves on antioxidant activity and signal transduction in termites. Overall, this study identified the key metabolites and pathways associated with the response of these termites to dietary changes and the effect of M. micrantha on termites.

Comparative Antennal Morphometry and Sensilla Organization in the Reproductive and Non-Reproductive Castes of the Formosan Subterranean Termite

Antennae are the primary sensory organs in insects, where a variety of sensilla are distributed for the perception of the chemical environment. In eusocial insects, colony function is maintained by a division of labor between reproductive and non-reproductive castes, and chemosensation is essential for regulating their specialized social activities. Several social species in Hymenoptera display caste-specific characteristics in antennal morphology and diversity of sensilla, reflecting their differential tasks. In termites, however, little is known about how the division of labor is associated with chemosensory morphology among castes. Using light and scanning electron microscopy, we performed antennal morphometry and characterized the organization of sensilla in reproductive (female and male alates) and non-reproductive (worker and soldier) castes in the Formosan subterranean termite, Coptotermes formosanus Shiraki. Here, we show that the antennal sensilla in alates are twice as abundant as in workers and soldiers, along with the greater number of antennal segments and antennal length in alates. However, all castes exhibit the same types of antennal sensilla, including basiconicum, campaniformium, capitulum, chaeticum I, chaeticum II, chaeticum III, marginal, trichodeum I, and trichodeum I. The quantitative composition of sensilla diverges between reproductive and non-reproductive castes, but not between female and male alates or between worker and soldier castes. The sensilla display spatial-specific distribution, with basiconicum exclusively and capitulum predominantly found on the ventral side of antennae. In addition, the abundance of chemosensilla increases toward the distal end of antennae in each caste. This research provides morphological signatures of chemosensation and their implications for the division of labor, and suggests future neurophysiological and molecular studies to address the mechanisms of chemical communication in termites.

Collaborative Response of the Host and Symbiotic Lignocellulytic System to Non-Lethal Toxic Stress in Coptotermes formosanus Skiraki

Disturbing the lignocellulose digestive system of termites is considered to be a promising approach for termite control. The research on the tolerance mechanism of the termite lignocellulose digestive system to harmful environment conditions is limited. In this study, we keep Coptotermes formosanus Skiraki under a non-lethal toxic condition by feeding the termites with filter paper containing the kojic acid (a low toxic insecticide). The effects of low toxic stress on the activities and gene expressions of host/symbiotic originated lignocellulases, and on the symbiotic microbial community structure of C. formosanus were explored. Our result showed that the low toxic stress would lead to the synchronous decrease of cellulase and hemicellulase activities, and supplementary increase of corresponding gene expressions. The symbiotic community maintained its role as the main force in the lignocellulolytic system of C. formosanus. Meanwhile, a large number of rare taxa were significantly enriched by kojic acid treatment. These numerically inconspicuous bacterial populations might be responsible for the functions similar to phenoloxidase or insecticide detoxification and enable C. formosanus to tolerate the harmful environment. Overall, our data suggested that the digestive adaptation of C. formosanus to physiotoxic feeding is closely related to the triple collaboration of termites–flagellates–bacteria.

Termite taxonomy from 2001–2021: the contribution of Zootaxa

Termites comprise a relatively small group of insects, with 3176 known species (2976 living and 200 fossil) (Constantino 2020). They include, however, very important urban and agricultural pests, and also major decomposers of plant matter in terrestrial ecosystems, especially in the tropics. For instance, the annual economic impact of a single invasive termite species, Coptotermes formosanus, was estimated as US$11 billion in the United States in 1999 (Su 2002), placing it among the most important insect pests in the world.

Genetic Analysis of Formosan Subterranean Termite (Blattodea: Rhinotermitidae) Populations in California

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 colonies that were collected in 1992 and 2018 in La Mesa, San Diego County. Maximum likelihood phylogeny of C. formosanus based on concatenated COI and COII sequences revealed that the two La Mesa populations (CA01 and CA02) and the Canyon Lake population (CA03) were from different maternal lineages. Based on the 14 COII haplotypes of C. formosanus found world-wide, CA01 and CA02 belonged to a haplotype widely distributed across the United States, while CA03 was grouped under a haplotype predominantly found in Asia. Microsatellite allele frequencies across all loci for both La Mesa populations were relatively similar, but significant genetic differences were found between CA02 and CA03 colonies (FST = 0.24; Dest = 0.30; G″ ST = 0.55; P < 0.01).

Colony Suppression and Possible Colony Elimination of the Subterranean Termites Coptotermes formosanus and Reticulitermes speratus by Discontinuous Soil Treatment Using a Diluent of Fipronil Suspension Concentrate

We assessed the efficacy of a discontinuous soil treatment using a diluent of fipronil suspension concentrate in controlling colonies of Coptotermes formosanus and Reticulitermes speratus. In-ground monitoring stations were installed at Isogi Park and Kindai University, and individual termites inhabiting the stations were collected for four or six years to determine the numbers and locations of colonies present in test areas before and after the discontinuous soil treatment. Microsatellite genotyping indicated that two C. formosanus and two R. speratus colonies in the test area at Isogi Park and five R. speratus colonies in the test area at Kindai University were active and that their territories fluctuated every year. One of the two C. formosanus colonies at Isogi Park and one of the five R. speratus colonies at Kindai University were subjected to discontinuous soil treatments with fipronil and were strongly affected by the treatment at the colony level, resulting in the suppression and possible elimination of colonies. Termite activity of the fipronil-treated colony of C. formosanus was detected within one week after the discontinuous soil treatment and was not found for more than two years (28 months), while termite activity of the fipronil-treated colony of R. speratus was detected within four days and three weeks after the discontinuous soil treatment and was not detected thereafter for three years. Fipronil residue analysis showed that workers of C. formosanus moved at least 28 m and that workers of R. speratus moved 6 m from the treated soil locations for up to three weeks.

Coptotermes formosanus (Formosan subterranean termite).

C. formosanus is often transported by boats and shipping containers to port cities before being carried further inland via landscape materials such as railroad ties (railway sleepers). This may explain the current C. formosanus distribution in the USA with coastal areas more densely infested than inland areas (Hochmair and Scheffrahn, 2010). Temperature and humidity are primary factors affecting the establishment of C. formosanus, and it is potentially invasive to areas of high humidity approximately 35° north and south of the equator (Su and Tamashiro, 1987). Competition from native species is another limiting factor for many exotic pests, but C. formosanus is more aggressive and is known to out-compete the endemic termites such as Reticulitermes species. Another factor that has allowed the successful establishment and spread of C. formosanus in exotic areas has been the pest control industry's heavy reliance on soil termiticide barriers for subterranean termite control since the 1950s. Numerous studies, using mark-recapture methods, have revealed that a single colony of C. formosanus might contain several million termites that forage up to 100 m in the soil (Lai, 1977; Su and Scheffrahn, 1988). These agree with the results of excavation studies for C. formosanus colonies (Ehrhorn, 1934; King and Spink, 1969). Because of the large colony size, the application of soil termiticides beneath a structure does not usually have a major impact on the overall population, and the surviving colony continues to produce alates that can further infest nearby areas. Once established, C. formosanus has never been completely eradicated from an area. The dependency of soil termiticide barriers as the primary tool for subterranean termite control is probably the main reason for the establishment and spread of C. formosanus from four isolated port cities in the 1960s in the USA to all south-eastern states by 2001.

Concentration-Dependent Feeding Deterrence to 20-Hydroxyecdysone for Three Subterranean Termite Species (Blattodea: Rhinotermitidae)

Effective active ingredients in toxicant bait formulations must be non-deterrent to insect feeding behavior at lethal concentrations. This study evaluated feeding deterrence for Coptotermes formosanus Shiraki, C. gestroi (Wasmann), and Reticulitermes flavipes (Kollar) when provided access to cellulose impregnated with various concentrations of the insect molting hormone, 20-hydroxyecdysone (20E). Termites were exposed to 20E concentrations of 200, 500, 1000 and 2000 ppm and to noviflumuron at 5000 ppm in a 24 h choice-test, and the mass of substrate consumption from treated and untreated media pads was compared for each treatment. 20E feeding deterrence was detected at 500, 1000 and 2000 ppm for C. gestroi, and at 2000 ppm for C. formosanus. No significant differences in consumption of treated and untreated substrate was detected at any concentration for R. flavipes. Potential methods for reducing deterrence are discussed.