Revisiting Coptotermes (Isoptera: Rhinotermitidae): a global taxonomic road map for species validity and distribution of an economically important subterranean termite genus

Coptotermes Wasmann (Isoptera: Rhinotermitidae) is one of the most economically important subterranean termite genera and some species are successful invaders. However, despite its important pest status, the taxonomic validity of many named Coptotermes species remains unclear. In this study, we reviewed all named species within the genus and investigated evidence supporting the validity of each named species. Species were systematically scrutinized according to the region of their original description: Southeast Asia, India, China, Africa, the Neotropics, and Australia. We estimate that of the currently 69 named species described by accepted nomenclatural rules, only 21 taxa have solid evidence for validity, 44 names have uncertain status, and the remaining species names should be synonymized or were made unavailable. Species with high degrees of invasiveness may be known under additional junior synonyms due to independent parochial descriptions. Molecular data for a vast majority of species are scarce and significant effort is needed to complete the taxonomic and phylogenetic revision of the genus. Because of the wide distribution of Coptotermes, we advocate for an integrative taxonomic effort to establish the distribution of each putative species, provide specimens and corresponding molecular data, check original descriptions and type specimens (if available), and provide evidence for a more robust phylogenetic position of each species. This study embodies both consensus and contention of those studying Coptotermes and thus pinpoints the current uncertainty of many species. This project is intended to be a roadmap for identifying those Coptotermes species names that need to be more thoroughly investigated, as an incentive to complete a necessary revision process.

Toxicidade de produtos sintetizados a partir de extratos de plantas e do bistrifluron para o cupim subterrâneo Coptotermes gestroi (Wasmann, 1896) (Isoptera; Rhinotermitidae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Rehydration after water stress in forager workers of coptotermes gestroi (wasmann) (blattaria: rhinotermitidae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

The taxonomic status of pest species of Coptotermes in Southeast Asia: Resolving the paradox in the pest status of the termites, Coptotermes gestroi, C-havilandi and C-travians (Isoptera : Rhinotermitidae)

The taxonomic status of Coptotermes gestroi (Wasmann), C. havilandi Holmgren, C. travians (Haviland) and C. borneensis Oshima (Isoptera: Rhinotermitidae) is revised. The apparent discrepancy between the reported importance of C. havitandi in countries to which it has been introduced and the region from which it originated is shown to be due to misidentification and taxonomic confusion between C. travians, C. havilandi and C. gestroi. Based on an examination of specimens from Southeast Asia, two species are recognized, namely C. gestroi and C. travians. Coptotermes havilandi, described from imagos, is shown to be the same species as C. gestro described earlier from the soldier caste, and is designated a junior synonym. Coptotermes gestroi occurs from Assam through Burma and Thailand to Malaysia and the Indonesian archipelago, and has been introduced into other geographic regions, including parts of North and South America and the Caribbean. It is frequently found damaging wood in buildings, and is often intercepted outside its range in cargo onboard ships and sailing vessels, which is a likely mechanism for its spread to new geographical areas. Coptotermes gestroi has been misidentified in much literature as C. travians. Conversely, C. travians has been misidentified in recent literature in Peninsular Malaysia as C. havilandi and was redescribed from Borneo as C. borneensis, which is here designated a junior synonym of C. travians. It has a known distribution from Peninsular Malaysia to Borneo, and has not been found infesting wood in buildings. It is envisaged that the resolution of this taxonomic problem will enable the deployment of common pest management strategies for C. gestro the primary pest species of Coptotermes originating from Southeast Asia.

Neotenic formation in laboratory colonies of the termite Coptotermes gestroi after orphaning

The termite Coptotermes gestroi (Wasmann 1896) ( Rhinotermitidae: Coptotermitinae) is an exotic species in Brazil and information concerning its reproductive developmental biology is scarce. We induced the formation of neotenics in laboratory colonies through orphaning experiments. Orphaning experiments were conducted in three-year old colonies of C. gestroi kept under laboratory conditions. After three months, eight nymphoid neotenics were observed in one colony after queen removal. Histological analysis showed that these neotenics were non-functional. The results suggest that these individuals may have arisen from the first nymphal instar (N1) or from an early N1 instar after one or two larval moults. Neotenics also were recorded on two incipient colonies of C. gestroi that lost the queen naturally.

Laboratory Assessment of two Active Ingredients for Control of Coptotermes gestroi (Isoptera: Rhinotermitidae)

The subterranean termite Coptotermes gestroi Wasmann (Isoptera: Rhinotermitidae) is a pest of major economic importance in urban environments of southeastern Brazil. Compared to using pesticides for termite control, termite bait products target termites more specifically and also address environmental contamination issues. In this point of view, we performed two different bioassays ill order to evaluate the efficacy of boric acid and fipronil against different populations of C. gestroi. The results showed that concentration between 2000 and 3000 ppm of boric acid caused approximately 100 percent mortality ill termites. Concentrations between 0.01 and 0.0001 ppm of fipronil resulted in 100% termite mortality after 2 wk exposure. The data displayed a fast mortality of termites contaminated with fipronil, even with small concentrations, and therefore it is riot a Suitable product to be used ill baits against C gestroi. The present Study showed a delayed toxicity of boric acid against the subterranean termite C gestroi which suggests a need for further field tests.

Nymphs in foraging populations of Coptotermes gestroi (Isoptera: Rhinotermitidae)

Food searching activity by subterranean termites involves workers, soldiers and nymphs. The present study was carried out for elucidating aspects of nymph foraging behavior in Brazil where the species Coptotermes gestroi (Wasmann, 1896) is exotic. Monitoring was conducted through making regular collections from seven different colonies for a period of 22 months, which exhibited the production of forager nymphs all year round. A biometric study was conducted to determine the instars of the forager nymphs collected. The identification of male and female nymphs was also conducted for establishing the sex proportion, which was slightly male-biased. From the 533 forager nymphs collected, 34 were from the 3rd instar, 280 from the 4th instar and 219 from 5th instar. The results suggest that 3rd instar nymphs were not too active in the foraging activities because they were collected only in one colony and in an insignificant quantity. (C) Koninklijke Brill NV, Leiden, 2011.

Reproductive strategy of Coptotermes gestroi (Isoptera: Rhinotermitidae) in Brazil

The termite Coptotermes gestroi (Wasmann) (Rhinotermitidae) is an exotic species in Brazil. This species forms colonies headed by primary reproductives, however, non-functional neotenics are commonly found even in the presence of the imaginai pair. The presence of non-functional neotenics in C. gestroi colonies may correspond to a strategy of this termite species for rapid colonization in foreign areas, instead of a response to chemical control methods.

Dynamics of foraging and recruitment behavior in the Asian subterranean termite Coptotermes gestroi (Rhinotermitidae)

The present study investigated the trail-following behavior of the subterranean termite Coptotermes gestroi (Wasmann Rhinotermitidae) under laboratory conditions. The results showed that workers were the first to initiate the exploration to the food source. When food was discovered they returned to the nest laying a trail for recruiting nestmates to the food source. In this situation, workers always traveled significantly faster when returning from the arenas. Both workers and soldiers were recruited to the food source; however, the soldier/worker proportion was higher during the first phase of the recruitment. When no food was available, the number of recruited nestmates and the speed on their way back to the nest were significantly lower. The results also showed that scout foragers always laid trail pheromones when entering into unknown territories, and that chemical signals found in the food could induce workers of C. gestroi to increase their travel speed. Copyright © 2012 Alberto Arab et al.

Seleção de ingredientes ativos para o controle do cupim subterrâneo Coptotermes gestroi (Isoptera: Rhinotermitidae)

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

The origins and radiation of Australian Coptotermes termites: From rainforest to desert dwellers

The termite genus Coptotermes (Rhinotermitidae) is found in Asia, Africa, Central/South America and Australia, with greatest diversity in Asia. Some Coptotermes species are amongst the world’s most damaging invasive termites, but the genus is also significant for containing the most sophisticated mound-building termites outside the family Termitidae. These mound-building Coptotermes occur only in Australia. Despite its economic and evolutionary significance, the biogeographic history of the genus has not been well investigated, nor has the evolution of the Australian mound-building species. We present here the first phylogeny of the Australian Coptotermes to include representatives from all described species. We combined our new data with previously generated data to estimate the first phylogeny to include representatives from all continents where the genus is found. We also present the first estimation of divergence dates during the evolution of the genus. We found the Australian Coptotermes to be monophyletic and most closely related to the Asian Coptotermes, with considerable genetic diversity in some Australian taxa possibly representing undescribed species. The Australian mound-building species did not form a monophyletic clade. Our ancestral state reconstruction analysis indicated that the ancestral Australian Coptotermes was likely to have been a tree nester, and that mound-building behaviour has arisen multiple times. The Australian Coptotermes were found to have diversified ∼13 million years ago, which plausibly matches with the narrowing of the Arafura Sea allowing Asian taxa to cross into Australia. The first diverging Coptotermes group was found to be African, casting doubt on the previously raised hypothesis that the genus has an Asian origin.

Borate protection of softwood from Coptotermes acinaciformis (Isoptera: Rhinotermitidae) damage: Variation in protection thresholds explained

Laboratory and field data reported in the literature are confusing with regard to “adequate” protection thresholds for borate timber preservatives. The confusion is compounded by differences in termite species, timber species and test methodology. Laboratory data indicate a borate retention of 0.5% mass/mass (m/m) boric acid equivalent (BAE) would cause >90% termite mortality and restrict mass loss in test specimens to ≤5%. Field data generally suggest that borate retentions appreciably >0.5% m/m BAE are required. We report two field experiments with varying amounts of untreated feeder material in which Coptotermes acinaciformis (Froggatt) (Isoptera: Rhinotermitidae) responses to borate-treated radiata (Monterey) pine, Pinus radiata D. Don, were measured. The apparently conflicting results between laboratory and field data are explained by the presence or absence of untreated feeder material in the test environment. In the absence of untreated feeder material, wood containing 0.5% BAE provided adequate protection from Coptotermes sp., whereas in the presence of untreated feeder material, increased retentions were required. Furthermore, the retentions required increased with increased amounts of susceptible material present. Some termites, Nasutitermes sp. and Mastotermes darwiniensis Froggatt, for example, are borate-tolerant and borate timber preservatives are not a viable management option with these species. The lack of uniform standards for termite test methodology and assessment criteria for efficacy across the world is recognized as a difficulty with research into the performance of timber preservatives with termites. The many variables in laboratory and field assays make “prescriptive” standards difficult to recommend. The use of “performance” standards to define efficacy criteria (“adequate” protection) is discussed.

Coptotermes gestroi (Isoptera: Rhinotermitidae) in Brazil: possible origins inferred by mitochondrial cytochrome oxidase II gene sequences

The Asian subterranean termite, Coptotermes gestroi, originally from northeast India through Burma, Thailand, Malaysia, and the Indonesian archipelago, is a major termite pest introduced in several countries around the world, including Brazil. We sequenced the mitochondrial COII gene from individuals representing 23 populations. Phylogenetic analysis of COII gene sequences from this and other studies resulted in two main groups: (1) populations of Cleveland (USA) and four populations of Malaysia and (2) populations of Brazil, four populations of Malaysia, and one population from each of Thailand, Puerto Rico, and Key West (USA). Three new localities are reported here, considerably enlarging the distribution of C. gestroi in Brazil: Campo Grande (state of Mato Grosso do Sul), Itajai (state of Santa Catarina), and Porto Alegre (state of Rio Grande do Sul).

Effect of biotic and abiotic factors on the tunneling behavior of Coptotermes gestroi and Heterotermes tenuis (Isoptera : Rhinotermitidae)

Using bidimensional arenas, the construction and spatial dispersion of tunnels constructed by Coptotermes gestroi and Heterotermes tenuis (Rhinotermitidae) was determined under different laboratory conditions. Workers of both species showed an increase of the tunneled area with the rise in temperature. The activity of workers of C gestroi also increased with the rise in soil moisture. Primary tunnels showed a high degree of dispersion in the arenas for all treatments. The presence of food did not influence the tunneling pattern of C. gestroi; but for H. tenuis there was a significant decrease in the number of tunnels. In addition, the type of substrate affected the tunneling pattern in both species. The understanding of food searching strategies of subterranean termites could contribute in the improvement of pest management programs based on the use of toxic baits. (c) 2005 Elsevier B.V. All rights reserved.

Morphology of the sternal gland in workers of Coptotermes gestroi (Isoptera, Rhinotermitidae)

The sternal gland is considered the only source of trail pheromones in termites. The morphology of the sternal gland was investigated in workers of Coptotermes gestroi using transmission and scanning electron microscopy. The results showed a small bilobed gland at the anterior part of the fifth abdominal sternite. The cuticular surface of the sternal gland showed a V-shaped structure with two peg sensilla in elevated socket and various campaniform sensilla. Pores and cuticular scale-like protuberances also occur in the glandular area. The ultrastructure showed a gland composed of class I cells and two different types of class 3 cells distinguished by location, different size and electron-density of secretory vesicles. Small class 3 cells (type 1) of the anterior lobe are inserted among class I cells and have weakly electron-dense vesicles associated with mitochondria, glycogen and smooth endoplasmic reticulum. The class 3 cells (type 2) of posterior lobe showed many round electron-lucent vesicles of secretion, abundant free ribosomes and a well-developed Golgi apparatus. Each class 3 cell is connected to the cuticle by a cuticular duct constituted by the receiving canal and the conducting canal. The secretion of class I cells is stored in an inner subcuticular reservoir that is delimited by the microvilli of these cells. This inner reservoir is large and crossed by the campaniform sensilla and ducts of two types of class 3 cells that open outside of the insect body. An exterior reservoir also is present between the fourth and fifth sternite. The complex structure of the sternal gland suggests multicomponents for the trail pheromone in the worker of C gestroi. (c) 2005 Elsevier Ltd. All rights reserved.