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Caste morphology and development in Termitogeton nr. planus (Insecta, Isoptera, Rhinotermitidae).

The termite family Rhinotermitidae displays a wide diversity in its patterns of social organization in castes. The genus Termitogeton probably branched off early in the evolution of this family. We studied the developmental pathways of a putative undescribed species from New Guinea, Termitogeton nr planus. The development begins with two white inactive instars (larvae), the second of which possesses small wing buds. These are followed by a relatively homogenous group of active immature stages (pseudergates) among which a biometric study revealed the presence of four instars. The first of these instars possesses wing buds that regress at subsequent molts. The external morphology of older instars resembles that of higher termite workers. Older pseudergates can differentiate into presoldiers and then into soldiers, but they are also able to molt into a unique stage with long wing pads (nymph), preceding the imago. Colony maturity can be reached with about 10(3) individuals. The sex ratio is near 1:1 in all castes except soldiers, among which females are more numerous than males. The caste pattern of T. nr. planus is reminiscent of those of the Kalotermitidae and Termopsidae, and of Prorhinotermes among the Rhinotermitidae. None of these taxa possess a true worker caste, permanently diverted from imaginal development: social tasks are done by unspecialized immatures that retain a full array of developmental options, including that of proceeding to the imago. The most remarkable trait of T. nr. planus is the presence in all second-instar individuals of wing buds that later regress to reappear in the single nymphal stage. We suggest that the traditional definitions of the terms larvae, nymphs, and pseudergates should be revised because they cannot be satisfactorily applied to the castes of Termitogeton.

Animals↗

Effect of hexaflumuron on mortality of the Western subterranean termite (Isoptera: Rhinotermitidae) during and following exposure and movement of hexaflumuron in termite groups.

The effects of hexaflumuron consumption on the mortality of workers of the Western subterranean termite, Reticulitermes hesperus Banks, were observed following various exposures to a substrate treated with 14C-labeled hexaflumuron at 5 g kg(-1). Uptake of hexaflumuron by workers was rapid, peaking at approximately 280 ng hexaflumuron per termite 12 days after exposure. The onset of mortality began at day 8, with cumulative mortality reaching approximately 96% by day 45. Excretion of hexaflumuron from worker termites was rapid following various exposure periods, half-lives ranging from 2.1 to 4.7 days. Trophallaxis among worker termites was efficient, hexaflumuron levels in recipient termites approaching two-thirds of levels present in donor termites feeding continuously on hexaflumuron-treated filter paper. The effect of donor-to-recipient group ratios was negligible on the amounts of hexaflumuron transferred from donors to recipients. In laboratory tests, hexaflumuron diffused from a feeding source within 7 days. The presence of additional food sources appeared to inhibit movement of hexaflumuron. Movement of hexaflumuron by cannibalism and coprophagy occurred, but was significant only when termites were starved. Hexaflumuron also appeared to adversely affect egg development by preventing hatch. Hexaflumuron was readily transferred through termite groups, effectively suppressing laboratory populations of R hesperus. Variable efficacy in field situations employing baiting with hexaflumuron in southern California may be a consequence of sporadic feeding at bait stations, the rapid clearance of hexaflumuron from individual termites, and the difficulties in bait presentation (low foraging activity, poor bait station foraging fidelity) inherent in the foraging characteristics of the Western subterranean termite.

Animals↗

Transfer of indoxacarb among workers of Coptotermes formosanus (Isoptera: Rhinotermitidae): effects of dose, donor:recipient ratio and post-exposure time.

Horizontal transfer of indoxacarb among workers of the Formosan subterranean termite, Coptotermes formosanus Shiraki, was examined under laboratory conditions. The effects of dose (0, 10, 20, 50, 100 or 200 ng AI per donor), donor:recipient ratio (1:1, 1:4 or 1:9) and post-exposure time (2, 4, 8, 16, 20 and 24 days) on lethal transfer of indoxacarb were investigated using a donor/recipient model in groups of 100 workers. Transfer of lethal doses from donors to recipients was evidenced by significant recipient mortality in 13 out of 15 treatments within 24 days post-exposure. Dosage significantly affected indoxacarb transfer. Higher doses resulted in greater recipient mortality than lower doses. The highest dose tested resulted in 100% death of recipients and donors within 20 days. A dose of 100 ng resulted in recipient mortalities ranging from 68 to 100%, whereas doses < or =50 ng killed <60% of recipients within 24 days. Donor:recipient ratio also had considerable effect on indoxacarb transfer. At 24 days post-exposure, greater recipient mortalities were observed at ratios > or =1:4 at doses > or =20 ng, but only at a ratio 1:1 at dose 10 ng. Recipient mortality increased significantly as post-exposure time increased. The higher the doses applied on donors, the shorter were the times required for the onset of recipient mortality to occur and for recipients to reach maximum mortality. Significantly greater recipient mortalities were not observed at doses 20 and 50 ng until more than 20 days post-exposure, indicating a delayed activity of indoxacarb. Possible transfer mechanisms are discussed.

Animals↗

United States Department of Agriculture-Agriculture Research Service research on targeted management of the Formosan subterranean termite Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae).

The Formosan subterranean termite, Coptotermes formosanus Shiraki is currently one of the most destructive pests in the USA. It is estimated to cost consumers over US dollars 1 billion annually for preventative and remedial treatment and to repair damage caused by this insect. The mission of the Formosan Subterranean Termite Research Unit of the Agricultural Research Service is to demonstrate the most effective existing termite management technologies, integrate them into effective management systems, and provide fundamental problem-solving research for long-term, safe, effective and environmentally friendly new technologies. This article describes the epidemiology of the pest and highlights the research accomplished by the Agricultural Research Service on area-wide management of the termite and fundamental research on its biology that might provide the basis for future management technologies. Fundamental areas that are receiving attention are termite detection, termite colony development, nutrition and foraging, and the search for biological control agents. Other fertile areas include understanding termite symbionts that may provide an additional target for control. Area-wide management of the termite by using population suppression rather than protection of individual structures has been successful; however, much remains to be done to provide long-term sustainable population control. An educational component of the program has provided reliable information to homeowners and pest-control operators that should help slow the spread of this organism and allow rapid intervention in those areas which it infests.

Agriculture↗

Survivorship, tunneling and feeding behaviors of Coptotermes formosanus (Isoptera: Rhinotermitidae) in response to 2'-acetonaphthone-treated sand.

In laboratory tests, 2'-acetonaphthone was incorporated into sand at different concentrations (4.16-200 mg kg(-1)) and evaluated for survivorship, feeding and tunneling effects on the Formosan subterranean termite, Coptotermes formosanus Shiraki. 2'-Acetonaphthone at 8.33 mg kg(-1) sand significantly reduced survivorship (89-94%), tunnel area (68-91%) and food consumption (84-100%) compared with the control. Most of the dead workers were found at the release point and a few had traveled to the surface of the treated sand. Malformed workers (< or =27%) with an unexpected symptom of molting failure were observed, particularly at the lowest concentration tested (4.16 mg kg(-1)). In choice assays, threshold concentration for significant reduction in survivorship, tunnel construction in treated sand and food consumption in the treated-sand side was 8.33mg kg(-1) with termites collected from New Orleans, LA and 35.0 mg kg(-1) for those from Lake Charles, LA. Termites actively fed and tunneled in the untreated sand, whereas 2'-acetonaphthone at 140mg kg(-1) completely inhibited consumption of food placed on the treated sand.

Animals↗

Molecular phylogeny and biogeography of the nasute termite genus Nasutitermes (Isoptera: Termitidae) in the pacific tropics.

The nasute termite genus Nasutitermes is widely distributed over all tropical regions. The phylogenetic relationships among 17 Nasutitermes species from the Pacific tropics were inferred from sequences of mitochondrial cytochrome oxidase II and 16S ribosomal RNA genes. Several methods of analysis yielded phylogenetic trees showing almost the same topology and in good agreement with reconstructions based on morphological or behavioral characters. Neotropical and Australian species came out as separate, apical clades. Asian species split between an apical branch, appearing as sister group to the neotropical clade, and basal taxa. New Guinean species were spread among several clades, suggesting a derivation from multiple origins. A well-supported clade includes the neotropical, Australian, and New Guinean species, with the southeast Asian N. takasagoensis and N. matangensis. It excludes the Asian species N. regularis, N. parvonasutus, and N. longinasus, which might deserve to be removed from Nasutitermes, as well as the long-legged Asian genera Hospitalitermes and Longipeditermes. A Gondwanan origin is proposed for the former clade, although an Old World origin of Nasutitermes followed by dispersal to Australia and South America cannot be excluded.

Animals↗

Phylogenetic analyses of two mitochondrial genes and one nuclear intron region illuminate European subterranean termite (Isoptera: Rhinotermitidae) gene flow, taxonomy, and introduction dynamics.

Phylogenetic analyses of multiple DNA sequences were conducted to elucidate gene flow, evolutionary patterns, taxonomy, and the dynamics of two accidental introductions: Reticulitermes lucifugus grassei into Devon, United Kingdom and R. flavipes into Europe. Two mitochondrial DNA genes totaling 1495 bp and a 380-bp ribosomal intergenic transcribed spacer were sequenced. Neighbor-joining and parsimony analyses revealed that multiple female lineages of R. lucifugus grassei were introduced into Devon possibly from southwestern France, where the species was indigenous. The taxonomic status of the European R. santonensis as a species separate from the North American R. flavipes has been questioned since it was described in 1924. Phylogenetic analyses revealed a close genetic relationship between R. flavipes from the United States and R. santonensis from France. These analyses, coupled with morphological and chemotaxonomic data, provide strong support for R. santonensis and R. flavipes being the same species. They also suggested that R. santonensis infestations likely resulted from R. flavipes being introduced into Europe.

Animals↗

Molecular relationships among European samples of Reticulitermes (Isoptera, Rhinotermitidae).

Taxonomy and phyletic relationships of the European termites of the holarctic genus Reticulitermes are highly debated and poorly known. Sequencing analyses of the complete cytochrome oxidase subunit II and of a fragment of 16S rDNA mitochondrial genes were performed on 21 Italian and French populations. Distance, parsimony, and maximum-likelihood evaluations on single and combined data sets confirm the presence in central peninsular Italy of R. lucifugus lucifugus, but demonstrate that a coastal Tuscan population and the Sardinian ones pertain to the Corsican subspecies R. lucifugus corsicus, whose distribution appears therefore transtyrrenian. Northeastern Italian samples are highly differentiated from presently analyzed R. lucifugus and constitute a new entity, possibly more related to the Japanese R. speratus. Finally, the French R. santonensis and the North American R. flavipes appear strongly related, so that the supposed synonymy between the two taxa must be accepted.

Animals↗

Mandibular morphogenesis during soldier differentiation in the damp-wood termite Hodotermopsis sjoestedti (Isoptera: Termopsidae).

The conspicuous morphogenesis during termite soldier differentiation is one of the most remarkable examples of specialized caste development in social insects. To clarify the mechanism of morphological changes during soldier differentiation, mandibular morphogenesis prior to the presoldier moult was examined in the damp-wood termite Hodotermopsis sjoestedti. Using experimental induction of presoldier differentiation from pseudergates (7th or later instar larvae) by the application of juvenile hormone analogue (JHA), we compared mandibular morphogenesis in a presoldier moult (from pseudergate to presoldier) with that of a stationary moult (from pseudergate to pseudergate). Future presoldier mandibles were formed under the old mandibular cuticle of the pseudergates just prior to the presoldier moult. Multiple folds of both epidermis and new cuticle were observed in the developing mandible, and large concavities between teeth were formed on the mandibular surface. While in the stationary moult, the future mandibles were formed with a smooth surface. The process of the mandibular morphogenesis in soldier differentiation explains the allometry of soldier mandibles. The morphological differences in mouth parts between castes arise from a switching mechanism that triggers caste-specific mandibular epithelial growth during development.

Animals↗

The first fossil fungus gardens of Isoptera: oldest evidence of symbiotic termite fungiculture (Miocene, Chad basin).

Higher termites of the subfamily Macrotermitinae (fungus-growing termites) are known to build fungus gardens where a symbiotic fungus (Termitomyces sp.) is cultivated. The fungus grows on a substrate called fungus comb, a structure built with the termites' own faeces. Here we present the first fossil fungus combs ever found in the world. They were extracted from 7-million-year-old continental sandstone (Chad basin). Fossilized fungus combs have an ovoid morphology with a more or less flattened concave base and a characteristic general alveolar aspect. Under lens, they display a typical millimetre-scale pelletal structure. The latter, as well as the general shape and alveolar aspect, are similar to the morphology of fungus combs from extant fungus-growing termites.

Animals↗

Discovery of a novel Wolbachia super group in Isoptera.

Wolbachia are one of the most abundant groups of bacterial endosymbionts in the biosphere. Interest in these heritable microbes has expanded with the discovery of wider genetic diversity in under-sampled host species. Here, we report on the putative discovery of a new genetic lineage, denoted super group H, which infects the Isopteran species Zootermopsis angusticollis and Z. nevadensis. Evidence for this novel super group is based on portions of new Wolbachia gene sequences from each species spanning 3.5 kilobases of DNA and the following genes: 16S rDNA, dnaA, gltA, groEL, and ftsZ. Single-gene and concatenated maximum likelihood phylogenies establish this new super group and validate the positioning of the other Wolbachia super groups. This discovery is the first example of a termite Wolbachia that is highly divergent from the Isopteran Wolbachia previously described in super group F. This study highlights the importance of multilocus approaches to resolving Wolbachia super group relationships. It also suggests that surveys of Wolbachia in more earlier-originating (and under-sampled) groups of arthropods are more apt to reveal novel genetic diversity.

Animals↗

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.

Animals↗

Identification of soldier caste-specific protein in the frontal gland of nasute termite Nasutitermes takasagoensis (Isoptera: Termitidae).

The termite soldier is unique because of its defensive task in a colony. In Nasutitermitinae (family Termitidae), soldiers use in their defense frontal glands, which contain various chemical substances. To isolate the gene products related to the chemical defense, we compared the sodium dodecyl sulfate-polyacrylamide gel electrophoresis protein profiles of soldier heads with those of workers of the nasute termite Nasutitermes takasagoensis. We identified a 26-kDa soldier-specific protein (Ntsp1) that exists most abundantly in the dorsal head including the frontal gland. We determined the N-terminal amino acid sequence of Ntsp1, and then cloned the Ntsp1 cDNA by rapid amplification of the cDNA ends-polymerase chain reaction (RACE-PCR). A putative signal peptide was detected upstream of the N-terminus and the Ntsp1 protein showed sequence homologies with known insect secretory carrier proteins, which bind to hydrophobic ligands such as juvenile hormone, suggesting that Ntsp1 belongs to this class of proteins. Northern blot analysis confirmed that the expression level of Ntsp1 was high only in the soldier head. In addition, the localization of Ntsp1 expression was limited in epithelial cells of the frontal gland reservoir, suggesting that this protein binds to some terpenoid(s) preserved in the frontal gland reservoir.

Amino Acid Sequence↗

Factors affecting post-flight behavior in primary reproductives of the Formosan subterranean termite, Coptotermes formosanus (Isoptera: Rhinotermitidae).

After swarming, reproductive dealates of the Formosan subterranean termite, Coptotermes formosanus, run together in tandem. The tandem running is an age-related behavioral activity in C. formosanus. This behavior was not evoked in pre-flight alates less than 6-d-old that were artificially dealated. Female age was more important than male age for evoking tandem behavior. Females and males, older than 35 days, did not exhibit this behavior. Mating status was not important for female and male dealates to form the tandem pairs. The titers of the major tergal gland component, trilinolein, did not decline significantly and remained high, not only in virgin females, but also in mated females for a period of time after swarming. On the other hand, increasing amounts of the female-specific compound trilinolein were detected in male dealates 7, 14, and 42 days after pairing. This suggests that trilinolein in the females might be transferred to the males as a nuptial gift in C. formosanus. In addition, females have structurally different lateral setae that may constitute a morphological factor involved in the tandem behavior in this species. Covering the setae with dimethyl sulfoxide prevented the tandem behavior.

Age Factors↗

Juvenile hormone III titers and regulation of soldier caste in coptotermes formosanus (Isoptera: Rhinotermitidae).

Juvenile hormone (JH) is an important growth hormone in insects that has also been implicated in caste determination in termites. Gas chromatography-mass spectrometry was used to establish that the JH in the Formosan subterranean termite, Coptotermes formosanus Shiraki, is JH III. JH III titers were measured in workers, pre-soldiers, and soldiers from samples collected from the field. The average titers of JH III in workers and soldiers were about 13 and 25 pg mg(-1), respectively. However, pre-soldiers contained a significantly higher amount, 596 pg mg(-1). As expected, treatment of workers with a JH-analogue, methoprene, triggered rapid formation of pre-soldiers. However, these pre-soldiers had a very low JH III titer (62 pg mg(-1)). It appears that the application of JHA, while inducing pre-soldier formation, does not increase the endogenous JH III titer. The titer, however, increased as the pre-soldiers aged and before transforming into soldiers.

Animals↗

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 1 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 1 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 1 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.

Animals↗

Fat body of Prorhinotermes simplex (Isoptera: Rhinotermitidae): ultrastructure, inter-caste differences and lipid composition.

Ultrastructure of the fat body was studied in following castes and developmental stages of Prorhinotermes simplex: larvae of the first and the second instar, pseudergates, presoldiers, soldiers, nymphs, imagoes and mature ergatoid neotenic reproductives of both sexes. Fat body always consists of two principal cell types: adipocytes and urocytes. Adipocytes are characterized by a presence of large amounts of storage substances, namely lipid droplets, glycogen rosettes and proteins in the form of either biocrystals or vacuoles. Proportion of these components strongly varies during ontogeny. Adipocytes are equipped by a large central vacuole in which lipid droplets may resolve. Cytoplasm of urocytes contain glycogen rosettes and spherical or irregular concretions, other organelles are rare. Only adipocytes change their inner structure in the course of ontogeny: amount of glycogen decreases during the postembryonic development, it is the major kind of inclusion in the larvae but lacks in nymphs and imagoes; opposite trend is performed by lipids. The changes in protein content are less obvious but are explained and discussed. The total amount of triacylglycerols (TAGs) was found to be roughly 100microg in a pseudergate, 250microg in a nymph, and 30microg in a soldier. The most abundant fatty acids in TAGs are oleic (O), stearic (S), palmitic (P) and linoleic (L) acid. TAGs form a complex mixture with OOO, OPO, OLO and OOS being the most abundant isomers. Only negligible differences exist among the castes.

Adipocytes↗