Interchange complexes in Italian populations of Reticulitermes lucifugus Rossi (Isoptera: Rhinotermitidae)

In six Italian populations of the rhinotermitid Reticulitermes lucifugus there is a diploid complement of 2n=42 with no visible sex chromosome difference between male and female reproductives. The first male meiotic division in this species is, however, characterised by the presence of inter-change multiples. Thus males from three mainland colonies at Udine, Tombolo and Barberino, as well as a Sicilian colony (Polizzi), are regularly heterozygous for a single translocation chain (C) multiple of IV (2n=19II+ C IV). A fourth mainland colony (Squinzano) proved to be polymorphic. Here, three of the six males analysed regularly formed 21 bivalents; two further males had nineteen bivalents and a chain of four multiple while a sixth male had a predominance of meiocytes with 19 bivalents and a chain of four together with a small number of cells with 18 bivalents and a chain of six multiple. Finally a sixth colony from Sardinia (Sennori) was characterized by larger multiple configurations involving chains or rings of eight.     

Caste differentiation and growth of laboratory colonies of Reticulitermes urbis (Isoptera, Rhinotermitidae)

A total of 280 laboratory colonies of Reticulitermes urbis were raised for 3 years, in order to observe caste differentiation and overall colony growth, and to investigate whether the initial composition (0, 2 or 4 nymphs; 0, 5, 10, 20 or 50 workers) affected colony survival, population growth and other colony dynamics. We also wanted to determine the minimum number of individuals needed to establish a vital colony. As a result, every colony beginning with less than 20 workers died within 60 weeks, while colonies beginning with at least 20 workers survived until the end of the tests, with varied survival rates dependent upon the initial size and the time of formation (summer or autumn). The number of nymphs did not affect colony survival and colony growth. Reproductives developed from nymphs within 2 weeks and later from workers. Many of the reproductives derived from nymphs were killed by workers. The majority of the colonies contained two reproductives 3 years after initiating the tests. In addition, a few individual workers were still alive at the end of the 3-year tests. At the end of the tests, the proportion of soldiers ranged anywhere from 5.23 to 7.69% of the total population. The number of viable “juveniles” from each colony was relatively low throughout the tests and the overall population growth was not sufficient enough to replace dead workers or increase the colony size considerably. These results indicate that the potential to establish viable and sustainable colonies for groups of R. urbis composed of 5–50 workers is very low.     

Septate Gregarines From Reticulitermes Flavipes and Reticulitermes Virginicus (Isoptera: Rhinotermitidae)

ABSTRACT. The gregarine parasites of Reticulitermes virginicus and Reticulitermes flavipes begin their development as trophozoites attached to the midgut epithelium by a small button-shaped epimerite. the epimerite is lost when the parasite becomes free-living in the gut lumen as a solitary gamont. Syzygy is late and was not observed. When full-grown, gamonts enter the hemocoel and fuse in pairs to form large gametocysts that are attached to the midgut of the termite by a duct. Thousands of sporocysts are formed within the original gametocyst. the mature sporocysts are released into the lumen of the midgut through the connecting duct. They are then passed out with the feces. These gregarines are believed to be identical to Gregarina termitis Leidy which was described from a single gamont and later erroneously placed in the genus Hirmocystis by Henry.     

Distribution and genetic variation of Reticulitermes (Isoptera: Rhinotermitidae) in Portugal

The genus Reticulitermes has a Holartic distribution and is widely represented in southern Europe. It is established in Portugal, across the full range of ambient environmental conditions, using different types of wood resources, as available, and nesting ubiquitously in many types of soil. Sequencing of part of the mtDNA COII gene revealed that all the specimens sampled on a north-south transect of the country were Reticulitermes grassei (considered synonymous with R. lucifugus grassei). Nevertheless, an association between the genetic and the geographical distances of sampled specimens was shown, leading to the conclusion that discrete termite populations have differentiated in situ. The geographic component of the genetic polymorphism should be the subject for further research.     

Isolation and characterization of norharmane from Reticulitermes termites (Isoptera: Rhinotermitidae)

The fluorescent alkaloid norharmane has been isolated from Reticulitermes termites and characterized by 1H NMR, UV/Vis, mass spectrometry, and gas chromatography/mass spectrometry. Microcoil 1H NMR spectroscopy allowed spectra to be obtained from mass-limited material, facilitating the identification of norharmane, which is the major component in termite fluorescence under UV light. Norharmane was uniformly present at approximately 1 ng/mg in Reticulitermes tibialis Banks workers, soldiers, and alates; Reticulitermes flavipes (Kollar) workers; and Reticulitermes virginicus (Banks) workers. Some termites were observed to fluoresce with less intensity, but no differences in norharmane levels were detected. Mechanisms that may account for fluorescent differences are discussed as are the possible ecological implications of norharmane in termites.     

Flight Speed of Tethered Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae) Alates

Alates of the Eastern subterranean termite, Reticulitermes flavipes (Kollar) were collected over two flight seasons (2002 and 2004) and flown on flight mills. Data were collected to test if alate mass, colony origin, or gender influenced flight speed. Flight speed ranged from 3.14 to 69.12 cm s⁻¹ and the maximum distance flown by an alate was 458.3 m. Alate mass (P = 0.9406), gender (P = 0.3976), colony origin (P = 0.1244), and the interaction of gender and colony (P = 0.7093) did not significantly influence flight speed. Additionally, an electronic counting device was used to provide instantaneous flight speeds and allowed flight speed to be modeled during acceleration, cruising, and deceleration periods of flight. Mean (±SEM) flight speeds in 2004 were 20.64 (±2.21) cm s⁻¹ (n = 13) for males and 17.76 cm s⁻¹ (n = 1) for the single female flown, falling within the range of the 2002 values.     

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.     

Insecticide susceptibility in Coptotermes formosanus and Reticulitermes virginicus (Isoptera: Rhinotermitidae).

Lethal time to mortality responses were established for eight insecticides against workers and soldiers of the Formosan subterranean termite, Coptotermes formosanus Shiraki, and workers of Reticulitermes virginicus (Banks). There were significant differences in the tolerance ratios between workers of C. formosanus colonies to all toxicants tested except fipronil. One colony was 16 times more tolerant than another to deltamethin. C. formosanus soldiers had significant differences in tolerance ratios among colonies exposed to all toxicants except chlorpyrifos. Methoxychlor, permethrin, deltamethrin, and fipronil did not kill soldiers from two, one, one, and three colonies, respectively, within 8 h. Seventy-five percent of R. virginicus colonies were significantly less susceptible than the most susceptible colony to chlordane, methoxychlor, chlorpyrifos, cypermethrin, and fipronil, with 50% of the colonies less susceptible to permethrin and bendiocarb. In 50% of C. formosanus colonies the worker lethal time curves displayed substantial flattening in response to permethrin, and deltamethrin. Lethal time curses for C. formosanus soldiers exposed to chlordane, chlorpyrifos, permethrin, cypermethrin, deltamethrin, and bendiocarb showed substantial flattening. R. virginicus workers demonstrated substantial curve flattening when exposed to chlordane, methoxychlor, chlorpyrifos, deltamethrin, and fipronil. These findings indicate substantial intercolony and intra-colony differences in susceptibility to insecticides.     

Factors affecting mate choice in the subterranean termite Reticulitermes chinensis (Isoptera: Rhinotermitidae)

Termites are a group of eusocial insects. Mate choice is the most important step which affects the subsequent new colony foundation and development in termites. This study investigated the effects of the four factors on mate choice in the subterranean termite Reticulitermes chinensis Snyder, including colony origin, entry time, physical damage of antennae, and fresh body weights of male dealates. The results showed that the pairing number of dealates from the different colonies was significantly higher than that from the same colonies. The male dealates with early entry time achieved a significantly higher pairing number than the male dealates with late entry time. The male dealates with complete antennae achieved a the significantly higher pairing number than the male dealates lacking the six terminal segments of antennae. Moreover, the heavier male dealates achieved a significantly higher pairing number than the lighter male dealates. However, there were no significant differences in the pairing time for all the experimental treatments. These results suggested that the male dealates, which are heterogenous, healthy, heavy, and more familiar with the environment in R. chinensis, have the obvious advantages in the mate choice. This rule of mate choice is helpful for termites to avoid inbreeding and to maintain the genetic diversity of offspring, which is very important for the environmental adaptability and development of termite colonies.     

Interspecific Interactions Between Coptotermes formosanus and Reticulitermes flavipes (Isoptera: Rhinotermitidae) in Laboratory Bioassays

Experiments were conducted to examine competitive interactions between the Formosan subterranean termite, Coptotermes formosanus Shiraki (FST), and the eastern subterranean termite, Reticulitermes flavipes (Kollar) (EST), using groups of termites with different worker:soldier proportions. Experiments were conducted using three connected test chambers: an FST chamber, an unoccupied center chamber, and an EST chamber. When groups of FST were comprised of 20% soldiers versus 2% EST soldiers, only 8% of center chambers were occupied exclusively by EST. When groups of FST were comprised of 10% soldiers versus 1% EST soldiers, 44% of center chambers were occupied exclusively by EST. When the only food source was located in the center chamber, 60% of center chambers were occupied by both species. FST did not completely displace EST in any of these experiments.     

Cuticular hydrocarbons suggest three lineages in Reticulitermes (Isoptera: Rhinotermitidae) from North America

Cuticular hydrocarbon mixtures can be used to discriminate insect taxa. They have utility for determining phylogenetic relationships where they are independent characters with discrete states and represent a hierarchical distribution of shared, derived characters. We report inferred degrees of relatedness among the chemical phenotypes of Reticulitermes from PAUP (phylogenetic analysis using parsimony) analyses of cuticular hydrocarbon characters. One hundred and forty-one Reticulitermes colonies collected from California, Georgia, New Mexico, Arizona and Nevada were used. Initial maximum parsimony analyses sorted the 141 colonies into 26 chemical phenotypes. Subsequent analyses, using the ancestral species Coptotermes formosanus and Heterotermes sp. as outgroups, sorted Reticulitermes taxa into three major lineages, each characterized by a different set of dominant methyl-branched or unsaturated hydrocarbon components. Reticulitermes in lineage I have cuticular hydrocarbon mixtures with a preponderance of internally branched monomethylalkanes and 11,15-dimethylalkanes. Those in lineage II are defined by a preponderance of 5-methylalkanes and 5,17-dimethylalkanes. Taxa in lineage III are characterized by the predominance of olefins and a relative paucity of n-alkanes and methyl-branched alkanes. Bootstrap analyses and decay indices provided statistical support and robustness for these chemical-based relationships.     

Phylogenetic relationships of nearctic Reticulitermes species (Isoptera: Rhinotermitidae) with particular reference to Reticulitermes arenincola Goellner

DNA sequence comparisons of the mitochondrial COII, 16S, and 12S rRNA genes were used to infer phylogenetic relationships among the six known US Reticulitermes species (Reticulitermes flavipes, Reticulitermes arenincola, Reticulitermes tibialis, Reticulitermes hageni, Reticulitermes virginicus, and Reticulitermes hesperus) and the closely related European species Reticulitermes santonensis. The interspecific pairwise sequence divergence, based on uncorrected "p" distance, varied up to 10% across the COII, 4.9% across the 16S, and 3% across the 12S fragments. Phylogenetic trees were constructed using maximum parsimony, likelihood, and distance methods. The combined results suggest several phylogenetic relationships including: (i) R. flavipes, R. arenincola, and European R. santonensis are possibly conspecific; (ii) R. virginicus and R. hageni are closely related species; and (iii) R. tibialis and R. hesperus are closely related species. Interestingly, while there is apparent synonymity between R. flavipes and R. arenincola by DNA sequence, there are clear morphological differences in the soldier caste. This finding suggests a combination of molecular and morphological approaches are necessary for accurate species identification. These data lend resolution to the complex problem of Reticulitermes systematics, and will assist future efforts directed toward characterizing species distribution and ecology.     

Intraspecific molecular phylogeny, genetic variation and phylogeography of Reticulitermes speratus (Isoptera: Rhinotermitidae)

Population structure was investigated in Reticulitermes speratus populations in the Korean Peninsula and the Japanese Archipelago. All trees derived from analyses of the combined sequence dataset of two mitochondrial genes, COII and COIII, showed that R. speratus populations cluster into two major clades comprising the Korean/southern Japanese populations and the north-ern Japanese populations. Analysis of population ge-netic structure showed strong genetic partitioning between populations of the two clades. To understand historical migration routes and current distributions, the phylogeographic history of R. speratus was inferred from intra-/interspecific phylogeny and diver-gence times estimated between the clades of the phylogenetic tree. The estimated migration route and divergence time of ancestral R. speratus are congruent with recent paleogeographic hypotheses involving land-bridge connections between the Asian continent and the Japanese Archipelago. We suggest that ancestral R. speratus separated into northern and southern Japanese populations after its migration into the Japanese main islands from East China during the early Pleistocene via the East China Sea basin, which may have been exposed during that period. The Korean populations seem to have diverged recently from southern Japanese populations; this may explain the current distribution of R. speratus in the Japanese Arachipelago, and account for why it is restricted to northern areas of the Tokara Strait.     

Feeding inhibition and mortality in Reticulitermes flavipes (Isoptera: Rhinotermitidae) after exposure to imidacloprid-treated soils

Feeding inhibition and mortality of Reticulitermes flavipes (Kollar) exposed to sand, sandy loam, loam, and silty clay loam soils treated with several concentrations of imidacloprid were studied using bioassay techniques under laboratory conditions. Termite workers stopped feeding after exposure to treated soils. Differences in feeding reduction varied among the soil types. Based on the magnitude of the F-statistics, the effect of imidacloprid on the reduction of termite feeding was greatest in sand followed by sandy loam, loam, and silty clay loam soils. Soil properties such as organic matter content, silt and clay proportions, pH, and cation exchange capacity were suggested to affect the bioavailability of imidacloprid. Similar soil effects on mortality were observed in termites continuously exposed to treated soil for 21 d. In three of four soils tested, susceptibility to imidacloprid was not affected by the source of the termites tested.     

Compound eye development during caste differentiation in the termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

We morphologically examined postembryonic compound eye development in Reticulitermes speratus (Kolbe) to understand developmental regulation during caste differentiation. The eye primordia were shown to exist from the larval stage. The number of ommatidia and compound eye size greatly increased over the course of imaginal development. Nymphoids (second-form reproductives) possessed a developed compound eye structure on the surface of the cuticle and thick optic nerves, but individual ommatidia were not clearly discriminated. However, in the line of apterous workers and soldiers, although the outer rim of the eye was observed from second-stage workers, there were few morphological differences among instars, including ergatoids (third-form reproductives). Both nymphoids and ergatoids are slightly physogastric and have highly developed reproductive organs. These results suggest that eye development in the apterous line could be strongly arrested and that there is a weak developmental correlation between the eyes and reproductive organs in R. speratus.     

Preadaptation for parthenogenetic colony foundation in subterranean termites Reticulitermes spp. (Isoptera: Rhinotermitidae)

Thelytokous (all-female producing) parthenogenesis, in some cases, involves reproductive advantages against obligate sexual reproduction. However, the completion of parthenogenesis takes multiple steps without the help of males, and thus preadaptation that meets those requirements will be an important factor for the evolution of parthenogenesis. The Japanese subterranean termite, Reticulitermes speratus, is known to have the ability of parthenogenetic colony foundation, where females that failed to mate with males found colonies cooperatively with partner females and reproduce by parthenogenesis. In this study, we compared the parthenogenetic ability and the colony initiation behavior among six Reticulitermes species in Japan. All species other than R. speratus were not able to reproduce parthenogenetically. Nevertheless, females of these species without the parthenogenetic ability performed homosexual female–female colony initiation and produced eggs without fertilization. In addition, in one species without parthenogenetic reproduction, R. kanmonensis, female–female pair initiated founding behavior as quickly as a heterosexual pair. These results suggest that female–female colony initiation and virgin egg-laying are predominant characters among the genus Reticulitermes and provide a preadaptive condition for parthenogenetic colony foundation in R. speratus.     

Effects of soil cations on the foraging behavior of Reticulitermes flavipes (Isoptera: Rhinotermitidae)

This study examined soil cation concentration gradients around decomposing logs and determined whether Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae) builds galleries in soil substrates containing those cations versus soils without, presumably in an effort to forage efficiently. Soil samples were collected at different depths and lateral distances from decomposing logs containing termites, freshly fallen logs without termites, and bare soil locations at 10 sites to determine whether cation gradients existed in a forest floor. Analyses of cation gradients were significant for potassium, calcium, and magnesium; however, only potassium showed a significant gradient where concentrations decreased with depth and lateral distance from a decomposing log. R. flavipes was then tested in laboratory bioassays to determine if they were attracted to salt cation solutions. Linear foraging distance was significantly greater in artificial K2CO3 gradients than in controls. The results from this study suggest that potassium is found in gradients around decomposing logs and that R. flavipes is attracted to this cation.     

Termite colony ontogeny: a long-term assessment of reproductive lifespan, caste ratios and colony size in Reticulitermes flavipes (Isoptera: Rhinotermitidae)

Thirty Reticulitermes flavipes (Kollar) colonies established by alates collected from two separate field sites were raised in the laboratory for eight years. Twenty-one of the colonies were founded by alates from one field source and nine from another, providing demographic data from two unrelated parental lineages. Colony totals ranged from 3620 to 11641 individuals, with no significant difference in size between lineages. Soldier caste proportion of the colony total and mean wet weights for workers, soldiers and kings were significantly different between the two lineages. This suggests that at least a portion of the variability observed in caste ratios and body size may be heritable. One founding reproductive had died in five of the colonies (17%); none lost both parents. The queenless colonies contained exclusively female replacement reproductives (neotenics); the kingless colony contained a female-skewed mixture of male and female neotenics. All the nests that lost a founding parent contained significantly more pre-alate nymphs than the nests with both a king and a queen. Comparisons with published reports of ontogenetic patterns in other termites and social insects are discussed.     

Effect of fipronil on subterranean termite colonies (Isoptera: Rhinotermitidae) in the field

To assess possible colony-level effects of fipronil, a commonly used nonrepellent termiticide, we conducted a field study of eight houses in the Raleigh, NC, area with infestations of the eastern subterranean termite Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae). We installed an extensive grid of in-ground monitoring stations on each property (mean = 68.6 monitors per property) and collected samples from these stations as well as from mud tubes in the structure and wood debris in the yard for approximately 6 mo. We genotyped all samples by using microsatellite markers to identify the number and locations of colonies present on each property. Houses were treated with either a full treatment (n = 5) or exterior/localized interior treatment (n = 3). After treatment, the monitors were checked monthly for 3 mo and then quarterly for 3 yr to track the fate of colonies. Wood debris in natural areas was checked semiannually for 3 yr. All 11 of the treated colonies (those attacking structures) disappeared within 90 d of treatment and were not found again. These colonies were presumed to be eliminated. In contrast, 60% of untreated colonies (those located > 6 m from the foundation wall at the time of treatment) continued to persist throughout the study, as did 25% of the likely treated colonies (those occupying monitors 0.5 m from the foundation wall where the treatment was applied). Our results provide strong evidence for potent colony wide effects of fipronil on subterranean termites leading to colony suppression and likely colony elimination under field conditions.     

Long-term impact of orphaning on field colonies ofCoptotermes lacteus (Froggatt) (Isoptera: Rhinotermitidae)

Summary Colonies ofCoptotermes lacteus (Froggatt) from a site in coastal south-eastern Australia were experimentally orphaned in early 1989. Sample colonies were examined 3, 6, 12, 18, 24 or 30 months later for their caste composition, the presence of replacement reproductives and brood. All replacement reproductives were nymphoid neotenics. The number of functional (physogastric) females ranged from 1 to 27; this variability was maintained irrespective of the length of time between orphaning and inspection of the colonies. The average live mass of individual females stayed at 30 to 40 mg over the period of 6 to 30 months after orphaning in groups of more than five neotenic queens, but increased from 38 mg three months after orphaning to about 125 mg after 24 months in colonies headed by fewer than five neotenic females. The combined live mass of neotenic females could approach or even exceed that of primary queens. Two key features characterized experimentally and naturally orphaned, neotenic-headed colonies: (1) Nymphs differentiated in significant numbers all year round for a period of at least 30 months right from the time neotenics commenced breeding (in primary-headed colonies nymph production is strictly seasonal). (2) All or most nymphs were males (in primary-headed colonies the sex ratio of nymphs is more or less balanced). The mechanism(s) for achieving the male-biased sex ratio is (are) unknown. Even when colonies have resumed breeding with the help of neotenics, colony survival is not guaranteed. Under such circumstances the gene pool is best preserved if colonies were to raise and release large numbers of alates as potential founders of new colonies. By producing largely male nymphs orphaned colonies ensure outbreeding and may prevent competition (and its disruptive impact an breeding) between existing reproducing neotenic queens and newly differentiating female neotenics. Competition between male neotenics is unlikely to have any impact on the rate of brood production and therefore would not require a mechanism to prevent it from occurring.