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Phylogeography of an Australian termite, Amitermes laurensis (Isoptera, Termitidae), with special reference to the variety of mound shapes.

In northern Australia, the debris-feeding termite Amitermes laurensis builds tall, wedge-shaped mounds in the northern part of Cape York Peninsula and Arnhem Land, where their habitats are seasonally flooded, and small dome shaped mounds in the southeastern part of Cape York Peninsula, where their habitats are well-drained. Phylogeographic analyses were conducted in 238 individuals from 30 populations using the mitochondrial cytochrome oxidase II (COII) gene. DNA sequences of 50 haplotypes were used to construct NJ, MP and ML trees. Phylogenetic trees for 16 Amitermes species showed monophyly of A. laurensis and the variation of A. laurensis mounds did not strongly correspond to the intraspecific phylogeny. It was observed that mounds with the same shape were constructed by phylogenetically different groups under similar environmental conditions and different mounds shapes were built by phylogenetically closely related groups under the different environmental conditions. Thus, phylogenetically close groups of A. laurensis, in different habitats, may adapt to environmental conditions by constructing different mound shapes. We also investigated the phylogeographic structure of A. laurensis. The significant positive correlation between genetic and geographic distances indicated isolation by distance, reflecting restricted dispersal ability of alates. Although the overall genetic structure of A. laurensis showed isolation by distance, we also identified two exceptions: (i) secondary contacts of genetically divergent lineages in southern Cape York Peninsula, and (ii) low genetic differences between geographically separated populations of Cape York Peninsula and Arnhem Land. Therefore, the phylogeography of A. laurensis may reflect continuous gene flow restricted to short distances and past changes of gene flow associated with the fluctuation of environmental conditions accompanying the changing sea levels in the Quaternary.

Animals↗

Energy consumption of termite colonies of Nasutitermes ephratae (Isoptera: Termitidae).

Measurements of CO2 production of whole termite colonies showed that respiration intensity varies slightly during the daily cycle, that the relation between standard metabolism and colony size is best described with a negative exponential, taking into account the proportion of the various castes. Larger colonies were more efficient in their energy use, suggesting that energetic considerations may contribute in explaining the maintenance in evolution of complex societies.

Adaptation, Biological↗

Origin of a new Reticulitermes termite (Isoptera, Rhinotermitidae) inferred from mitochondrial and nuclear DNA data.

The Holoarctic termite genus Reticulitermes is widely distributed in Europe. A new Reticulitermes species, R. sp. nov, was recently found in France and Italy. Its phylogenetic position was investigated using a 743-bp fragment of mitochondrial 16S rRNA-ND1 genes and 382-bp of the nuclear ITS2 region. Phylogenies for these sequences were estimated by neighbor-joining, maximum-parsimony and maximum-likelihood analysis. The results strongly supported a relationship between R. sp. nov. and the termite species from the eastern Mediterranean area including Reticulitermes balkanensis from the Balkans, Reticulitermes lucifugus from Turkey and Reticulitermes clypeatus from Israel. The hypothesis of a relationship between R. sp. nov. and the Japanese Reticulitermes speratus was rejected by parametric bootstrap. The current distribution of R. sp. nov. could be linked to postglacial colonization routes between Balkan refuge and northern regions.

Animals↗

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.

Animals↗

Cyclic CO(2) release in Cryptotermes cavifrons Banks, Incisitermes tabogae (Snyder) and I. minor (Hagen) (Isoptera: Kalotermitidae).

CO(2) release patterns of three drywood termite species were investigated using flow-through respirometry techniques. Eight hours of real-time CO(2) release data were recorded for pseudergates of Cryptotermes cavifrons Banks, Incisitermes minor (Hagen), and I. tabogae (Snyder) at 20-40 degrees C. Cyclic release of CO(2) was observed in 20-90% of C. cavifrons, 70-100% of I. tabogae, and 87-100% of I. minor pseudergates. Variability of the recordings (calculated as the coefficient of variability or CV) was used to estimate the level of cycling in each recording. CV ranged from 14.53+/-2.57 (40 degrees C) to 32.33+/-1.12% (30 degrees C) in C. cavifrons, 20.24+/-2.44 (35 degrees C) to 67.3+/-10.3% (20 degrees C) in I. minor, and 15.9+/-1.46 (35 degrees C) to 34.15+/-6.18% (20 degrees C) in I. tabogae. The relationship between temperature and CV for each species was modeled using non-linear regression. CV of both Incisitermes spp. decreased exponentially with temperature, while C. cavifrons CV followed a Gaussian model, indicating an optimal cycling temperature of approximately 30 degrees C. Mean V.CO(2) values were determined for each species as a function of temperature, and ranged from 0.1 ml CO(2) g(-1) h(-1) (I. minor at 20 degrees C) to 0.8 ml CO(2) g(-1) h(-1) (C. cavifrons at 40 degrees C). For all three species, V.CO(2) significantly increased linearly with temperature. Colinearity tests indicated that different models described the V.CO(2) relationship with temperature for both genera. Q(10) values for V.CO(2) over the range of 20-40 degrees C were 1.92 for I. minor, 1.66 for I. tabogae, and 1.62 for C. cavifrons pseudergates.

Adaptation, Physiological↗

Cuticular permeability of two species of Coptotermes Wasmann (Isoptera: Rhinotermitidae).

Cuticular permeability (CP) values of worker and soldier castes of Coptotermes formosanus Shiraki and Coptotermes vastator Light were determined using gravimetric techniques. Comparisons were made between castes, species, and between colonies of C. formosanus and C. vastator. CP values did not differ among either species or colony, however they did differ between castes. Data reported here and that of established data in the literature for C. formosanus are inconsistent, with the current report having values as small as half that of previously published data. Examining mean CP data by colony indicated that there is a range for Coptotermes spp. termites, from 6.84+/-1.2 to 25.61+/-1.49 microg H(2)Ocm(-2)mmHg(-1)h(-1) dependent upon caste. A novel method of examining intercaste CP variability is the use of worker:soldier CP ratios. Ratios for C. formosanus were consistent between the current study and previous reports. Mean percentage total body water (%TBW) lost at the time of death was 33.75+/-1.15% (soldiers) and 54.24+/-1.43% (workers) for C. formosanus, and 37.69+/-3.31% (soldiers) and 52.87+/-3.65% (workers) for C. vastator. Consistently, %TBW (and fresh mass) was greater in worker than in soldier castes. These data suggest the use of water storage mechanisms in worker termites, as shown for other rhinotermitids.

Animals↗

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.

Animals↗

Toxicity and repellency of patchouli oil and patchouli alcohol against Formosan subterranean termites Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae).

Patchouli oil obtained from Pogostemon cablin (Blanco) Benth and its main constituent, patchouli alcohol, were tested for their repellency and toxicity against Formosan subterranean termites (Coptotermes formosanus Shiraki). Both were found to be toxic and repellent. Unusual tissue destruction was noted inside the exoskeleton of the termite after patchouli alcohol was topically applied to the dorsum.

Animals↗

Food exploitation in termites: indication for a general feeding-stimulating signal in labial gland secretion of isoptera.

The paired labial glands are located in all termite species in the thorax. During food exploitation workers of the French termite Reticulitermes santonensis and the African termite Schedorhinotermes lamanianus release the secretion of their labial glands directly onto the food. The secretion carries a water-soluble, heat-resistant, nonvolatile signal that stimulates gnawing and feeding and leads to aggregations of feeding workers. In a feeding bioassay, extracts of the labial glands of 11 termite species from five families all proved to have this feeding-stimulating effect both on R. santonensis and S. lamanianus. The heat resistance of the feeding-stimulating signal also could be shown for selected species from all termite families tested. A combined thin layer chromatography-feeding bioassay on cellulose TLC plates showed that after chromatography of labial gland secretion, the feeding-stimulating signal is located in all 11 species in the same area from Rf 0.46 to 0.88. An extract of labial glands of cockroaches stimulated feeding in R. santonensis and S. lamanianus as well, but was not active after heat treatment and after TLC. This points towards a general feeding-stimulating signal having evolved only in the labial gland secretion of termites.

Animals↗

Characterization of Nasutitermes globiceps (Isoptera: Termitidae) esterases.

Esterases of Nasutitermes globiceps termites which occur on the Upper Paraná River floodplain (Brazil) were characterized. The electrophoretic pattern of the termite esterases Nasutitermes globiceps was obtained by starch gel electrophoresis. Six esterase activity zones were obtained and numbered, with esterase-1 being the most anodall one and esterase-6 the most cathodal one. Esterase-2 was detected only with substrates derived from the 4-methylumbelliferyl radical. The esterases of N. globiceps present wide substrate specificity, having been observed with substrates derived from alpha-naphthyl (acetate, propionate, and butyrate) and beta-naphthyl (acetate, butyrate) and from 4-methylumbelliferyl (acetate, propionate and butyrate). Esterase-6 is a caste-specific enzyme detected in soldiers. Only esterases 1, 3 and 5 were detected in nymphs. No genetic polymorphism has been detected thus far in the esterases of Nasutitermes globiceps. This study suggests that allozyme variation can be explored to understand Nasutitermes social structure.

Animals↗

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.

Animals↗

Genetic variation of symbiotic fungi cultivated by the macrotermitine termite Odontotermes formosanus (Isoptera: Termitidae) in the Ryukyu Archipelago.

Fungus-growing termites have a mutualistic relationship with their cultivated fungi. To improve understanding of genetic aspects of this relationship, we examined molecular markers in the fungus-growing termite Odontotermes formosanus and its fungi Termitomyces spp. from the Ryukyu Archipelago. Based on the polymorphic band patterns obtained from arbitrarily primed polymerase chain reaction methods, we constructed cladograms for related colonies of the termites and fungi. The resulting trees indicated that the termites display little genetic variation among the colonies, while the symbiotic fungi consist of two major genetic types. In addition, molecular phylogenetic trees of the symbiotic fungi based on internal transcribed spacer and 18S rDNA suggested that these two types of fungi are different species. We also demonstrated that the fungi comprising the fruiting bodies and fungus combs are identical, and that fungus combs are probably a monoculture within a single termite colony. Our results indicate that horizontal transmission of symbiotic fungi among termite colonies occurred during the evolutionary history of this symbiosis.

Animals↗

Soldier caste-specific gene expression in the mandibular glands of Hodotermopsis japonica (Isoptera: termopsidae).

Although "polymorphic castes" in social insects are well known as one of the most important phenomena of polyphenism, few studies of caste-specific gene expressions have been performed in social insects. To identify genes specifically expressed in the soldier caste of the Japanese damp-wood termite Hodotermopsis japonica, we employed the differential-display method using oligo(dT) and arbitrary primers, compared mRNA from the heads of mature soldiers and pseudergates (worker caste), and identified a clone (PCR product) 329 bp in length termed SOL1. Northern blot analysis showed that the SOL1 mRNA is about 1.0 kb in length and is expressed specifically in mature soldiers, but not in pseudergates, even in the presoldier induction by juvenile hormone analogue, suggesting that the product is specific for terminally differentiated soldiers. By using the method of 5'- and 3'-rapid amplification of cDNA ends, we isolated the full length of SOL1 cDNA, which contained an ORF with a putative signal peptide at the N terminus. The sequence showed no significant homology with any other known protein sequences. In situ hybridization analysis showed that SOL1 is expressed specifically in the mandibular glands. These results strongly suggest that the SOL1 gene encodes a secretory protein specifically synthesized in the mandibular glands of the soldiers. Histological observations revealed that the gland actually develops during the differentiation into the soldier caste.

Amino Acid Sequence↗

Tunnel specificity and forager movement in subterranean termites (Isoptera: Rhinotermitidae and Termitidae).

The movement of foragers of two species of Australian, subterranean, mound-building termites, Coptotermes lacteus (Froggatt) (Rhinotermitidae) and Nasutitermes exitiosus (Hill) (Termitidae), was investigated in their natural habitat using artificial feeding sites along trenches dug to mimic natural forager tunnels that radiate out from the central mound-nests. Termites were dyed by self-feeding on cardboard soaked with histological fat-stains on one or two trenches and then termites were collected from other feeding sites at two and four weeks after the fat-stains were placed. At two and four weeks after marking commenced, 60-75% of marked termites were found in trenches containing the marked paper, and 2-16% were found in trenches on the opposite side of the nest. The proportion of marked termites in a sample was three to eight times greater in the trenches containing the marked paper relative to other trenches. Although difficulties with fat-stains used as markers might explain some of the observed patterns, it is evident that C. lacteus and N. exitiosus foragers do not move randomly between feeding sites in their natural habitat.

Animals↗

Feeding site selection by workers of the Formosan subterranean termite Coptotermes formosanus (Isoptera: Rhinotermitidae) - a re-analysis of field data from a mark-recapture study.

During the early stages of the development of termite baits, dyed paper was placed in specified feeding stations to ascertain whether a slow-acting toxicant could be placed in a few bait stations to be delivered to the entire colony members of the Formosan subterranean termite, Coptotermes formosanus Shiraki. Feeding frequency data, as measured by the dye concentration in individual termites, suggested the absence of feeding site fidelity. However, these results were often misinterpreted as random movement of termites that were marked and released for population estimate studies, or the random search of food in soil by subterranean termites. A computer simulation program was constructed to re-examine this feeding frequency data, and confirmed the earlier conclusion that the lack of feeding site fidelity was the most likely explanation for the data.

Animals↗