The sternal glands of termites: segmental pattern,phylogenetic implications

Summary In Mastotermitidae, 3 sternal glands are observed on the 3rd, 4th and 5th abdominal segments. All other families only bear one gland, set on the 4th segment in Termopsidae and Hodotermitidae, and on the 5th in Kalotermitidae, Rhinotermitidae, Serritermitidae and Termitidae. This character may be useful for a phylogenetic analysis.     

Morphological phylogenetics of termites (Isoptera)

Isoptera (termites) are an ecologically important order, with both a high abundance and biomass in tropical ecosystems. However, there have been few phylogenetic hypotheses for termites, and we present here the first comprehensive cladistic analysis for the group. We analysed relationships between all seven termite families, including representatives of all known feeding group, plus a number of systematically critical taxa. Termite species richness is biased towards the higher termites (Termitidae), and our taxon sampling reflects this. Our analysis was based essentially on morphological characters (96 workers, 93 soldiers) plus seven biological characters. The cladistic analysis gave four equally parsimonious trees, representing two islands of topologies. The strict consensus tree is fully resolved for the higher termites, but less so for the lower termites. Overall there is low statistical support for the suggested topology, and this can be explained by the high incongruence between the data sets (worker, soldier and biological). This study highlights the particular problems of coding morphological characters in social insects with multiple castes. Without the input of additional data sets, e.g. alates, biological, behavioural and molecular, it will not be possible to obtain a well-supported termite phylogeny.     

Tarsomere and distal tibial glands: Structure and potential roles in termites (Isoptera: Rhinotermitidae,Termitidae)

Social insects have numerous exocrine glands, but these organs are understudied in termites compared to hymenopterans. The tarsomere and distal tibial glands of the termites Heterotermes tenuis, Coptotermes gestroi and Silvestritermes euamignathus were investigated by scanning and transmission electron microscopy. Pore plates are visible in scanning micrographs on the distal tibial surfaces and on the ventral surface of the first and second tarsomeres of workers of H. tenuis and C. gestroi. In contrast, workers of S. euamignathus have isolated pores spread throughout the ventral surfaces of the first, second, and third tarsomeres and the distal tibia. In all three species each pore corresponds to the opening of a class-3 secretory unit, composed of one secretory and one canal cell. Clusters of class-3 glandular cells are arranged side by side underneath the cuticle. The main characteristics of these exocrine glands include their presence on all the legs and the electron-lucent secretion in the secretory cells. Possible functions of these glands are discussed.     

The fine structure of colleterial glands in two cockroaches and three termites, including a detailed study of Cryptocercus punctulatus (Blattaria, Cryptocercidae) and Mastotermes darwiniensis (Isoptera, Mastotermitidae)

The colleterial glands of insects are organs associated with the female genital apparatus. In cockroaches, these glands produce secretions that cover two parallel rows of eggs during oviposition, and in oviparous species, these secretions become the tanned, sculpted, rigid outer casing of the ootheca. The goal of this study was to compare the gross anatomy of the colleterial glands and the ultrastructure of their component tubules in the phylogenetically significant genera Cryptocercus (Blattaria) and Mastotermes (Isoptera). Recent studies indicate that cockroaches in the genus Cryptocercus are the sister group of termites, and Mastotermes is the only termite known to produce a cockroach-like ootheca. One additional oviparous cockroach, Therea, and two additional termites, Zootermopsis and Pseudacanthotermes, were also examined. As in other cockroaches, the colleterial glands of Cryptocercus and Therea are asymmetrical, with a well developed bipartite left gland and a smaller right gland. In the termites Mastotermes, Zootermopsis, and Pseudacanthotermes, the colleterial glands are composed of a well-developed, paired, anterior gland and a small posterior gland; histological staining and cytological evidence suggest that these are homologues of the left and the right colleterial glands of cockroaches, respectively. At the ultrastructural level, colleterial gland tubules are made of cells belonging to a modified class 1 type cell in the cockroaches, in Mastotermes, and in Zootermopsis; the latter lays its eggs singly, without a surrounding ootheca-like structure. In the advanced termite Pseudacanthotermes, the tubules are made of secretory units belonging to the class 3 cell type. This study demonstrates that the cytological characteristics of colleterial glands in basal termites are similar to those of cockroaches, whether the termite secretes an oothecal casing that covers two parallel rows of eggs, as in Mastotermes, or lays its eggs singly, as in Zootermopsis. The function of colleterial glands in non-mastotermitid termites is unknown.     

Cryoprotection in dampwood termites (Termopsidae, Isoptera)

In contrast to the majority of the Order, the dampwood termites of the family Termopsidae found in colder regions can experience frost and snow, either in cool temperate areas at high latitudes (45°), or alpine areas at high elevations (>1000 m). This suggests that dampwood termites are adapted to cold climates. We investigated this hypothesis in two dampwood termites, Porotermes adamsoni Froggatt and Stolotermes victoriensis Hill. We measured nest temperatures and atmospheric temperatures of their alpine habitat during winter, and measured survival and recovery at subzero temperatures. We also determined the minimum temperature at which these species remain active and the LT50 values. We used a novel gas chromatographic strategy to examine eight metabolites from individuals of both species collected in winter and summer to identify possible cryoprotectants. Both P. adamsoni and S. victoriensis had significantly higher levels of trehalose, a known cryoprotectant, in winter than in summer; in addition S. victoriensis also had higher levels of unsaturated fatty acid ligands in winter than in summer, consistent with patterns observed for cold adaptation in other organisms. These results are the first to reveal that dampwood termites are adapted to cold climates and use trehalose and unsaturated lipids as cryoprotectants.     

Arolia in termites (Isoptera): functional significance and evolutionary loss

Summary. Termite workers from all families examined had no arolia (=adhesive pads) on their tarsi and are unable to climb smooth vertical surfaces such as glass or polypropylene plastic. This contrasts with ants where both workers and alates of most species possess arolia and are able to climb these surfaces. Arolia were present in alates of the majority of species investigated from three of the four most basal termite families (Mastotermitidae, Termopsidae and Kalotermitidae), though absent from the basal family Hodotermitidae that contains only three genera. Alates in the two kalotermitid species tested readily climbed glass walls. The complete evolutionary loss of arolia from alates in the specious two most apical termite families (Rhinotermitidae and Termitidae) suggests paedomorphosis. Very smooth surfaces probably cannot be used to completely prevent entry of rhinotermitid termites into buildings because these termites can eventually build galleries of feces and soil over these surfaces. However, an experiment with Coptotermes formosanus showed that a smoother surface significantly slows down the rate of gallery building.Received 12 February 2004; revised 17 June 2004; accepted 29 June 2004.     

两种散白蚁两性生殖与孤雌生殖胚胎发育比较研究

【目的】探究散白蚁两性生殖胚胎和兼性孤雌生殖胚胎各自的发育特点。【方法】以黑胸散白蚁Reticulitermes chinensis和尖唇散白蚁R.aculabialis各自的受精卵(雌雄配对所产的卵)和未受精卵(雌雌配对所产的卵)为研究对象,采用激光共聚焦显微镜观测两种散白蚁的受精卵和未受精卵的卵裂状态,数码显微系统拍照观察卵的外部形态变化。【结果】黑胸散白蚁蚁后所产的受精卵和未受精卵在巢中均能进行卵裂,但是未受精卵在24和48 h时的卵核数无显著性差异,而受精卵卵核数在24和48 h时有显著性差异;15 d时未受精卵的体积没有发生显著变化,而受精卵的体积显著性增大;15-20 d时未受精的蚁卵死亡,而受精卵正常发育。尖唇散白蚁受精卵和未受精卵在相同的时间段内卵核数没有显著性差异,48 h时的卵核数明显比24 h时多;第10天时受精卵长度和宽度发生显著性变化,同时体积也明显增大,而未受精卵在第15天时长度和宽度才开始发生显著性变化,同时体积也开始显著增大。【结论】具有兼性孤雌生殖能力的尖唇散白蚁,受精卵和未受精卵卵裂速度相同,但受精卵外形体积变化比未受精卵早。黑胸散白蚁未受精卵能继续进行卵裂,但发育异常,不能孵化;两种白蚁卵发育过程中卵的长度和宽度同时发生变化。黑胸散白蚁孤雌卵的卵裂特性可能是白蚁两性生殖向兼性孤雌生殖进化的过渡适应阶段。     

Intraspecific interactions in a community of arboreal nesting termites (Isoptera: Termitidae)

In coconut plantations of northern New Guinea, the arboreal nesting termite community comprises three species:Nasutitermes princeps, N. novarumhebridarum, andMicrocerotermes biroi. In orde to assess the importance of intraspecific interactions in this community, we conducted pairwise encounters between batches of individuals in the laboratory and between entire nest populations in seminantural conditions. Three levels of agonism were defined in laboratory bioassays: anagonism, moderate agonism, and strong agonism. Anagonism was observed during all control tests with homocolonical groups and in some tests with allocolonial groups of all species. Moderate agonism included initial aggressiveness that subsequently faded out, and initially passive encounters where aggression progressively built up and led to fighting. Strong agonism corresponded to initial aggressiveness and fighting. Results obtained in alboratory bioassays were consistent with bioassays in seminatural conditions. WhenNasutitermes colonies were anagonists in laboratory bioassays, their foraging trails merged without aggression in field tests.N. princeps nests that were moderately agonistic in laboratory tests fought and either continued to avoid each other or finally joined after elimination of the most aggressive individuals. The most aggressiveM. biroi andN. princeps colonies fought and their foraging trails diverged afterward. Direct attacks on alien nests were winnessed inM. biroi. In all species, anagonism occurred in 21–34% of the combinations tested, between either geographically close or distantcolonies. An exeption was a group of 112 anagonist nests ofN. princeps, which most probably constituted a supercolony. The level of agonism betweenNasutitermes colonies was constant during the wet and dry season. Termite colonies excluded each other, both intra- and interspecifically, from the coconut trees, and their territories seem distributed in a mosaic pattern. Agonism between colonies may result in the elimination of the weakest colonies or in trail divergence, maintaining this mosaic. In cotrast, lack of agonism between some colonies suggests the possibility of colony fusion and gene exchanges without nuptial flights.     

两种鼻白蚁之间的入侵关系

设计了5个实验以研究台湾乳白蚁(Coptotermes formosanus)和黄肢散白蚁(Reticulitermes flavipes)的种间斗争关系。所试白蚁来自阿拉巴马州。实验1将两组白蚁的兵蚁和工蚁以自然比例混合(台湾乳白蚁工蚁：兵蚁＝20：5,黄肢散白蚁工蚁：兵蚁＝24：1);实验2观察了双方同数量(各25头)工蚁的种间斗争能力;实验3观察两者同数量(各5头)兵蚁的种间斗争关系;实验4、5分别研究双方工蚁与兵蚁的斗争关系。因两种工蚁形态相似,所以在实验前用0．1％Nile blue A染过的滤纸饲喂一方白蚁24h使虫体着色以便观察。所有实验在22℃,65％RH荧光灯下进行。每个实验将两方白蚁同时放入一培养皿中,观察至一方100％受伤,然后把培养皿移到培养箱内,24小时后统计死亡率。结果表明,实验一开始,双方的工蚁和兵蚁都立即进入残酷的厮杀斗争,台湾乳白蚁兵蚁和工蚁的斗争能力都明显强于黄肢散白蚁;兵蚁的斗争行为不受工蚁的影响,工蚁的斗争能力也不受兵蚁的制约。实验中台湾乳白蚁的工蚁与黄肢散白蚁的兵蚁攻击能力相当,但黄肢散白蚁的工蚁却远弱于台湾乳白蚁的兵蚁。上述结果说明,自然界中不可能出现此两种白蚁群体的偶然性融合;外来的台湾乳白蚁很可能在其群体建立的地区占据优势[动物学报49(3)：295—302,2003]。     

Hermetically sealed baits for subterranean termites (Isoptera: Rhinotermitidae)

Cellulose baits containing 0.5% hexaflumuron were hermetically sealed in a closed cell polyethylene sheet envelope and placed in soil to test their durability and efficacy against field colonies of the Formosan subterranean termite, Coptotermes formosanus Shiraki, or the eastern subterranean termite, Reticulitermes flavipes (Kollar). The closed cell polyethylene sheet was readily tunneled through by termites, yet it was impervious to water and protected the cellulose baits and hexaflumuron from the environment. Only in a few incidents did the polyethylene envelope become infiltrated by plant roots, resulting in water intrusion and apparent degradation of cellulose baits. After consuming one to three sealed baits, three colonies each of both termite species were eliminated. The sealed baits may be placed in soil for months or years without the need of monitoring, and they are readily penetrated and fed upon by termites when they are present. Application of baits hermetically sealed in a protective sheet may save labor costs by bypassing the monitoring phase, circumvent the station avoidance by some termite species, and enable the use of baiting technologies in large areas such as agricultural fields in which the manual monitoring is impractical.     

The effect of bait design on bait consumption in termites (Isoptera: Rhinotermitidae)

The efficacy of baiting as a pest control method relies on the bait appealing to the pest species. In the case of wood-eating termites, bait stations should be designed to encourage termite presence and to maximize their consumption of bait matrix in order to expedite control in minimal time. A field experiment examined the effect of bait size (one large bait or four small baits of equivalent total size, with commensurate inspection and replacement schedules), compaction (tightly rolled or loosely folded) and composition (paper only or paper plus wood) on termite presence and on untreated bait paper removal rates over four months. All three factors were significant, with bait size the most important factor, followed by compaction and then composition. The least effective baits were small, compacted (rolled) paper-only baits with monthly inspections; these had the highest abandonment rate (70%) and had the least paper removed (mean of 24 g). The most effective baits were large, folded paper-plus-wood baits with inspections at two months; these had the lowest abandonment rate (20%) and had the highest paper removal (mean of 112 g). The more than four-fold difference between these baits types demonstrates that bait efficacy can be altered considerably merely by changing bait design without adding new ingredients to the bait matrix.     

On the respiratory quotient (RQ) of termites (Insecta: Isoptera)

The respiratory quotient (RQ) at 28 degrees C was determined by Warburg manometry in 23 species of termites from the Mbalmayo Forest Reserve (Cameroon) and three sub-tropical species cultured under laboratory conditions in the U.K. or freshly collected in Australia. The data are tabulated with other recently reported RQs (determined by manometry or GC) and with measured CH(4) emission rates to provide a survey of 29 species covering both lower and higher termites in all major trophic (functional) categories. In all species, except the wood-feeding Coptotermes acinaciformis and the soil-feeding Cubitermes fungifaber, the observed mean values (with manometry corrected for known fluxes of H(2) and CH(4)) were at or well above 1.00. Soil-feeding forms (except C. fungifaber) generally showed a high apparent RQ (not corrected for H(2)), with nine species (out of 13) above 1.20 and six species above 1.30. Well-replicated laboratory experiments with Reticulitermes lucifugus showed that there was a tendency for RQ to fall with time over a 4-h incubation, although remaining greater than 1.00.The observed RQs are consistent with carbohydrate being the principal substrate supporting respiration in all trophic and taxonomic categories, with little or no contribution from the degradation of lignin or other polyaromatic materials. However, in many species (especially soil-feeders), the observed RQ is greater than that expected from known fluxes of O(2), CO(2) and CH(4) on the assumption that carbohydrate is the respiratory substrate. This presupposes that there is a large hydrogen sink (additional to CH(4) production), possibly the emission of H(2) gas, and/or the existence of unresolved digestive mechanisms or electron routings. Uncertainties in the use of manometry with termites are discussed.     

Control of subterranean termites (Isoptera: Rhinotermitidae) infesting power poles

A trial was conducted to determine the efficacy of termiticidal dusts (arsenic trioxide, triflumuron, and Metarhizium anisopliae), a timber fumigant (dazomet) and liquid termiticides (bifenthrin, chlorfenapyr, chlorpyrifos, fipronil, and imidacloprid) for controlling subterranean termites (Isoptera: Rhinotermitidae) infesting in-service power poles in New South Wales, Australia. Dusts were applied to parts of the pole where termites were present. Fumigant was inserted into holes drilled into the base of the pole. Liquid termiticides were mixed with soil around the base of the pole and injected into internal voids if present. Poles were inspected for up to 5 yr, and the time taken for reinfestation to occur was recorded. Before the start of the trial, the major Australian pole owners were surveyed to obtain an estimate of the annual national cost of termite infestation to the power supply industry. The annual costs of termite treatment and replacing damaged poles were estimated at AU$2 million and AU$13 million, respectively. Infestation rates were lower for all treatments compared with controls within the first 12 mo of the study. Dazomet, arsenic trioxide, fipronil, and chlorpyrifos were the most efficacious treatments. Efficacy was positively related to the amount of termiticide applied and negatively related to the infestation severity but was unaffected by geographical location. Survival curves were calculated of the time elapsed before the recurrence of termite infestations (survival absence of reinfestation). Survival was highest for poles treated with liquid termiticides.     

Attraction of subterranean termites (Isoptera) to carbon dioxide

Subterranean termites, Reticulitermes spp., were attracted to carbon dioxide (CO2) in laboratory and field tests. In behavioral bioassays, Reticulitermes flavipes (Kollar), Reticulitermes tibialis Banks, and Reticulitermes virginicus Banks were attracted to CO2 concentrations between 5 and 50 mmol/mol. In further bioassays, R. tibialis and R. virginicus were attracted to the headspace from polyisocyanurate construction foam that contained 10-12 mmol/mol CO2. In soil bioassays in the laboratory, more termites foraged in chambers containing CO2-generating formulations than in unbaited control chambers. In field tests, stations containing CO2-generating baits attracted R. tibialis away from wooden fence posts at rangeland sites in Colorado. For all of the CO2 formulations tested, termites foraged in significantly more bait stations at treatment fenceposts than in bait stations at the control fenceposts. By the end of the 8-wk study, the number of bait stations located by termites at treatment fenceposts ranged from 40 to 90%. At control fenceposts, termites foraged in only a single station and the one positive station was not located by termites until week 5 of the study. At treatment fenceposts, termites foraged equally in active stations (containing a CO2-generating bait) and passive stations (with no CO2-generating bait), indicating that bait stations may benefit passively from a proximal CO2 source in the soil. CO2 used as an attractant in current baiting systems could improve their effectiveness by allowing earlier exposure of termites to an insecticide.     

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.     

Ultrastructure of posterior sternal glands of Macrotermes annandalei (Silvestri): new members of the sexual glandular set found in termites (Insecta)

In female alates of Macrotermes annandalei, two types of abdominal glands are involved in the secretion of sex pheromone. Tergal glands are found at the anterior margin of tergites 6-10 and posterior sternal glands (PSGs) are located at the anterior margin of sternites 6-7. The cytological features of both types of glands are quite similar. The fine structural organization of PSGs is studied more precisely and described for the first time. The glandular cuticle is pitted with narrow apertures corresponding to the openings of numerous subcuticular pouches. Several Class 3 glandular units open in each pouch. One canal cell and one secretory cell make an individual glandular unit. The canal cell is enlarged apically and is connected with the other canal cells to form a common pouch. Based on the structural features found in these glands, we propose a common secretory process for PSGs and tergal glands. During the physiological maturation of alates inside the nest, secretory vesicles amass in the cytoplasm of secretory cells, while large intercellular spaces collapse the cuticular pouches. At the time of dispersal flight, pouches are filled with the content of secretory vesicles while intercellular spaces are sharply reduced. After calling behavior, no secretion remains in the glands and pouches collapse again, while secretory cells are drastically reduced in size. The structure and the secretory processes of PSGs and tergal glands are compared to those of abdominal sexual glands known in termites.     

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.     

Latitudinal gradients in the species richness of Australian termites (Isoptera)

Abstract Using data on the geographic range of 260 described species in the Atlas of Australian Termites, seven ‘regions’ with more complete data, across a wide range of latitudes were selected for further analysis. For these regions, mean species richness (± SE) was calculated for (i) all species from all families, (ii) Termitidae (197 spp.), (iii) Amitermes spp. (Termitidae, 58 spp.), (iv) all families excluding Amitermes spp. (139 spp.), (v) Termopsidae (5 spp.), (vi) Kalotermitidae (32 spp.) and (vii) Rhinotermitidae (25 spp.). In addition, we compared the Atlas data with species richness for five regions, across a comparable range of latitudes, based on the pooled species richness of described and un-described species given in community studies. No group of termites showed a consistent decline in species richness from tropical to temperate latitudes for either data set. The Atlas data showed similar total species richness from the tropics to the mediterranean southwest, before declining to lowest species richness at the highest latitudes. Species richness of Amitermes spp. and Rhinotermitidae was highest in the southwest. Termopsidae and Kalotermitidae showed no latitudinal pattern in species richness. Community studies showed highest and lowest total species richness in the southwest and at the highest latitudes (south-coastal Western Australia), respectively, and similar species richness from the tropics to arid central Australia. Species richness of. Amitermes spp. was highest in the southwest (31 spp.). Kalotermitidae and Rhinotermitidae showed no clear latitudinal pattern. The latitudinal patterns of species richness for the Australian termites is consistent with that for the Australian vertebrates and ants in that they differ from patterns established for these taxa on other continents.