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.     

Parthenogenetic reproduction in neotenics of the subterranean termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

Experiments with artificial colonies consisting of only female workers and those of female nymphs and workers of a subterranean termite, Reticulitermes speratus, showed that neotenics (ergatoids and nymphoids) can reproduce parthenogenetically. The larvae hatched from the eggs produced partheno‐genetically by the neotenics were all female and did not have any aberrant external morphology. They had 42 chromosomes and were diploid. The analysis of genotypes at microsatellite loci revealed that the eggs produced parthenogenetically had only one of the two alleles that the mother neotenics possessed, indicating that the eggs were produced by automixis.     

Polymorphism in the giant cocoa termite,Neotermes papua (Desneux)

Summary The direct development ofNeotermes papua (Isoptera: Kalotermitidae) comprises four larval and three nymphal instars before the alate. The first five instars can be easily characterized. The second stage nymphs come morphologically close to the pseudergates, characterized by reduced wing buds. These nymphs can moult stationarily, i.e. with little morphological change, or to presoldiers, or proceed to the alate via the third nymphal stage. Pseudergates originate through a late and reversible deviation from the straight development to the alate. Presoldiers may derive from several stages, up to the last nymphal one; their production is subject to an inhibition by extant soldiers. This developmental schema is congruent with those described in other Kalotermitidae and the Termopsidae. By pinpointing the existence of a large pool of pluripotent individuals, in which the penultimate nymphal stage mingles with pseudergates, the present study also reveals a great similarity betweenNeotermes andProrhinotermes, and suggests that this developmental schema might be generally applicable to termites devoid of a permanent worker caste.King Léopold III Biological Station, Laing Island, Papua New Guinea, contribution No. 235     

Diazotrophs in the digestive tract of termite Neotermes castaneus

Normal vital activity of termites Neotermes castaneus requires the presence and continuous replenishment of transient nitrogen-fixing bacteria in their digestive tract, which is realized by coprophagy and repeated utilization of substrate enriched in termite feces. This is the first demonstration of significant changes in the complex of aerobic and facultatively anaerobic nitrogen fixers mediated by an extension of microbial group composition and a shift in dominant taxa in termites that cannot reutilize their feces.     

Revision of the termite family Rhinotermitidae (Isoptera) in New Guinea

Recently, we completed a revision of the Termitidae from New Guinea and neighboring islands, recording a total of 45 species. Here, we revise a second family, the Rhinotermitidae, to progress towards a full picture of the termite diversity in New Guinea. Altogether, 6 genera and 15 species are recorded, among which two species, Coptotermes gambrinus and Parrhinotermes barbatus, are new to science. The genus Heterotermes is reported from New Guinea for the first time, with two species restricted to the southern part of the island. We also provide the first New Guinea records for six species of the genera Coptotermes and Schedorhinotermes. We briefly describe soldiers and imagoes of each species and provide a key based on soldier characters. Finally, we discuss the taxonomic and biogeographical implication of our results. A replacement name, Schedolimulus minutides Bourguignon, is proposed for the termitophilous staphylinid Schedolimulus minutus Bourguignon, to solve a question of secondary homonymy.     

Divergence times in the termite genus Macrotermes (Isoptera: Termitidae)

The evolution of fungus-growing termites is supposed to have started in the African rain forests with multiple invasions of semi-arid habitats as well as multiple invasions of the Oriental region. We used sequences of the mitochondrial COII gene and Bayesian dating to investigate the time frame of the evolution of Macrotermes, an important genus of fungus-growing termites. We found that the genus Macrotermes consists of at least 6 distantly related clades. Furthermore, the COII sequences suggested some cryptic diversity within the analysed African Macrotermes species. The dates calculated with the COII data using a fossilized termite mound to calibrate the clock were in good agreement with dates calculated with COI sequences using the split between Locusta and Chortippus as calibration point which supports the consistency of the calibration points. The clades from the Oriental region dated back to the early Tertiary. These estimates of divergence times suggested that Macrotermes invaded Asia during periods with humid climates. For Africa, many speciation events predated the Pleistocene and fall in range of 6-23 million years ago. These estimates suggest that savannah-adapted African clades radiated with the spread of the semi-arid ecosystems during the Miocene. Apparently, events during the Pleistocene were of little importance for speciation within the genus Macrotermes. However, further investigations are necessary to increase the number of taxa for phylogenetic analysis.     

Morphometric changes during soldier differentiation of the damp-wood termite Hodotermopsis japonica (Isoptera, Termopsidae)

Summary. To clarify the allometric development of body parts accompanying soldier differentiation in termites, we measured 16 body parts of soldiers, presoldiers, pseudergates (workers), nymphs and larvae of the damp-wood termite Hodotermopsis japonica. Principal component analysis (PCA) was performed using these parameters, which revealed that differentiation into soldiers differed distinctly from development into adult (reproductive) individuals. In particular, the anterior body parts enlarged during development of soldiers. Similarly, elongation of the apical portion of both mandibles was noted during soldier differentiation. X-ray analysis of mandibles revealed sclerotization of the soldier mandibles after differentiation into terminal soldiers. These morphological changes during soldier differentiation are associated with changes in their roles within the colony. Through soldier differentiation, the morphology of this caste of termite becomes functionally suited for attacking predators, and unsuitable for feeding on wood using their mandibles. Based on these data, we suggest that there must be some morphogenetic factors leading caste specific morphology such as soldier mandibles.     

Distribution of Formosan subterranean termite (Isoptera: Rhinotermitidae) in Mississippi

An extensive monitoring and survey program in Mississippi was conducted from 2000 to 2004 to investigate the distribution of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae). Seventy-two towns from 22 counties in southern Mississippi were monitored with a total of 3914 traps that catch alates during the swarming season. In addition, 259 licensed pest management professionals in Mississippi were surveyed to determine the locations of termite infestations treated. The alates of C. formosanus were recovered in 12 counties with light traps, and termite infestations were documented in an additional 13 from data collected in the termite survey. Infestations of C. formosanus have been documented in urban, urban cluster, rural, and forested areas of Mississippi. However, the distribution in mean total capture of alates for 4 yr differed significantly among the four ecological areas with the highest percentage in forested areas (31%) and the lowest percentage in urban cluster areas (17%). Most of the infestations of C. formosanus were geographically distributed along the coastal areas of southern Mississippi from Gulfport to Pascagoula. The greatest total number of alates captured in light traps was documented in Pearl River County. Mass swarming of C. formosanus occurred primarily in May or June, depending on weather conditions. The number of documented counties with the evidence of large and widely dispersed swarms of C. formosanus in different ecological areas, and the increase in total annual alate captures from 2000 to 2003, suggest that this invasive termite species is now firmly established in Mississippi.     

Soldier-like intercastes in the rotten-wood termite Hodotermopsis sjostedti (Isoptera: Termopsidae)

All termite species possess a distinct sterile-soldier caste in their colonies, although reproductive soldiers, with soldier characteristics and reproductive ability, have been reported from several species of the family Termopsidae. Such intercastes have been considered the primitive-soldier caste, and based on this many researchers have discussed the evolutionary origin of termite soldiers. We investigated whether such soldier-reproductive intercastes also exist in the Japanese rotten-wood termite Hodotermopsis sjostedti. Abnormal individuals with soldier-like characteristics were found and designated as soldier-like intercastes, which appeared to have both soldier and reproductive characteristics. Based on our morphometric analyses and histological examinations, we suggest that the developmental origin of this intercaste is a pseudergate, nymph or sixth-instar larva. In addition, the intercaste was found to have relatively well-developed gonads, although mature oocytes and spermatozoa were not found in female ovaries and male seminal vesicles, respectively. We conclude that the soldier-like intercaste of H. sjostedti does not have reproductive ability, which is different from all other known soldier-like intercastes in Termopsidae.     

Alate differentiation and compound-eye development in the dry-wood termite <Emphasis Type="Italic">Neotermes koshunensis</Emphasis> (Isoptera,Kalotermitidae)

In social insects, caste-specific characters develop in the postembryonic differentiation processes. However, the mechanisms of caste-specific organ development have yet to be elucidated. In order to obtain insights into the relationship between caste differentiation and the regulation of organ development, we determined the caste-developmental pathway and observed compound-eye development accompanying alate differentiation in the dry-wood termite, Neotermes koshunensis. As previously reported in other Neotermes, this species has a linear caste-developmental pathway, comprising six larval- and two nymphal-instar stages. Although the apparent eye formation occurs during the last nymphal stages, just prior to the imaginal molt, individuals possess eye primordia from the first larval-instar stage. The outer morphological structure of the eye was observed from the third larval-instar stage. The detailed differentiation of cells constituting ommatidia appeared to occur in relatively young larval instars (fourth stage), although the pigmentation of pigment cells and detailed structural formation of ommatidia occurred during the final stage of alate development, i.e., during the late second nymphal-instar stage. This suggests that eye development is arrested in the larval stages, and then resumed during the late nymphal stage to complete functional eye formation, which is required for nuptial flight. In comparison to major hemimetabolous insects, which possess functional compound eyes even at the first instar larva, this termite species shows the heterochronic shift in terms of compound-eye development. Received 20 March 2006; revised 24 September 2006; accepted 4 October 2006.     

Antennal cropping in the Asian dry-wood termite, Neotermes koshunensis

Antennal cropping, a behavior inferred to exist because queens and kings have shorter antennae than fresh alates, is widespread in termites. However, the proximate and ultimate mechanisms underlying this phenomenon remain unclear. We studied the occurrence of antennal cropping in queens and kings of the dry-wood termite Neotermes koshunensis (Kalotermitidae). Observation of the antennal tip structure with scanning electron microscopy and the occurrence of antennal cropping in new kings and queens reared in isolation indicated that self-cropping is an important proximate mechanism. Previous studies inferred that antennal cropping may play a key role in the life-history of alates at the colony-founding stage. However, we also found antennal cropping in adultoid reproductives (secondary reproductives) that had not experienced a colony founding. We propose a new hypothesis that antennal cropping is important for individuals in regulating their physiology when they change from the non-reproducing to the reproducing phase.     

Microsatellite markers for the polygamous termite Nasutitermes corniger (Isoptera: Termitidae)

We developed eight highly variable microsatellite markers for the termite Nasutitermes corniger. Allele number per locus ranged from nine to 34, and expected heterozygosity from 0.45 to 0.94, in samples from seven sites in the former canal zone of Panama. The utility of these markers was assessed for five congeners varying in phylogenetic distance to N. corniger. The markers will be useful for fine‐scale examination of population and colony genetic structure in N. corniger and other closely related species.     

Habitat-specific life-history variation in the Caribbean termite Nasutitermes acajutlae (Isoptera: Termitidae)

Abstract.  1. Population and individual colony characteristics of Nasutitermes acajutlae on St John, U.S. Virgin Islands were studied over a 5-year period.
2. Four habitat types (dry forests, moist forests, sparse vegetation, and woodlands) were sampled for N. acajutlae , using quadrat-based methods. Lying dead wood was also quantified in the same sites.
3. Habitats differed significantly in the volume of lying dead wood, and total termite nest volume within quadrats was positively related to wood volume. It was hypothesised that the volume of dead wood affected growth, reproduction, and yearly survival of individual colonies.
4. In a sample of > 100 colonies assessed in 1998, 2000, 2001, and 2002, no significant differences in the growth rate of colonies (= increase in nest volume) could be found between habitat types; however, colonies in sparse vegetation were smaller, less likely to survive, and reproduced at a smaller size compared with those in the other three habitats.
5. Overall nest volume in sparse vegetation was high, compared with dry and moist forest, but colonies showed, on average, poorer survival (interpreted as lower fitness), and produced alates at an early stage of development. Optimum habitat characteristics for N. acajutlae may include not only favourable microclimate but also the reliable acquisition of new dead wood after storms.     

Caste differentiation pathways in the Neotropical termite Silvestritermes euamignathus (Isoptera: Termitidae)

Termites are hemimetabolous insects that commonly exhibit a bifurcation in post‐embryonic development, with the presence of two lines: (i) the nymphal–imaginal line; and (ii) the apterous line. The former comprises the reproductive imagoes of the colonies and their preceding immature stages, whereas the latter is composed of workers, soldiers and their precursors. In this study, we analyzed the developmental pathways of caste differentiation in the Neotropical termite Silvestritermes euamignathus and compared it to the known sibling species, Silvestritermes holmgreni. Our results show that the apterous line is composed of male and female workers without sexual dimorphism and that soldiers are exclusively females. The imaginal line comprises five instars of nymphs, which may develop into either nymphoid neotenics or alates. Similarities found in our study between caste patterns of S. euamignathus and of S. holmgreni suggest a shared evolutionary history in these termite species. Some biological differences between the species are also discussed.