Termitidae

Abstract: Summary 1. Gut content analysis of termites was undertaken using microscopical techniques. The 46 study species covered the entire range of taxonomic and feeding forms within the Order. 2. Inter-specific gut contents data were analysed using principal components analysis, placing species along a clear humification gradient based on variations in the amount of silica and plant tissue fragments in the gut. 3. Redundancy analysis was used to find morphological correlates of the observed variation in gut contents. A total of 22 morphological characters (out of 45 candidate characters) were correlated significantly with the gut contents. 4. Three of the 22 significantly correlated characters unambiguously defined feeding groups, which were designated groups I to IV in increasing order of humification of the feeding substrate. Group I contains lower termite dead wood and grass-feeders; group II contains Termitidae with a range of feeding habits including dead wood, grass, leaf litter, and micro-epiphytes; group III contains Termitidae feeding in the organic rich upper layers of the soil; group IV contains the true soil-feeders (again all Termitidae), ingesting apparently mineral soil. These groupings were generally supported statistically in a canonical covariance analysis, although group II apparently represents termite species with a rather wide range of feeding habits. 5. Using existing hypotheses of termite phylogenetic relationships, it seems probable that group I feeders are phylogenetically basal, and that the other groupings have arisen independently on a number of occasions. Soil-feeding (i.e. group III and group IV feeding) may have evolved due to the co-option of faecal material as a fungal substrate by Macrotermitinae-like ancestral forms. As a consequence, these forms would have been constrained to build nest structures from soil and would therefore have passed at least some soil through their guts.

Abstract: We investigated the bacterial gut microbiota from 32 colonies of wood-feeding termites, comprising four Microcerotermes species (Termitidae) and four Reticulitermes species (Rhinotermitidae), using terminal restriction fragment length polymorphism analysis and clonal analysis of 16S rRNA. The obtained molecular community profiles were compared statistically between individuals, colonies, locations, and species of termites. Both analyses revealed that the bacterial community structure was remarkably similar within each termite genus, with small but significant differences between sampling sites and/or termite species. In contrast, considerable differences were found between the two termite genera. Only one bacterial phylotype (defined with 97% sequence identity) was shared between the two termite genera, while 18% and 50% of the phylotypes were shared between two congeneric species in the genera Microcerotermes and Reticulitermes, respectively. Nevertheless, a phylogenetic analysis of 228 phylotypes from Microcerotermes spp. and 367 phylotypes from Reticulitermes spp. with other termite gut clones available in public databases demonstrated the monophyly of many phylotypes from distantly related termites. The monophyletic “termite clusters” comprised of phylotypes from more than one termite species were distributed among 15 bacterial phyla, including the novel candidate phyla TG2 and TG3. These termite clusters accounted for 95% of the 960 clones analyzed in this study. Moreover, the clusters in 12 phyla comprised phylotypes from more than one termite (sub)family, accounting for 75% of the analyzed clones. Our results suggest that the majority of gut bacteria are not allochthonous but are specific symbionts that have coevolved with termites and that their community structure is basically consistent within a genus of termites.

Abstract: Termites have colonized many habitats and are among the most abundant animals in tropical ecosystems, which they modify considerably through their actions. The timing of their rise in abundance and of the dispersal events that gave rise to modern termite lineages is not well understood. To shed light on termite origins and diversification, we sequenced the mitochondrial genome of 48 termite species and combined them with 18 previously sequenced termite mitochondrial genomes for phylogenetic and molecular clock analyses using multiple fossil calibrations. The 66 genomes represent most major clades of termites. Unlike previous phylogenetic studies based on fewer molecular data, our phylogenetic tree is fully resolved for the lower termites. The phylogenetic positions of Macrotermitinae and Apicotermitinae are also resolved as the basal groups in the higher termites, but in the crown termitid groups, including Termitinae + Syntermitinae + Nasutitermitinae + Cubitermitinae, the position of some nodes remains uncertain. Our molecular clock tree indicates that the lineages leading to termites and Cryptocercus roaches diverged 170 Ma (153-196 Ma 95% confidence interval [CI]), that modern Termitidae arose 54 Ma (46-66 Ma 95% CI), and that the crown termitid group arose 40 Ma (35-49 Ma 95% CI). This indicates that the distribution of basal termite clades was influenced by the final stages of the breakup of Pangaea. Our inference of ancestral geographic ranges shows that the Termitidae, which includes more than 75% of extant termite species, most likely originated in Africa or Asia, and acquired their pantropical distribution after a series of dispersal and subsequent diversification events.