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Fig. 8 in Discovery of termitophilous rove beetles associated with Formosan subterranean termite Coptotermes formosanus in Taiwan, with the first larval description for the tribe Termitohospitini (Coleoptera: Staphylinidae)
Fig. 8. Japanophilus hojoi Maruyama & Iwata, 2002, larval instar 1. A – abdominal tergites; B – abdominal sternites I and II; C – abdominal segments VIII, IX, and X, lateral view; D – abdominal tergites VIII, IX, and X; E – abdominal sternites IX and X. Abbreviations: A – anterior setae; Ah – anal hooks; D – discal setae, rows a–d; L – lateral setae; P – posterior setae; Sp – spiracle; Tgrs – tergal glandular reservoir sac; Tgo – tergal gland opening; Ug – urogomphus.
Fig. 6 in Discovery of termitophilous rove beetles associated with Formosan subterranean termite Coptotermes formosanus in Taiwan, with the first larval description for the tribe Termitohospitini (Coleoptera: Staphylinidae)
Fig. 6. Japanophilus hojoi Maruyama & Iwata, 2002, larval instar 1. A – mandibles, dorsal view; B – labrum, dorsal view; C–D – right antenna, dorsal (C) and lateral (D) view; E – right maxilla, ventral view; F–G – curved seta of mala, ventrolateral (F) and lateral (G) view; H – labium, ventral view. Abbreviations: I, II, III – antennal articles; IIIS (1–3) – solenidia of article 3; Cdo – cardo; Fm – frontal marginal seta; Ld – labral dorsal setae; Lg – ligula; Ll – labral lateral seta; Lm – labral marginal setae; Ma – mala; Mnt – mentum; Pf – palpifer; Pmnt – prementum; Sa – sensory appendage; Smnt – submentum; Stp – stipes.
Variation in susceptibility of Eucalyptus grandis and selected hybrid clones to two termite species Macrotermes bellicosus and M. subhyalinus in Uganda
<p>The maximum productivity of plantation forestry and its role in climate change mitigation, adaptation and resilience cannot be met without proper management. Termites in the genus <i>Macrotermes</i> have been reported as a major challenge to <i>Eucalyptus</i> plantation forestry establishment. The current study evaluated the susceptibility of four <i>Eucalyptus</i> hybrid clones; GU 7, GC 796, GC 550 and GC 796/2 and <i>E. grandis </i>to the most damaging <i>Macrotermes bellicosus </i><span>(Smeathman) </span>and <i>Macrotermes subhyalinus </i><span>(Rambur) to identify tolerant material that can be planted in high incidence areas. The study involved exposure of moisture dry pieces of wood from <i>E. grandis</i> and the four hybrid clones to damage by <i>M. bellicosus</i> and <i>M. subhyalinus</i>. Results confirmed that <i>M. bellicosus</i> is the most aggressive. Results further revealed that <i>E. grandis</i> and GC 550 are the most susceptible whereas GC 796 is the most tolerant clone. The findings from the study will contribute to improved management of termites by planting tolerant material in high risk areas. </span></p>
Figure 5 from: Scheffrahn RH, Vasconcellos A (2020) Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil. ZooKeys 954: 75-84. https://doi.org/10.3897/zookeys.954.52335
Figure 5 Map of Tauritermes from the literature and UFTC. Biomes are shown for Brazil. See Scheffrahn (2019b) for UFTC data and Table 3 for literature references.
Figure 4 from: Scheffrahn RH, Vasconcellos A (2020) Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil. ZooKeys 954: 75-84. https://doi.org/10.3897/zookeys.954.52335
Figure 4 Soldier of Tauritermes taurocephalus (BO722) A dorsal B lateral C oblique, and D ventral views of head and pronotum. DH = dorsal horn.
Figure 2 from: Scheffrahn RH, Vasconcellos A (2020) Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil. ZooKeys 954: 75-84. https://doi.org/10.3897/zookeys.954.52335
Figure 2 Soldier of Tauritermes bandeirai sp. nov. (SA499) A dorsal B lateral C oblique, and D ventral views of head and pronotum. AC = antennal carina, DH = dorsal horn, FH = frontal horn, and GH = genal horn.
Figure 3 from: Scheffrahn RH, Vasconcellos A (2020) Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil. ZooKeys 954: 75-84. https://doi.org/10.3897/zookeys.954.52335
Figure 3 Soldier of Tauritermes triceromegas (PA942) A dorsal B lateral C oblique, and D ventral views of head and pronotum. DH = dorsal horn and FH = frontal horn.
Figure 1 from: Scheffrahn RH, Vasconcellos A (2020) Tauritermes bandeirai: A new drywood termite (Isoptera, Kalotermitidae) from the Caatinga and Atlantic Forest of Brazil. ZooKeys 954: 75-84. https://doi.org/10.3897/zookeys.954.52335
Figure 1 Imago of Tauritermes bandeirai sp. nov. (SA502) A dorsal and B lateral views of head and pronotum C right forewing.
Figure 4 from: Scheffrahn RH, Pinzón Florian OP (2020) Rugitermes tinto: A new termite (Isoptera, Kalotermitidae) from the Andean region of Colombia. ZooKeys 963: 37-44. https://doi.org/10.3897/zookeys.963.55843
Figure 4 Anteriodorsal views of Rugitermes soldier heads from South America. Margins of left frontolateral ridges enhanced. AR. bicolor, PU946 BR. flavicinctus, TT88 CR. laticollis, EC1465 DR. magninotus, PU1087 ER. occidentalis, AG380 FR. niger, AG500 GR. nodulosus (modified from fig. 14, Krishna 1961) HR. rugosus PA1186. Accession numbers from UFTC (Scheffrahn 2019b). Scale bars: 1 mm.
Figure 1 from: Scheffrahn RH, Pinzón Florian OP (2020) Rugitermes tinto: A new termite (Isoptera, Kalotermitidae) from the Andean region of Colombia. ZooKeys 963: 37-44. https://doi.org/10.3897/zookeys.963.55843
Figure 1 Type localities for all Rugitermes species described from South America and collection localities for Rugitermes spp. in the University of Florida Termite Collection (UFTC).
Figure 2 from: Scheffrahn RH, Pinzón Florian OP (2020) Rugitermes tinto: A new termite (Isoptera, Kalotermitidae) from the Andean region of Colombia. ZooKeys 963: 37-44. https://doi.org/10.3897/zookeys.963.55843
Figure 2 Rugitermes tinto sp. nov. A Dorsal B lateral views of head and thorax of imago C dorsal D frontal views of anterior soldier head capsule (white arrows mark frontal flange, yellow arrows mark frontolateral ridge, and black arrows mark protuberance of frontolateral ridge).
Termite males enhance mating encounters by changing speed according to density
<ol> <li><span>Search theory predicts that animals evolve efficient movement patterns to enhance encounter rates with specific targets. The optimal movements vary with the surrounding environments, which may explain the observation that animals often switch their movement patterns depending on conditions. However, the effectiveness of behavioral change during search is rarely evaluated because it is difficult to examine the actual encounter dynamics.</span></li> <li><span>Here we studied how partner-seeking termites update their search strategies depending on the local densities of potential mates. After a dispersal flight, termites drop their wings and walk to search for a mate; when a female and a male meet, they form a female-led tandem pair and search for a favorable nesting site. If a pair is separated, they have two search options: reunite with their stray partner, or seek a new partner. We hypothesized that the density of individuals affects separation-reunion dynamics and thus the optimal search strategy.</span></li> <li><span>We observed the searching process across different densities and found that termite pairs were often separated but obtained a new partner quickly at high mate density. After separation, while females consistently slowed down, males increased their speed according to the density. Under high mate density, separated males obtained a partner earlier than females, who do not change movement with density.</span></li> <li><span>Our data-based simulations confirmed that the observed behavioral change by males contributes to enhancing encounters. Males at very low mate densities did best to move slowly and thereby reduce the risk of missing their stray partner, who is the only available mate. On the other hand, males that experienced high mate densities did better in mating encounters by moving fast because the risk of isolation is low, and they must compete with other males to find a partner.</span></li> <li><span>These results demonstrate that termite males adaptively update their search strategy depending on conditions. Understanding the encounter dynamics experienced by animals is key to connecting empirical work to the idealized search processes of theoretical studies. </span></li> </ol>
Supplementary material 2 from: Basset Y, Donoso DA, Hajibabaei M, Wright MTG, Perez KHJ, Lamarre GPA, De León LF, Palacios-Vargas JG, Castaño-Meneses G, Rivera M, Perez F, Bobadilla R, Lopez Y, Ramirez JA, Barrios H (2020) Methodological considerations for monitoring soil/litter arthropods in tropical rainforests using DNA metabarcoding, with a special emphasis on ants, springtails and termites. Metabarcoding and Metagenomics 4: e58572. https://doi.org/10.3897/mbmg.4.58572
Appendix S2
Data from: Lack of aggression and apparent altruism towards intruders in a primitive termite
In eusocial insects, the ability to discriminate nest-mates from non-nest-mates is widespread and ensures that altruistic actions are directed towards kin and agonistic actions are directed towards non-relatives. Most tests of nest-mate recognition have focused on hymenopterans, and suggest that cooperation typically evolves in tandem with strong antagonism towards non-nest-mates. Here, we present evidence from a phylogenetically and behaviourally basal termite species that workers discriminate members of foreign colonies. However, contrary to our expectations, foreign intruders were the recipients of more rather than less cooperative behaviour and were not subjected to elevated aggression. We suggest that relationships between groups may be much more peaceable in basal termites compared with eusocial hymenoptera, owing to energetic and temporal constraints on colony growth, and the reduced incentive that totipotent workers (who may inherit breeding status) have to contribute to self-sacrificial intergroup conflict.
Data from: Hemimetabolous genomes reveal molecular basis of termite eusociality
Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the 2-Gb genome of the German cockroach, Blattella germanica, and the 1.3-Gb genome of the drywood termite Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolutionary transitions in biological complexity.
Data from: Catechol 2,3-dioxygenase and other meta-cleavage catabolic pathway genes in the 'anaerobic' termite gut spirochete Treponema primitia
Microorganisms have evolved a spectacular diversity of metabolisms, some of which allow them to overcome environmental constraints, utilize abundant but inaccessible resources and drive nutrient cycling in various ecosystems. The termite hindgut microbial community is optimized to metabolize wood, and in recent years, the in situ physiological and ecological functions of community members have been researched. Spirochetes are abundant in the termite gut, and herein, putative aromatic meta-cleavage pathway genes typical of aerobic pseudomonads were located in genomes of homoacetogenic termite hindgut 'anaerobes', Treponema primitia str. ZAS-1 and ZAS-2. Phylogenetic analyses suggest the T. primitia catechol 2,3-dioxygenase and several other essential meta-pathway genes were acquired from an α-proteobacterium in the distant past to augment several genes T. primitia acquired from anaerobic firmicutes that do not directly catabolize aromatics but can contribute to the final pathway steps. Further, transcripts for each meta-pathway gene were expressed in strictly anaerobic cultures of T. primitia str. ZAS-2 indicative of constitutive pathway expression. Also, the addition of catechol + O2 to T. primitia liquid cultures resulted in the transient accumulation of trace amounts of the yellow ring cleavage product, hydroxymuconic semialdehyde. This is the first evidence of aromatic ring cleavage in the phylum (division) Spirochetes. Results also support a possible role for T. primitia in termite hindgut O2/lignin aromatic monomer metabolism. Potential O2-dependent yet nonrespiratory microbial metabolisms have heretofore been overlooked and warrant further investigation. These metabolisms could describe the degradation of plant-derived and other aromatics in microoxic environments and contribute significantly to carbon turnover.
Data from: Breaking the cipher: ant eavesdropping on the variational trail pheromone of its termite prey
Predators may eavesdrop on their prey using innate signals of varying nature. In regards to social prey, most of the prey signals are derived from social communication and may therefore be highly complex. The most efficient predators select signals that provide the highest benefits. Here, we showed the use of eusocial prey signals by the termite-raiding ant Odontoponera transversa. O. transversa selected the trail pheromone of termites as kairomone in several species of fungus-growing termites (Termitidae: Macrotermitinae: Odontotermes yunnanensis, Macrotermes yunnanensis, Ancistrotermes dimorphus). The most commonly predated termite, O. yunnanensis, was able to regulate the trail pheromone component ratios during its foraging activity. The ratio of the two trail pheromone compounds was correlated with the number of termites in the foraging party. (3Z)-Dodec-3-en-1-ol (DOE) was the dominant trail pheromone component in the initial foraging stages when fewer termites were present. Once a trail was established, (3Z,6Z)-dodeca-3,6-dien-1-ol (DDE) became the major recruitment component in the trail pheromone and enabled mass recruitment of nest-mates to the food source. Although the ants could perceive both components, they revealed stronger behavioural responses to the recruitment component, DDE, than to the common major component, DOE. In other words, the ants use the trail pheromone information as an indication of suitable prey abundance, and regulate their behavioural responses based on the changing trail pheromone component. The eavesdropping behaviour in ants therefore leads to an arms race between predator and prey where the species specific production of trail pheromones in termites is targeted by predatory ant species.
Data from: Dopamine regulates termite soldier differentiation through trophallactic behaviours
Caste polyphenism in social insects is regulated by social interactions among colony members. Trophallaxis is one of the most frequently observed interactions, but no studies have been conducted identifying the intrinsic factors involved in this behaviour and caste differentiation. Dopamine (DA) has multiple roles in the modulation of behaviours and physiology, and it produces species-specific behaviours in animals. Here, to verify the role of DA in termite soldier differentiation, we focused on the first soldier in an incipient colony of Zootermopsis nevadensis, which always differentiates from the oldest 3rd instar (No. 1 larva) via a presoldier. First, brain DA levels of the No. 1 larva at day 3 after its appearance were significantly higher than day 0. Second, DA synthesis gene expression levels were extraordinarily high in the No. 1 larva at day 0–1 after appearance. Finally, injection of a DA receptor antagonist into the No. 1 larva resulted in the inhibition of presoldier differentiation. Behavioural observations of the antagonist or control-injected larvae suggested that brain DA and signalling activity regulate the frequencies of trophallaxis from reproductives and presoldier differentiation. Because trophallaxis is a social behaviour frequently observed in natural conditions, the role of DA should be investigated in other social insects with frequent trophallactic and allogrooming behaviour.
Data from: Diet is the primary determinant of bacterial community structure in the guts of higher termites
The gut microbiota of termites plays critical roles in the symbiotic digestion of lignocellulose. While phylogenetically 'lower termites' are characterized by a unique association with cellulolytic flagellates, higher termites (family Termitidae) harbour exclusively prokaryotic communities in their dilated hindguts. Unlike the more primitive termite families, which primarily feed on wood, they have adapted to a variety of lignocellulosic food sources in different stages of humification, ranging from sound wood to soil organic matter. In this study, we comparatively analysed representatives of different taxonomic lineages and feeding groups of higher termites to identify the major drivers of bacterial community structure in the termite gut, using amplicon libraries of 16S rRNA genes from 18 species of higher termites. In all analyses, the wood-feeding species were clearly separated from humus and soil feeders, irrespective of their taxonomic affiliation, offering compelling evidence that diet is the primary determinant of bacterial community structure. Within each diet group, however, gut communities of termites from the same subfamily were more similar than those of distantly related species. A highly resolved classification using a curated reference database revealed only few genus-level taxa whose distribution patterns indicated specificity for certain host lineages, limiting any possible cospeciation between the gut microbiota and host to short evolutionary timescales. Rather, the observed patterns in the host-specific distribution of the bacterial lineages in termite guts are best explained by diet-related differences in the availability of microhabitats and functional niches.
Data from: Extreme genetic mixing within colonies of the wood-dwelling termite Kalotermes flavicollis (Isoptera, Kalotermitidae)
The existence of altruism in social insects is commonly attributed to altruistic individuals gaining indirect fitness through kin selection. However, recent studies suggest that such individuals might also gain direct fitness through reproduction. Experimental studies on primitive wood-dwelling termites revealed that colony fusion often causes the death of primary reproductives (queen and king), allowing opportunities for workers to inherit the nest by developing into replacement reproductives (neotenics). Therefore, colony fusion has been proposed as an important factor that may have favoured sociality in termites. However, whether colony fusion occurs frequently in natural populations of wood-dwelling termites remains an open question. We analysed eleven colonies of the wood-dwelling termite Kalotermes flavicollis (Kalotermitidae), using two mitochondrial and five nuclear microsatellite markers. Nine of eleven colonies (82%) were mixed families, with offspring of three or more primary reproductives. To our knowledge, this result represents the highest frequency of mixed-family colonies ever reported in termites. Moreover, genetic mixing of colonies appeared extreme in two ways. First, the number of haplotypes per colony was exceptionally high (up to nine), indicating that colonies were composed of multiple queens' offspring. Second, some mixed-family colonies included individuals belonging to two highly divergent genetic lineages. F-statistics and relatedness values suggest that mixed-family colonies most likely result from colony fusion, giving support to the accelerated nest inheritance theory. These findings raise important questions about the mode of foundation of mixed-family colonies and the evolutionary forces that maintain them within populations.
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