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The Tea Bag Index (TBI) is a standardised and cheap method to quantify microbial-driven decomposition by measuring the mass loss of tea within tea bags. Termites are known to damage the bags to access the content, rendering the method less suitable for termite-rich ecosystems. Extension of the TBI to accommodate and incorporate the influence of termites would broaden its applicability to include termite-rich ecosystems, such as tropical forests. We extended the original TBI by applying physical and chemical termite-exclusion methods. Tea mass loss and the proportion of tea bags detected by termites in the original TBI were also recorded to infer the role of termites in litter decomposition. TBI estimates derived from the original and extended TBI were compared, benchmarked against global estimates, and validated with time-series mass loss data. Using the original TBI, we found that termites damaged up to 80 % of tea bags and consumed the recalcitrant fraction of tea in several of them, leaving only 20 % of tea bags from which TBI estimates could be retrieved. The physical termite-exclusion treatment completely eliminated termite-infringement, thus preserving the full sample size for estimating TBI parameters. The chemical termite-exclusion treatment also successfully excluded termites, but potentially inhibited microbial decomposition and made TBI estimates unreliable. In the absence of termite-infringement, both the TBI estimates and time-series analysis revealed a low decomposition rate compared to other measurements in tropical and temperate regions. We propose an extended TBI, in which the physical termite-exclusion treatment is used to preserve the retrieval rate of TBI parameters and reliably measure microbial-driven decomposition, while the original TBI is used to incorporate the contribution of termites in driving litter mass loss. By characterising both termite- and microbial-driven decomposition, the extended TBI will provide a comprehensive understanding of decomposition and its drivers in termite-rich ecosystems, and permit global comparisons.

A - C absolute weight-increase in g and D - F weight-increase relative to dryweight for beam geometry; humidification (relative humidity in air) and moistening (water dipping) of Po. alba, P. radiata and E. regnans over time. Average values are marked by a solid thick line. Period I describes the process of saturation at 80% RH (28 ºC), period II is the increase in weight owing to the wood being dipped in water and in period III the wood is oven-dried again

Animals use cues to find their food, in microhabitats within their physiological tolerances. Termites build and modify their microhabitat, to transform hostile environments into benign ones, which raises questions about the relative importance of cues. Termites are desiccation intolerant and foraging termites are attracted to water, so most research has considered moisture to be a cue. However, termites can also transport water to food, and so moisture may play other roles than previously considered. To examine the role of moisture, we compared Coptotermes acinaciformis termite foraging decisions in laboratory experiments when they were offered dry and moist wood, with and without load. Without load, termites preferred moist wood and ate it without any building, whereas they moistened dry wood after wrapping it in a layer of clay. For the ‘With load’ units, termites substituted some of the wood for load-bearing clay walls, and kept the wood drier than on the unloaded units. As drier wood has higher compressive strength and higher rigidity, it allows more of the wood to be consumed. These results suggest that moisture plays a more important role in termite ecology than previously thought. Termites manipulate the moisture content according to the situational context and use it for multiple purposes: increased moisture levels soften the fibre, which facilitates foraging, yet keeping the wood dry provides higher structural stability against buckling which is especially important when foraging on wood under load.

Animals use cues to find their food, in microhabitats within their physiological tolerances. Termites build and modify their microhabitat, to transform hostile environments into benign ones, which raises questions about the relative importance of cues. Termites are desiccation intolerant and foraging termites are attracted to water, so most research has considered moisture to be a cue. However, termites can also transport water to food, and so moisture may play other roles than previously considered. To examine the role of moisture, we compared Coptotermes acinaciformis termite foraging decisions in laboratory experiments when they were offered dry and moist wood, with and without load. Without load, termites preferred moist wood and ate it without any building, whereas they moistened dry wood after wrapping it in a layer of clay. For the ‘With load’ units, termites substituted some of the wood for load-bearing clay walls, and kept the wood drier than on the unloaded units. As drier wood has higher compressive strength and higher rigidity, it allows more of the wood to be consumed. These results suggest that moisture plays a more important role in termite ecology than previously thought. Termites manipulate the moisture content according to the situational context and use it for multiple purposes: increased moisture levels soften the fibre, which facilitates foraging, yet keeping the wood dry provides higher structural stability against buckling which is especially important when foraging on wood under load.

Termite mounds built by representatives of the family Termitidae are among the most spectacular constructions in the animal kingdom, reaching 6–8 m in height and housing millions of individuals. Although functional aspects of these structures are well studied, their evolutionary origins remain poorly understood. Australian representatives of the termitid subfamily Nasutitermitinae display a wide variety of nesting habits, making them an ideal group for investigating the evolution of mound building. Because they feed on a variety of substrates, they also provide an opportunity to illuminate the evolution of termite diets. Here, we investigate the evolution of termitid mound building and diet, through a comprehensive molecular phylogenetic analysis of Australian Nasutitermitinae. Molecular dating analysis indicates that the subfamily has colonized Australia on three occasions over the last approximately 20 Myr. Ancestral-state reconstruction showed that mound building arose on multiple occasions and from diverse ancestral nesting habits, including arboreal and wood or soil nesting. Grass feeding appears to have evolved from wood feeding via ancestors that fed on both wood and leaf litter. Our results underscore the adaptability of termites to ancient environmental change, and provide novel examples of parallel evolution of extended phenotypes.

Termite mounds built by representatives of the family Termitidae are among the most spectacular constructions in the animal kingdom, reaching 6–8 m in height and housing millions of individuals. Although functional aspects of these structures are well studied, their evolutionary origins remain poorly understood. Australian representatives of the termitid subfamily Nasutitermitinae display a wide variety of nesting habits, making them an ideal group for investigating the evolution of mound building. Because they feed on a variety of substrates, they also provide an opportunity to illuminate the evolution of termite diets. Here, we investigate the evolution of termitid mound building and diet, through a comprehensive molecular phylogenetic analysis of Australian Nasutitermitinae. Molecular dating analysis indicates that the subfamily has colonized Australia on three occasions over the last approximately 20 Myr. Ancestral-state reconstruction showed that mound building arose on multiple occasions and from diverse ancestral nesting habits, including arboreal and wood or soil nesting. Grass feeding appears to have evolved from wood feeding via ancestors that fed on both wood and leaf litter. Our results underscore the adaptability of termites to ancient environmental change, and provide novel examples of parallel evolution of extended phenotypes.