the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
| 01 Jun 2026
Ideas and perspectives: Beyond Microbes: Integrating Termites into Global Soil Carbon Cycling Models
Abstract. Termites are major detritivores in tropical and subtropical ecosystems, yet their contributions to the terrestrial carbon cycle remains absent from process-based soil organic carbon (SOC) models. Here, we present a termite carbon module that explicitly represents termite-mediated litter consumption and transfer of ingested carbon into gaseous (CO2, CH4) and SOC pools. The module integrates biome-specific termite biomass with spatially explicit productivity inputs to quantify termite-driven carbon fluxes within a mass-balance framework. Model simulations show that termites act as spatially heterogeneous carbon processors, accelerating litter turnover while modifying the pathways through which carbon is redistributed between atmospheric and SOC pools. Global sensitivity analysis identifies termite biomass and ingestion capacity as the dominant controls on flux magnitude, whereas carbon partitioning governs the fate of processed carbon. Including termite-mediated pathways in SOC models provides a mechanism for representing faunal controls on decomposition, soil carbon formation, and trace gas emissions, particularly in tropical and seasonally dry ecosystems. Globally, we estimate termites process 1569.4 ± 800.4 Tg C yr-1, releasing 864.7 ± 444.5 Tg C yr-1 as CO2 and 7.9 ± 4.9 Tg C yr-1 as CH4, while transferring 689.3 ± 367.4 Tg C yr-1 into labile and mineral-associated SOC. Explicit representation of termite-driven carbon fluxes will therefore be important for improving predictions of litter decomposition, SOC formation, and terrestrial carbon-climate feedbacks.
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Status: open (until 14 Aug 2026)
- RC1: 'Comment on egusphere-2026-2844', Omar Flores, 22 Jun 2026 reply
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General comments
The study by Farooq et al. presents a novel work that explicitly includes termites within Microbial Explicit Soil Carbon (MES-C), a process-based SOC model. The relevance of such work is well justified, particularly due to the key role of termites in regulating SOC sequestration and CO2 and CH4 emissions, which is missing in global SOC models.
The main technical weaknesses of the presented work (e.g. uncertainty in some assumptions, simplification of key soil processes, homogeneous spatial representation of termites despite their highly heterogeneous distribution) are reasonable taking into account issues in the availability of data, the novelty of the approach, and common limitations in model structure. Moreover, they have been also properly addressed by the authors.
Nevertheless, there is one issue with the perspective of a detailed representation of termites into global models. The authors have divided termites in three feeding guilds, with their own parameters and specialized interactions with litter and SOC pools. Such level of detail is very reasonable for ecosystem-scale models, but it might be excessive for global models, which is the claimed goal of the authors. Despite it can be done, as shown by the authors, a question remains: should we really add all that complexity in global models? There is no doubt that termites are an important biota group with high impact, but the same can be said for several other groups, like ants, earthworms, nematodes, etc. Global models normally do not represent any of them, and if they were to include them, it would not be feasible to add each taxon separately (let alone with several guilds or subgroups per taxon). While the specific framework developed by the authors can be interesting for specific simulations related to termites, or as reference for future model developments, its future integration into global models seeking to improve the representation of soil fauna will probably need a simplification. After reading this study, I see right now two possibilities (for future developments):
1) Justify (if that is the case) that termites must remain separately from any other biota due to unique roles, but perhaps combining all guilds into one pool, and parameterize the different roles of each guild within that common pool. For example, equations S7-S9 could be modified using one single TER pool, but adding in each case a multiplier with the fraction of the TER pool that corresponds to the fraction of biomass of each feeding guild (at each time step). Some processes could retain the specialized effects of each guild that way, while other processes should be simplified with an average effect of the entire termite biomass across guilds, which would reduce the number of parameters (together with reduced interactions and fluxes) and therefore reduce complexity.
2) Discuss if termites could be mixed with other fauna groups, splitting if needed their guilds into different functional groups of soil fauna, in the same way that for example in some soil food web models nematodes are divided into different functional groups depending on their body size and/or feeding roles (e.g. microbivores, herbivores, predators). Maybe the way to go could be to modify generic functional groups (once they are added to global models) to account for the specific roles played by termites, as for example including the role of TERSF within microbivores, and the roles of TERFG and TERXP within detritivores, taking into account that termites will only be a fraction of such pools.
I don’t think there is any need to modify the study for that reason, the presented perspectives (and all equations, results, etc.) can stay as they are, but I would suggest to add to the discussion also those challenges related to adding termites to global models while keeping complexity as low as possible. Despite this work has merit on its own and can certainly help to improve models, I would like the authors to address, from their expertise on modelling termites, what would be the best option to deal with future global modelling frameworks including termites along with other soil fauna groups.
Overall, the manuscript is interesting, well written, it represents a first step towards a potentially relevant improvement of global SOC models, and it should be suitable for publication after a minor revision.
Specific comments
Table 2 – The total Area was apparently calculated as the sum of all the areas by vegetation type. If I understand the represented concept correctly, termites occupy 17.3 m/km2 of tropical evergreen, 5.9 m/km2 of tropical deciduous, etc. Then what you are suggesting with that total is that, in general over all vegetation types, termites occupy 72.3 m/km2, right? I don’t think that is correct. You should calculate an average instead of a sum for such variable, or even a weighted average considering the different total areas of each vegetation type. If my interpretation is not correct, please explain better what exactly are you showing there.
You explain in the manuscript (L210) that GPP was smoothed using a 7-day running mean, while in the supplementary (L58) you also mention that you obtained GPP from MODIS 8-day productivity product, and that you constructed daily climatological time series. Can you explain in more detail how did you process the 8-day data? In particular, for generating the daily data; the use of a 7-day running mean from daily data is more obvious and should not require more details, but it would be better to explain the method used to go from 8-day to daily.
Supplementary:
L44 – the symbols ω and δ determine the fractions of decomposed carbon entering the LMWC pool for frass and necromass, respectively, with their complementary fractions determining what goes to MAOC in each case; the text wrongly describes them as if they refer to the different SOC pools, when they refer to different output fluxes from termite biomass.
Table S3 – the values shown for fractions of total termite biomass for Asia and Africa not always add up to 1. In particular, the first two rows for tropical vegetation (evergreen and deciduous) sum in total 0.91, and savanna and dense shrubland only have 0.33. Please explain why is that. Conversely, for grassland, there is already 1 for soil-feeding termites (SF), but also shows a zero for xylophagous termites (XP), while other cases in which one guild has 1 the others show a hyphen instead of a zero. Is that because the shown values for grassland are rounded, but SF guild is actually lower than 1, and XP higher than 0?
Equation S14 – add an explanation of why the denominator is the square root of 12.
Technical corrections
The manuscript title is not very fluent to read, with the double colon. I would suggest to remove “Ideas and perspectives:” entirely, or if not, then at least remove the first colon, or rewrite the title in a more fluent way (not a correction, just a suggestion).
L17 – “their contributions … remain” (instead of “remains”).
L198 – “carbon biomass” instead of “biomass carbon”.
L256 – In Fig. 2 caption, better make a clearer reference to the latitudinal mean representation, e.g. “The subset in each plot (blue line) shows…”. The same applies for Fig. S2.
L407 – “is available” instead of “are available”
Supplementary:
L21 – missing space in “times(Wang et al., 2010)”.
L26 – missing space in “underwentmicrobial”.
Equations S13 – kB should be kn
Table S3 – “from Sanderson (1996)”, the author name is explicitly referenced, so it should not be inside the parenthesis.
L68 – “carbon” instead of “substrates”, within the full name of LMWC.