Nest composition, stable isotope ratios and microbiota unravel the feeding behaviour of an inquiline termite.

Termites are eusocial insects having evolved several feeding, nesting and reproductive strategies. Among them, inquiline termites live in a nest built by other termite species: some of them do not forage outside the nest, but feed on food stored by the host or on the nest material itself. In this study, we characterized some dimensions of the ecological niche of Cavitermes tuberosus (Termitidae: Termitinae), a broad-spectrum inquiline termite with a large neotropical distribution, to explain its ecological success. We used an integrative framework combining ecological measures (physico-chemical parameters, stable isotopic ratios of N and C) and Illumina MiSeq sequencing of 16S rRNA gene to identify bacterial communities and to analyse termites as well as the material from nests constructed by different termite hosts (the builders). Our results show that (1) nests inhabited by C. tuberosus display a different physico-chemical composition when compared to nests inhabited by its builder alone; (2) stable isotopic ratios suggest that C. tuberosus feeds on already processed, more humified, nest organic matter; and (3) the gut microbiomes cluster by termite species, with the one of C. tuberosus being much more diverse and highly similar to the one of its main host, Labiotermes labralis. These results support the hypothesis that C. tuberosus is a generalist nest feeder adapted to colonize nests built by various builders, and explain its ecological success.

Optimizing the choice of a spatial weighting matrix in eigenvector-based methods.

Eigenvector-mapping methods such as Moran's eigenvector maps (MEM) are derived from a spatial weighting matrix (SWM) that describes the relations among a set of sampled sites. The specification of the SWM is a crucial step, but the SWM is generally chosen arbitrarily, regardless of the sampling design characteristics. Here, we compare the statistical performances of different types of SWMs (distance-based or graph-based) in contrasted realistic simulation scenarios. Then, we present an optimization method and evaluate its performances compared to the arbitrary choice of the most-widely used distance-based SWM. Results showed that the distance-based SWMs generally had lower power and accuracy than other specifications, and strongly underestimated spatial signals. The optimization method, using a correction procedure for multiple tests, had a correct type I error rate, and had higher power and accuracy than an arbitrary choice of the SWM. Nevertheless, the power decreased when too many SWMs were compared, resulting in a trade-off between the gain of accuracy and the loss of power. We advocate that future studies should optimize the choice of the SWM using a small set of appropriate candidates. R functions to implement the optimization are available in the adespatial package and are detailed in a tutorial.

Use of phytoextraction with Noccaea caerulescens to limit the transfer of cadmium and zinc to subsequent rocket crops.

Soil trace metal (TM) contamination is a worldwide issue and threatens food production and security. Remediation of cadmium (Cd) and zinc (Zn) contaminated soils by phytoextraction with the Zn/Cd hyperaccumulator Noccaea caerulescens is widely studied but few studies have investigated the efficiency of this technique to reduce Cd and Zn soil-to-crop transfers to subsequent vegetable crops. The vegetable biomonitor rocket Diplotaxis tenuifolia was grown in pots on 13 moderately contaminated soils that had previously been cropped with N. caerulescens. Using mixed-effects models, we show the drivers of rocket biomass, Cd and Zn concentrations. Our models show, for our study soils, the benefit of previous N. caerulescens uptake of Cd and Zn in decreasing Cd and Zn concentrations in a subsequent rocket crop. We also show a slight positive impact of N. caerulescens biomass (and therefore uptake) on rocket growth. Our data show that exchangeable soil concentrations are major drivers of Cd and Zn rocket concentrations. Other soil variables negatively driving rocket Cd and Zn concentrations are NO3- content, organic matter content, cation exchange capacity, and soil manganese which stimulate rocket biomass and/or influence TM bioavailability. Rocket D. tenuifolia seems to be a good biomonitor for contaminated soils as it is tolerant to relatively high TM soil concentrations. We demonstrate that 40 % of rockets grown on soils below 2 mg total Cd kg-1 dry soil have foliar Cd concentrations above the European maximum allowed level confirming the need to review soil legal thresholds to protect consumers' health. In conclusion, our study suggests promising use of N. caerulescens phytoextraction for bioavailable contaminant stripping which is all the more interesting given the increasing demand for urban growing spaces.

Influence of edaphic conditions and nitrogen fertilizers on cadmium and zinc phytoextraction efficiency of Noccaea caerulescens.

The success of cadmium phytoextraction operations with Noccaea caerulescens varies by a factor of 70 between sites of trials. However, soil factors driving the efficiency of cadmium (Cd) and zinc (Zn) phytoextraction are still poorly understood, as are the effects of nitrogen fertilizers. We studied biomass production and Cd and Zn uptake by two contrasting populations of N. caerulescens, Ganges (metallicolous) and Wilwerwiltz (non-metallicolous) grown in pots on a range of 24 field contaminated soils for 20 weeks. The addition of KNO3 and NH4NO3 fertilizers was also tested. Using model averaging of multiple regression models, we show that the major drivers of N. caerulescens growth are physical soil factors such as organic matter and soil bulk density while trace metal accumulation mainly relies on soil Cd and Zn exchangeable concentrations. We confirm the negative effect of soil copper (Cu) on growth, even at exchangeable concentrations below 30 mg kg-1, and therefore on uptake efficiency, while increasing soil lead (Pb) content was related to increased biomass probably due to a protective effect against soil pathogens. Finally, there is a small positive effect of nitrogen fertilization on biomass production only in soils with low initial nitrogen content (under 25 µg g-1 NO3-), while above this value, the positive impact of initial nitrogen content is offset by lower shoot Cd and Zn concentrations. Our data bring substantial information regarding the physico-chemical properties to ensure N. caerulescens growth: a soil bulk density under 1.05 kg/dm3, organic matter above 7% and pH under 7.5. We show that phytoextraction efficiency is maximal for moderate soil contamination in Cd (2-10 mg kg-1) and Zn (300-1000 mg kg-1).

Phytoremediation of urban soils contaminated with trace metals using Noccaea caerulescens: comparing non-metallicolous populations to the metallicolous 'Ganges' in field trials.

Urban soil contamination with trace metals is a major obstacle to the development of urban agriculture as crops grown in urban gardens are prone to accumulate trace metals up to toxic levels for human consumption. Phytoextraction is considered as a potentially cost-effective alternative to conventional methods such as excavation. Field trials of phytoextraction with Noccaea caerulescens were conducted on urban soils contaminated with Cd, Cu, Pb, and Zn (respectively around 2, 150-200, 400-500, and 400-700 µg g-1 of dry soil). Metallicolous (Ganges population) and non-metallicolous (NMET) populations were compared for biomass production and trace metal uptake. Moreover, we tested the effect of compost and fertilizer addition. Maximal biomass of 5 t ha-1 was obtained with NMET populations on some plots. Compared to Ganges- the high Cd-accumulating ecotype from South of France often used in phytoextraction trials- NMET populations have an advantage for biomass production and for Zn accumulation, with an average Zn uptake of 2.5 times higher. The addition of compost seems detrimental due to metal immobilization in the soil with little or no effect on plant growth. In addition to differences between populations, variations of growth and metal accumulation were mostly explained by soil Cd and Zn concentrations and texture. Our field trials confirm the potential of using N. caerulescens for both Cd and Zn remediation of moderately contaminated soils-with uptake values of up to 200 g Cd ha-1 and 47 kg Zn ha-1-and show the interest of selecting the adequate population according to the targeted metal.

Multiscale assemblage of an ectomycorrhizal fungal community: the influence of host functional traits and soil properties in a 10-ha miombo forest.

Ectomycorrhizal fungi (EMF) are highly diversified and dominant in a number of forest ecosystems. Nevertheless, their scales of spatial distribution and the underlying ecological processes remain poorly understood. Although most EMF are considered to be generalists regarding host identity, a preference toward functional strategies of host trees has never been tested. Here, the EMF community was characterised by DNA sequencing in a 10-ha tropical dry season forest-referred to as miombo-an understudied ecosystem from a mycorrhizal perspective. We used 36 soil parameters and 21 host functional traits (FTs) as candidate explanatory variables in spatial constrained ordinations for explaining the EMF community assemblage. Results highlighted that the community variability was explained by host FTs related to the 'leaf economics spectrum' (adjusted R(2) = 11%; SLA, leaf area, foliar Mg content), and by soil parameters (adjusted R(2) = 17%), notably total forms of micronutrients or correlated available elements (Al, N, K, P). Both FTs and soil generated patterns in the community at scales ranging from 75 to 375 m. Our results indicate that soil is more important than previously thought for EMF in miombo woodlands, and show that FTs of host species can be better predictors of symbiont distribution than taxonomical identity.

Correction: Influence of Soil Properties on Soldierless Termite Distribution.

[This corrects the article DOI: 10.1371/journal.pone.0135341.].

Influence of Soil Properties on Soldierless Termite Distribution.

In tropical rainforests, termites constitute an important part of the soil fauna biomass, and as for other soil arthropods, variations in soil composition create opportunities for niche partitioning. The aim of this study was twofold: first, we tested whether soil-feeding termite species differ in the foraging substrate; second, we investigated whether soil-feeding termites select their foraging sites to enhance nutrients intake. To do so, we collected termites and analysed the composition and structure of their feeding substrates. Although Anoplotermes-group members are all considered soil-feeders, our results show that some species specifically feed on abandoned termite nests and very rotten wood, and that this substrate selection is correlated with previous stable isotope analyses, suggesting that one component of niche differentiation among species is substrate selection. Our results show that the composition and structure of bare soils on which different termite species foraged do not differ, suggesting that there is no species specialization for a particular type of bare soil. Finally, the bare soil on which termites forage does not differ from random soil samples. Overall, our results suggest that few species of the Anoplotermes-group are specialized toward substrates rich in organic matter, but that the vast majority forage on soil independently of its structural and chemical composition, being ecologically equivalent for this factor.

Soil charcoal to assess the impacts of past human disturbances on tropical forests.

The canopy of many central African forests is dominated by light-demanding tree species that do not regenerate well under themselves. The prevalence of these species might result from ancient slash-and-burn agricultural activities that created large openings, while a decline of these activities since the colonial period could explain their deficit of regeneration. To verify this hypothesis, we compared soil charcoal abundance, used as a proxy for past slash-and-burn agriculture, and tree species composition assessed on 208 rainforest 0.2 ha plots located in three areas from Southern Cameroon. Species were classified in regeneration guilds (pioneer, non-pioneer light-demanding, shade-bearer) and characterized by their wood-specific gravity, assumed to reflect light requirement. We tested the correlation between soil charcoal abundance and: (i) the relative abundance of each guild, (ii) each species and family abundance and (iii) mean wood-specific gravity. Charcoal was found in 83% of the plots, indicating frequent past forest fires. Radiocarbon dating revealed two periods of fires: "recent" charcoal were on average 300 years old (up to 860 BP, n = 16) and occurred in the uppermost 20 cm soil layer, while "ancient" charcoal were on average 1900 years old (range: 1500 to 2800 BP, n = 43, excluding one sample dated 9400 BP), and found in all soil layers. While we expected a positive correlation between the relative abundance of light-demanding species and charcoal abundance in the upper soil layer, overall there was no evidence that the current heterogeneity in tree species composition can be explained by charcoal abundance in any soil layer. The absence of signal supporting our hypothesis might result from (i) a relatively uniform impact of past slash-and-burn activities, (ii) pedoturbation processes bringing ancient charcoal to the upper soil layer, blurring the signal of centuries-old Human disturbances, or (iii) the prevalence of other environmental factors on species composition.