Identification of a queen primer pheromone in higher termites.

It is long established that queens of social insects, including termites, maintain their reproductive dominance with queen primer pheromones (QPPs). Yet, the QPP chemistry has only been elucidated in a single species of lower termites. By contrast, the most diversified termite family Termitidae (higher termites), comprising over 70% of termite species, has so far resisted all attempts at QPP identification. Here, we show that the queen- and egg-specific sesquiterpene (3R,6E)-nerolidol acts as the QPP in the higher termite Embiratermes neotenicus. This species has a polygynous breeding system, in which the primary queen is replaced by multiple neotenic queens of parthenogenetic origin. We demonstrate that (3R,6E)-nerolidol suppresses the development of these parthenogenetic queens and thus mimics the presence of mature queen(s). It acts as an airborne signal and may be used to optimize the number of queens, thus being the key regulatory element in the special breeding system of E. neotenicus.

Insights into coastal phytoplankton variations from 1979 to 2018 derived from Ba/Ca records in scallop shells (Chlamys islandica) from a fishing ground in the northern Gulf of St. Lawrence.

Bivalve growth is affected by phytoplankton quality and availability, but long-term, coastal environmental time series related to these parameters are often lacking. Therefore, it is crucial to develop methods to accurately quantify trends in phytoplankton dynamics over time. This would be especially important for the fished scallop beds in the northern Gulf of St. Lawrence, where landings sharply declined since the early-2000s. Over the past decade, many studies have highlighted the potential of Ba/Ca ratios in bivalve shells as an environmental proxy for phytoplankton dynamics. This study presents records of Ba/Ca ratios in 31 young Chlamys islandica shells sampled in the Mingan Archipelago from 1979 to 2018. The Ba/Ca master chronology showed a decreasing trend since 2002, which could reflect changes in local phytoplankton bloom taxonomic composition, and coincides with the aforementioned decline in scallop landings. Investigations of environmental controls on barium incorporation into the shells highlight the importance of bottom, nutrient-rich waters to support diatom production or export in this fishing area. The use of such high-resolution seasonal records extracted from bio-archives may identify essential environmental mechanisms that will then assist with the development of an ecosystem-based fishery management strategy.

[Training session on healthy environments: evaluation of an intervention for local stakeholders]. / Sensibiliser aux environnements favorables aux saines habitudes de vie : évaluation d’une intervention auprès des personnes-relais.

BACKGROUND: Around the world, various interventions have been developed to encourage the adoption of healthy lifestyles, particularly nutrition and physical activity. Physical, political, economic and socio-cultural environments have a major influence on individual attitudes in relation to healthy lifestyle. However, stakeholders with the greatest impact on improving these environments are not always well informed about the theory and their roles on the creation of environments favourable to healthy lifestyles. Various stakeholders from the province of Quebec were therefore invited to attend training sessions in order to prepare them to act on these four environments. OBJECTIVES: 1) To describe the perceptions of the stakeholders who attended these sessions concerning the content and teaching methods and 2) to identify stakeholders' changes of perceptions and practices following the training session. METHODS: Twelve (12) focus groups and 52 individual interviews were conducted across Quebec with stakeholders who attended a training session. RESULTS: Our results indicate increased awareness of stakeholders on the importance of their role but also the need to more precisely target those aspects requiring increased awareness. A content better suited to the level of expertise is therefore proposed to maximize the benefits of these training sessions. CONCLUSION: Training sessions must be addressed to influential stakeholders with a limited knowledge on the subject, which is often the case for municipal decision-makers known to play a major role in promoting environments favourable to healthy eating and physical activity.

The role of the glucose-sensing transcription factor carbohydrate-responsive element-binding protein pathway in termite queen fertility.

Termites are among the few animals that themselves can digest the most abundant organic polymer, cellulose, into glucose. In mice and Drosophila, glucose can activate genes via the transcription factor carbohydrate-responsive element-binding protein (ChREBP) to induce glucose utilization and de novo lipogenesis. Here, we identify a termite orthologue of ChREBP and its downstream lipogenic targets, including acetyl-CoA carboxylase and fatty acid synthase. We show that all of these genes, including ChREBP, are upregulated in mature queens compared with kings, sterile workers and soldiers in eight different termite species. ChREBP is expressed in several tissues, including ovaries and fat bodies, and increases in expression in totipotent workers during their differentiation into neotenic mature queens. We further show that ChREBP is regulated by a carbohydrate diet in termite queens. Suppression of the lipogenic pathway by a pharmacological agent in queens elicits the same behavioural alterations in sterile workers as observed in queenless colonies, supporting that the ChREBP pathway partakes in the biosynthesis of semiochemicals that convey the signal of the presence of a fertile queen. Our results highlight ChREBP as a likely key factor for the regulation and signalling of queen fertility.

Profiling the Succession of Bacterial Communities throughout the Life Stages of a Higher Termite Nasutitermes arborum (Termitidae, Nasutitermitinae) Using 16S rRNA Gene Pyrosequencing.

Previous surveys of the gut microbiota of termites have been limited to the worker caste. Termite gut microbiota has been well documented over the last decades and consists mainly of lineages specific to the gut microbiome which are maintained across generations. Despite this intimate relationship, little is known of how symbionts are transmitted to each generation of the host, especially in higher termites where proctodeal feeding has never been reported. The bacterial succession across life stages of the wood-feeding higher termite Nasutitermes arborum was characterized by 16S rRNA gene deep sequencing. The microbial community in the eggs, mainly affiliated to Proteobacteria and Actinobacteria, was markedly different from the communities in the following developmental stages. In the first instar and last instar larvae and worker caste termites, Proteobacteria and Actinobacteria were less abundant than Firmicutes, Bacteroidetes, Spirochaetes, Fibrobacteres and the candidate phylum TG3 from the last instar larvae. Most of the representatives of these phyla (except Firmicutes) were identified as termite-gut specific lineages, although their relative abundances differed. The most salient difference between last instar larvae and worker caste termites was the very high proportion of Spirochaetes, most of which were affiliated to the Treponema Ic, Ia and If subclusters, in workers. The results suggest that termite symbionts are not transmitted from mother to offspring but become established by a gradual process allowing the offspring to have access to the bulk of the microbiota prior to the emergence of workers, and, therefore, presumably through social exchanges with nursing workers.

Asexual queen succession in the higher termite Embiratermes neotenicus.

Asexual queen succession (AQS), in which workers, soldiers and dispersing reproductives are produced sexually while numerous non-dispersing queens arise through thelytokous parthenogenesis, has recently been described in three species of lower termites of the genus Reticulitermes. Here, we show that AQS is not an oddity restricted to a single genus of lower termites, but a more widespread strategy occurring also in the most advanced termite group, the higher termites (Termitidae). We analysed the genetic structure in 10 colonies of the Neotropical higher termite Embiratermes neotenicus (Syntermitinae) using five newly developed polymorphic microsatellite loci. The colonies contained one primary king accompanied either by a single primary queen or by up to almost 200 neotenic queens. While the workers, the soldiers and most future dispersing reproductives were produced sexually, the non-dispersing neotenic queens originated through thelytokous parthenogenesis of the founding primary queen. Surprisingly, the mode of thelytoky observed in E. neotenicus is most probably automixis with central fusion, contrasting with the automixis with terminal fusion documented in Reticulitermes. The occurrence of AQS based on different mechanisms of ploidy restoration raises the hypothesis of an independent evolutionary origin of this unique reproductive strategy in individual lineages of lower and higher termites.

The distribution of Wolbachia in Cubitermes (Termitidae, Termitinae) castes and colonies: a modelling approach.

Wolbachia are endosymbiotic bacteria of arthropods and nematodes that are able to manipulate host reproduction. Although vertically transmitted via the cytoplasm in eggs, horizontal transmission of Wolbachia among and within arthropod species has been shown to be common. Eusocial insects represent interesting models for studying Wolbachia transmission due to colonial organization and close interaction between nestmates. Here we conducted a detailed screening of Wolbachia infection for 15 colonies of the very common soil-feeding termites Cubitermes spp. affinis subarquatus (Termitidae, Termitinae) that consist of four distinct phylogenetic species in the Lopé forest Reserve, Gabon. Infection tests showed that 50% of the individuals were Wolbachia positive (N = 555) with 90% of reproductives and 48% of offspring infected. White soldiers, which are transitional stages preceding mature soldiers, had a significantly higher mean infection rate (74%) than the other castes and stages (63%, 33% and 39% for larvae, workers and mature soldiers, respectively). We used a maximum likelihood method and Akaike's Information Criterion in order to explain the non-expected high rate of Wolbachia infection in white soldiers. The best model included a species effect for the stochastic loss of Wolbachia and a caste effect for the rate of gain. After fitting, the best model selected was for a species-specific rate of loss with a null rate of new gain for larvae, workers and soldiers and a probability of 0.72 whatever the species, that a white soldier becomes newly contaminated during that stage. The mean expected infection rate in white soldiers without a new gain was estimated to 17% instead of the 74% observed. Here we discuss the possible explanations to the high infection rate observed in white soldiers.

Environmental drivers of the Canadian Arctic megabenthic communities.

Environmental gradients and their influence on benthic community structure vary over different spatial scales; yet, few studies in the Arctic have attempted to study the influence of environmental gradients of differing spatial scales on megabenthic communities across continental-scales. The current project studied for the first time how megabenthic community structure is related to several environmental factors over 2000 km of the Canadian Arctic, from the Beaufort Sea to northern Baffin Bay. Faunal trawl samples were collected between 2007 and 2011 at 78 stations from 30 to 1000 m depth and patterns in biomass, density, richness, diversity, and taxonomic composition were examined in relation to indirect/spatial gradients (e.g., depth), direct gradients (e.g., bottom oceanographic variables), and resource gradients (e.g., food supply proxies). Six benthic community types were defined based on their biomass-based taxonomic composition. Their distribution was significantly, but moderately, associated with large-scale (100-1000 km) environmental gradients defined by depth, physical water properties (e.g., bottom salinity), and meso-scale (10-100 km) environmental gradients defined by substrate type (hard vs. soft) and sediment organic carbon content. We did not observe a strong decline of bulk biomass, density and richness with depth or a strong increase of those community characteristics with food supply proxies, contrary to our hypothesis. We discuss how local- to meso-scale environmental conditions, such as bottom current regimes and polynyas, sustain biomass-rich communities at specific locations in oligotrophic and in deep regions of the Canadian Arctic. This study demonstrates the value of considering the scales of variability of environmental gradients when interpreting their relevance in structuring of communities.

Species delimitation and phylogeny in the genus Nasutitermes (Termitidae: Nasutitermitinae) in French Guiana.

Species delimitation and identification can be arduous for taxa whose morphologic characters are easily confused, which can hamper global biodiversity assessments and pest species management. Exploratory methods of species delimitation that use DNA sequence as their primary information source to establish group membership and estimate putative species boundaries are useful approaches, complementary to traditional taxonomy. Termites of the genus Nasutitermes make interesting models for the application of such methods. They are dominant in Neotropical primary forests but also represent major agricultural and structural pests. Despite the prevalence, pivotal ecological role and economical impact of this group, the taxonomy of Nasutitermes species mainly depends on unreliable characters of soldier external morphology. Here, we generated robust species hypotheses for 79 Nasutitermes colonies sampled throughout French Guiana without any a priori knowledge of species affiliation. Sequence analysis of the mitochondrial cytochrome oxidase II gene was coupled with exploratory species-delimitation tools, using the automatic barcode gap discovery method (ABGD) and a generalized mixed Yule-coalescent model (GMYC) to propose primary species hypotheses (PSHs). PSHs were revaluated using phylogenetic analyses of two more loci (mitochondrial 16S rDNA and nuclear internal transcribed spacer 2) leading to 16 retained secondary species hypotheses (RSSH). Seven RSSHs, represented by 44/79 of the sampled colonies, were morphologically affiliated to species recognized as pests in the Neotropics, where they represent a real invasive pest potential in the context of growing ecosystem anthropization. Multigenic phylogenies based on combined alignments (1426-1784 bp) were also reconstructed to identify ancestral ecological niches and major-pest lineages, revealing that Guyanese pest species do not form monophyletic groups.

Regulation of soil organic C mineralisation at the pore scale.

Little is known about the factors that regulate C mineralisation at the soil pore scale or how these factors vary throughout the pore network. This study sought to understand how the decomposition of organic carbon varies within the soil pore network and to determine the relative importance of local environmental properties relative to biological properties as controlling factors. This was achieved by sterilising samples of soil and reinoculating them with axenic bacterial suspensions using the matric potential to target different locations in the pore network. Carbon mineralisation curves were described with two-compartment first-order models to distinguish CO2 derived from the labile organic carbon released during sterilisation from CO2 derived from organic C unaffected by sterilisation. The data indicated that the size of the labile pool of organic C, possibly of microbial origin, varied as a function of location in the pore network but that the organic carbon unaffected by sterilisation did not. The mineralisation rate of the labile C varied with the bacterial type inoculated, but the mineralisation rate of the organic C unaffected by sterilisation was insensitive to bacterial type. Taken together, the results suggest that microbial metabolism is a less significant regulator of soil organic carbon decomposition than are microbial habitat properties.

Beaver ponds increase methylmercury concentrations in Canadian shield streams along vegetation and pond-age gradients.

Beaver impoundments flood forested areas and may be important production sites for methylmercury (MeHg) because of the resulting enhanced microbial activity and oxygen depletion. The influence of 17 beaver impoundments on streamwater chemistry (total mercury (THg), MeHg, nutrients, cations, and anions)] was investigated by sampling sites located along vegetation and pond-age gradients in southwestern Quebec (Canada). Recently inundated beaver ponds (< 10 years old) and those located in coniferous watersheds had the highest MeHg concentrations (range, 0.10-4.53 ng L(-1)) and greatest methylation efficiencies (% THg as MeHg; range, 10-74%). High heterotrophic activity likely occurred in the beaver ponds as suggested by depletions of dissolved oxygen, sulfate and nitrite-nitrate concentrations, and increases in nutrients (e.g., dissolved organic carbon, total phosphorus, and total nitrogen) in outlets compared to inlets. Acidic waters at coniferous sites may have stimulated more MeHg production than in mixed woodland regions. Lower methylation efficiencies in older ponds (> 20 years old) may be due to the degradation of less labile organic matter as ponds age. Beavers actively alter watersheds by building impoundments, and our findings indicate that this landscape disturbance may be a significant source of MeHg to downstream water bodies.

Diversity of Wolbachia isolated from the Cubitermes sp. affinis subarquatus complex of species (Termitidae), revealed by multigene phylogenies.

Wolbachia are endosymbiotic bacteria that may alter the reproductive mechanisms of arthropod hosts. Eusocial termites provide considerable scope for Wolbachia studies owing to their ancient origin, their great diversity and their considerable ecological, biological and behavioral plasticity. This article describes the phylogenetic distribution of Wolbachia infecting termites of the Cubitermes genus, which are particularly abundant soil-feeders in equatorial Africa. Fourteen colonies of the Cubitermes sp. affinis subarquatus complex of species were screened using five bacterial genes (wsp, ftsZ, coxA, fbpA and 16S rRNA genes) and a striking diversity of Wolbachia strains was identified within these closely related species. In the host complex, three Wolbachia variants were found that were not in the super groups usually reported for termites (F and H), each infecting one or two Cubitermes species.

Genetic differentiation in the soil-feeding termite Cubitermes sp. affinis subarquatus: occurrence of cryptic species revealed by nuclear and mitochondrial markers.

BACKGROUND: Soil-feeding termites are particularly interesting models for studying the effects of fragmentation, a natural or anthropic phenomenon described as promoting genetic differentiation. However, studying the link between fragmentation and genetics requires a method for identifying species unambiguously, especially when morphological diagnostic characters are lacking. In humivorous termites, which contribute to the fertility of tropical soils, molecular taxonomy and phylogenetic relationships are rarely studied, though mitochondrial and nuclear molecular markers are widely used in studies of pest termites. Here, we attempt to clarify the taxonomy of soil-feeding colonies collected throughout the naturally fragmented Lopé Reserve area (Gabon) and morphologically affiliated to Cubitermes sp. affinis subarquatus. The mitochondrial gene of cytochrome oxidase II (COII), the second nuclear rDNA internal transcribed spacer (ITS2) and five microsatellites were analyzed in 19 colonies. RESULTS: Bayesian Inference, Maximum Likelihood and Maximum Parsimony phylogenetic analyses, which were applied to the COII and ITS2 sequences, and Neighbor-Joining reconstructions, applied to the microsatellite data, reveal four major lineages in the Cubitermes sp. affinis subarquatus colonies. The concordant genealogical pattern of these unlinked markers strongly supports the existence of four cryptic species. Three are sympatric in the Reserve and are probably able to disperse within a mosaic of forests of variable ages and savannahs. One is limited to a very restricted gallery forest patch located in the North, outside the Reserve. CONCLUSION: Our survey highlights the value of combined mitochondrial and nuclear markers for exploring unknown groups such as soil-feeding termites, and their relevance for resolving the taxonomy of organisms with ambiguous morphological diagnostic characters.