First report of black leaf streak disease in bananas caused by Pseudocercospora fijiensis on Mauritius island.

Pseudocercospora fijiensis, the causal agent of the black leaf streak disease of bananas (plants in the genus Musa) (BLSD), is considered to be the major economic threat to export-banana cultivation (de Bellaire, Fouré, Abadie, & Carlier, 2010). The disease has a worldwide distribution throughout the humid tropical regions and has been previously reported in the Southwestndian Ocean (SWIO) area: in 1993 in Mayotte and Comoros islands (DR Jones & Mourichon, 1993), in 2000 in Madagascar (Jones, 2003; Rivas, Zapater, Abadie, & Carlier, 2004) and in 2018 in Reunion Island (Rieux et al., 2019). In Mauritius, the presence of Pseudocercospora fijiensis was suspected in 1996 (Soomary & Benimadhu, 1998) but has never been confirmed, as symptoms could have been confounded with Pseudocercospora musae or Pseudocercospora eumusae, two causal agents of others leaf spot diseases of banana which were previously described in Mauritius in 1959 (Orieux & Felix, 1968) and 2000 (Carlier, Zapater, Lapeyre, Jones, & Mourichon, 2000), respectively. In March 2022, typical BLSD symptoms were observed at relatively low prevalence in a Cavendish crop located in the "Balance John" area (site S1 on Fig. S1-A) of Mauritius island. Typical early symptoms (stages 2) were 1- to 4-mm long brown streaks at the abaxial leaf surface, and typical older streaks (stages 3 and 4) were also observed (Fig. S1-B). These symptoms were mixed with symptoms of ELSD caused by P. eumusae. Since both species cannot be clearly distinguished only on the description of symptoms, conidial sporulation on stages 2 was checked in the laboratory (Ngando et al., 2015) since P. eumusae does not produce conidia on these young stages. In April 2022, banana leaves bearing symptoms of leaf spot diseases were collected in 7 different sites (Fig. S1-A). All leaf fragments were sent to the CIRAD laboratories where molecular diagnosis was performed following the protocol developed by Arzanlou et al. (2007). In brief, genomic DNA was extracted from ground leaf fragments displaying symptoms using the DNeasy® Plant Mini Kit (Qiagen®, Courtaboeuf, France). At each site, a total of 6 lesions cut from 6 different leaves were pooled. The DNA extracts were added as templates for real-time PCR assay designed to specifically detect the presence of P. fijiensis, P. musae and P. eumusae using MFbf/MFbrtaq/MFbp, MEbf/MEbrtaq/FMep and MMbf/Mmbrtaq/FMep primers and probes, respectively (Arzanlou et al., 2007). Both positive and negative controls were included in the assay and every sample reaction was duplicated. P. fijiensis was detected from 2 out of 7 sites (S2 and S7, see Fig.S2-B). P. eumusae was detected at all sites while P. musae was found in one site only (S6). Interestingly, our results also showed coinfection by P. fijiensis - P. eumusae & P. musae - P. eumusae on several sites. The presence of P. fijiensis was further confirmed by several investigations performed on conidia isolated from S2 samples including i) morphological observations of conidia displaying P. fijiensis type description (Pérez-Vicente, Carreel, Roussel, Carlier, & Abadie (2021), Fig. S2-A), ii) DNA sequencing of 16S ribosomal gene with ITS1 & ITS4 primers (GenBank accessions Nos. OR515818-OR515810) with BLAST results displaying percentages of identity > 99.70% with type strains and iii) Koch's postulates were fulfilled by artificial inoculation of detached leaf pieces as described in Pérez-Vicente, Carreel, Roussel, Carlier, & Abadie (2021) (Fig. S2-D). In brief, for the artificial inoculation, symptoms obtained after inoculation of both a strain isolated in Mauritius (S2-MAU) and a positive control (T+) were compared and shown to be typical of P. fijiensis species for the 3 replicates. To the best of our knowledge, this is the first official report of P. fijiensis and BLSD in Mauritius Island. This revelation holds significant importance for both the agricultural and scientific communities, shedding light on the potential spread and impact of this devastating pathogen in previously unaffected regions. From a global perspective, this discovery underscores the interconnectedness of agricultural ecosystems and the need for vigilance in monitoring and responding to emerging plant diseases in an increasingly interconnected world (Vega et al. 2022). Future investigations will be required to monitor the spread of BLSD on the island, describe the genetic structure of populations and identify routes of invasion at the SWOI scale.

Ecological Regulation of Black Leaf Streak Disease Driven by Plant Richness in Banana Agroecosystems.

Black leaf streak disease (BLSD), caused by the fungus Mycosphaerella fijiensis, is an important threat to banana production. Although its control relies on costly and unsustainable use of fungicides, ecological regulation of BLSD linked to field-scale plant diversity has received little attention. We monitored banana phytometers in plots in banana-based fields where no fungicides were applied. Within each plot, we measured plant richness in three strata, canopy openness, necrotic leaf removal, Musa abundance and richness. We quantified ecological regulation of five BLSD parameters (inoculum sources, spore abundance, lesion density, incubation time, and the area under the disease progression curve) and identified, using structural equation modeling, the characteristics of the plant community and the mechanisms likely responsible for the regulation. Regulation occurred, but most effectively before lesion formation, and was mainly related to plant richness between 1.5 and 5m high. A barrier effect, rather than a dilution effect, more likely limited spore abundance. Our results support the hypothesis that the potential effects of plant richness on leaf-scale microclimate variability and on the diversity of epiphyllic microorganisms are involved in the regulation of incubation time and lesion density. Field-scale management of plant diversity may be a promising lever to foster ecological regulation of BLSD.

Assessing the effect of complex ground types on ground-dwelling arthropod movements with video monitoring: Dealing with concealed movements under a layer of plant residues.

Understanding the effect of ground types on foraging movements of ground-dwelling arthropods is a key step to managing their spatial distribution as required for successful conservation biological control. Indeed, fine movements at the centimeter scale can strongly influence the foraging ability of pest predators. However, because radio frequency identification or harmonic tracking techniques are not yet suitable for small species and video tracking focuses on uniform and light backgrounds, foraging movements have rarely been studied in relation to ground types. We present a method to track a ground-dwelling arthropod (the earwig Euborellia caraibea) at night, walking on two contrasted ground types: bare soil and soil partly covered with a stratum of banana plant residues allowing individuals to hide periodically. The tracking of individuals within these ground types was achieved by infrared light, tagging individuals, video treatments, and semi-automatic cleaning of trajectories. We tested different procedures to obtain segments with identical durations to quantify speeds and sinuosities. These procedures were characterized by the junction time gap between trajectory fragments, the rediscretization time of trajectories, and whether or not to use interpolation to fill in missing points in the trajectories. Earwigs exhibited significantly slower and more sinuous movements on soil with banana plant residues than on bare soil. Long time gaps for trajectory junction, extended rediscretization times, and interpolation were complementary means to integrate concealed movements in the trajectories. The highest slowdown in plant residues was detected when the procedure could account for longer periods under the residues. These results suggest that earwigs spent a significant amount of time concealed by the residues. Additionally, the residues strongly decreased the earwigs' movement. Since the technical solutions presented in this study are inexpensive, easy to set up, and replicate, they represent valuable contributions to the emerging field of video monitoring.

Caught on camera: Field imagery reveals the unexpected importance of vertebrates for biological control of the banana weevil (Cosmopolites sordidus Col. Curculionidae).

Understanding of ecological interactions is necessary for the application of biological control. Banana is the second most produced fruit worldwide and the banana weevil (Cosmopolites sordidus) is the most important pest of banana and plantain. Its biological control remains challenging because of the robustness and cryptic behaviour of the adult and the hidden development of larval stages. Researchers therefore tend to favour conservation biological control of this pest. The commonly used methods for measuring the effects of natural enemies on the regulation of this pest focus on invertebrates and may underestimate the role of vertebrates on biological control. Using cameras, we recorded the predation of sentinel adult weevils in banana plots in La Réunion island that differed in weevil infestation levels and in animal biodiversity. To facilitate image analysis, we used background subtraction to isolate moving parts of image sequences and thus detect predators and predation events. Our cameras recorded only vertebrates as predators of adult banana weevils. The most important predator appeared to be the Asian shrew (Suncus murinus), which was responsible for 67% of the predation events. Other predators included the house mouse (Mus musculus), the oriental garden lizard (Calotes versicolor), and the guttural toad (Sclerophrys gutturalis). The exact time of predation events were determined from the images metadata. It was thus possible to identify predator foraging periods that coincided with activity of adult weevils. Our results confirm that images provide useful information for biological and ecological studies. Along with other recent studies, our results suggest that the role of vertebrates in biological control may be underestimated. Based on these results, we advocate for several management implications such as the installation of hedges, grasslands, and ponds to favour these vertebrate predators of the banana weevil, possibly also favouring other vertebrate and invertebrate natural enemies.

Vegetation structure of plantain-based agrosystems determines numerical dominance in community of ground-dwelling ants.

In tropics, ants can represent an important part of animal biomass and are known to be involved in ecosystem services, such as pest regulation. Understanding the mechanisms underlying the structuring of local ant communities is therefore important in agroecology. In the humid tropics of Africa, plantains are cropped in association with many other annual and perennial crops. Such agrosystems differ greatly in vegetation diversity and structure and are well-suited for studying how habitat-related factors affect the ant community. We analysed abundance data for the six numerically dominant ant taxa in 500 subplots located in 20 diversified, plantain-based fields. We found that the density of crops with foliage at intermediate and high canopy strata determined the numerical dominance of species. We found no relationship between the numerical dominance of each ant taxon with the crop diversity. Our results indicate that the manipulation of the densities of crops with leaves in the intermediate and high strata may help maintain the coexistence of ant species by providing different habitat patches. Further research in such agrosystems should be performed to assess if the effect of vegetation structure on ant abundance could result in efficient pest regulation.

Dataset on the regulation of banana weevil abundance and corm damage associated with plant richness and the ground-dwelling arthropods' food web.

The data presented in this article are related to the research article entitled "Plant richness enhances banana weevil regulation in a tropical agroecosystem by affecting a multitrophic food web " [1]. It provides information about plant species richness, weevil corm damage and the abundance of different arthropod groups, including the banana weevil and its potential natural enemies and alternative preys.

Kin discrimination and altruism in the larvae of a solitary insect.

Kin selection theory predicts altruism between related individuals, which requires the ability to recognize kin from non-kin. In insects, kin discrimination associated with altruistic behaviour is well-known in clonal and social species but in very few solitary insects. Here, we report that the solitary larvae of a non-social insect Aleochara bilineata Gyll. (Coleoptera; Staphylinidae) show kin discrimination and sibling-directed altruistic behaviour. Larvae superparasitize more frequently the hosts parasitized by non-kin individuals than those hosts parasitized by siblings. Kin discrimination probably occurs by self-referent phenotype matching, where an individual compares its own phenotype with that of a non-familiar related individual, a mechanism rarely demonstrated in animals. The label used to recognize kin from non-kin corresponds to substances contained in the plug placed on the hosts by the resident larvae during the parasitization process. Kin competition induced by a limited larval dispersion may have favoured the evolution of kin recognition in this solitary species.

Data on the abundance of the banana weevil Cosmopolites sordidus and of the earwig Euborellia caraibea in bare soil and cover crop plots.

The data presented in this article are related to the research article entitled "Cover cropping reduces the abundance of the banana weevil Cosmopolites sordidus but does not reduce its damage to the banana plants" (Carval et al., in press) [1]. This article describes how the abundance of the banana weevil, Cosmopolites sordidus, and the abundance of the earwig Euborellia caraibea were affected by the addition of a cover crop. The field data set is made publicly available to enable critical or extended analyzes.

Dataset on the abundance of ants and Cosmopolites sordidus damage in plantain fields with intercropped plants.

The data presented in this article are related to the research article entitled "Ant abundance and Cosmopolites sordidus damage in plantain fields as affected by intercropping" (A.G. Dassou, D. Carval, S. Dépigny, G.H Fansi, P. Tixier, 2015) [1]. This article describes how associated crops maize (Zea mays), cocoyam (Xanthosoma sagittifolium) and bottle gourd (Lagenaria siceraria) intercropped in the plantain fields in Cameroun modify ant community structure and damages of banana weevil Cosmopolites sordidus. The field data set is made publicly available to enable critical or extended analyzes.

Balancing competition for resources with multiple pest regulation in diversified agroecosystems: a process-based approach to reconcile diversification and productivity.

Agroecosystem plant diversification can enhance pest biological regulation and is a promising alternative to pesticide application. However, the costs of competition for resources between plants may exceed the benefits gained by pest regulation. To disentangle the interactions between pest regulation and competition, we developed a generic process-based approach that accounts for the effects of an associated plant and leaf and root pests on biomass production. We considered three crop-plant associations that differ in competition profiles, and we simulated biomass production under wide ranges of both pest regulation rates and resources' availability. We analyzed outputs to quantify the pest regulation service level that would be required to attain monoculture yield and other production goals. Results showed that pest regulation requirements were highly dependent on the profile of resource interception of the associated plant and on resources' availability. Pest regulation and the magnitude of competition between plants interacted in determining the balance between nitrogen and radiation uptake by the crop. Our findings suggest that productivity of diversified agroecosystems relative to monoculture should be optimized by assembling plants whose characteristics balance crops' resource acquisition. The theoretical insights from our study draw generic rules for vegetation assemblage to optimize trade-offs between pest regulation and production. Our findings and approach may have implications in understanding, theorizing and implementing agroecosystem diversification. By its generic and adaptable structure, our approach should be useful for studying the effects of diversification in many agroecosystems.

Dominance in a ground-dwelling ant community of banana agroecosystem.

In tropical ecosystems, ants represent a substantial portion of the animal biomass and contribute to various ecosystem services, including pest regulation and pollination. Dominant ant species are known to determine the structure of ant communities by interfering in the foraging of other ant species. Using bait and pitfall trapping experiments, we performed a pattern analysis at a fine spatial scale of an ant community in a very simplified and homogeneous agroecosystem, that is, a single-crop banana field in Martinique (French West Indies). We found that the community structure was driven by three dominant species (Solenopsis geminata, Nylanderia guatemalensis, and Monomorium ebeninum) and two subdominant species (Pheidole fallax and Brachymyrmex patagonicus). Our results showed that dominant and subdominant species generally maintained numerical dominance at baits across time, although S. geminata, M. ebeninum, and B. patagonicus displayed better abilities to maintain dominance than P. fallax and N. guatemalensis. Almost all interspecific correlations between species abundances, except those between B. patagonicus and N. guatemalensis, were symmetrically negative, suggesting that interference competition prevails in this ground-dwelling ant community. However, we observed variations in the diurnal and nocturnal foraging activity and in the daily occurrence at baits, which may mitigate the effect of interference competition through the induction of spatial and temporal niche partitioning. This may explain the coexistence of dominant, subdominant, and subordinate species in this very simplified agroecosystem, limited in habitat structure and diversity.

Local and neighboring patch conditions alter sex-specific movement in banana weevils.

Understanding the mechanisms underlying the movements and spread of a species over time and space is a major concern of ecology. Here, we assessed the effects of an individual's sex and the density and sex ratio of conspecifics in the local and neighboring environment on the movement probability of the banana weevil, Cosmopolites sordidus. In a "two patches" experiment, we used radiofrequency identification tags to study the C. sordidus movement response to patch conditions. We showed that local and neighboring densities of conspecifics affect the movement rates of individuals but that the density-dependent effect can be either positive or negative depending on the relative densities of conspecifics in local and neighboring patches. We demonstrated that sex ratio also influences the movement of C. sordidus, that is, the weevil exhibits nonfixed sex-biased movement strategies. Sex-biased movement may be the consequence of intrasexual competition for resources (i.e., oviposition sites) in females and for mates in males. We also detected a high individual variability in the propensity to move. Finally, we discuss the role of demographic stochasticity, sex-biased movement, and individual heterogeneity in movement on the colonization process.

A unified model for the coevolution of resistance, tolerance, and virulence.

We present a general host-parasite model that unifies previous theory by investigating the coevolution of virulence, resistance, and tolerance, with respect to multiple physiological, epidemiological, and environmental parameters. Four sets of new predictions emerge. First, compared to virulence coevolving with resistance or tolerance, three-trait coevolution promotes more virulence and less tolerance, and broadens conditions under which pure defenses evolve. Second, the cost and efficiency of virulence and the epidemiological rates are the key factors of virulence coevolving with resistance and tolerance. Maximum virulence evolves for intermediate infection rate, at which coevolved levels of resistance and tolerance are both high. The influence of host and parasite background mortalities is strong on the evolution of defenses and weak on the coevolution of virulence. Third, evolutionary correlations between defenses can switch sign along single-parameter gradients. The evolutionary trade-off between resistance and tolerance may coevolve with virulence that either increases or decreases monotonically, depending on the underlying parameter gradient. Fourth, despite global attractiveness and stability of coevolutionary equilibria, not-so-rare and not-so-small mutations can beget large variation in virulence and defenses around equilibrium, in the form of transient "evolutionary spikes." Implications for evolutionary management of infections are discussed and directions for future research are outlined.