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Infection dynamics in vertebrates are driven by biological and ecological processes. For bats, population structure and reproductive cycles have major effects on RNA virus transmission. On Reunion Island, previous studies have shown that parturition of pregnant females and aggregation of juvenile Reunion free-tailed bats (Mormopterus francoismoutoui) are associated with major increase in the prevalence of bats shedding RNA viruses. The synchronicity of such shedding pulses, however, is yet to be assessed between viruses but also maternity colonies. Based on 3422 fresh faeces collected every 2–5 weeks during four consecutive birthing seasons, we report the prevalence of bats shedding astroviruses (AstVs), coronaviruses (CoVs) and paramyxoviruses (PMVs) in two maternity colonies on Reunion Island. We found that the proportion of bats shedding viruses is highly influenced by sampling collection periods, and therefore by the evolution of the population age structure. We highlight that virus shedding patterns are consistent among years and colonies for CoVs and to a lesser extent for PMVs, but not for AstVs. We also report that 1% of bats harbour co-infections, with two but not three of the viruses, and most co-infections were due to CoVs and PMVs.

Co-infections have a key role in virus transmission in wild reservoir hosts. We investigated the simultaneous presence of astroviruses, coronaviruses, and paramyxoviruses in bats from Madagascar, Mayotte, Mozambique, and Reunion Island. A total of 871 samples from 28 bat species representing 8 families were tested by polymerase chain reactions (PCRs) targeting the RNA-dependent RNA-polymerase genes. Overall, 2.4% of bats tested positive for the presence of at least two viruses, only on Madagascar and in Mozambique. Significant variation in the proportion of co-infections was detected among bat species, and some combinations of co-infection were more common than others. Our findings support that co-infections of the three targeted viruses occur in bats in the western Indian Ocean region, although further studies are needed to assess their epidemiological consequences. Supplementary Information The online version contains supplementary material available at 10.1186/s12985-021-01673-2.

Leptospirosis, an environmental infectious disease of bacterial origin, is the infectious disease with the highest associated mortality in Seychelles. In small island territories, the occurrence of the disease is spatially heterogeneous and a better understanding of the environmental factors that contribute to the presence of the bacteria would help implement targeted control. The present study aimed at identifying the main environmental parameters correlated with animal reservoirs distribution and Leptospira infection in order to delineate habitats with highest prevalence. We used a previously published dataset produced from a large collection of rodents trapped during the dry and wet seasons in most habitats of Mah&eacute, the main island of Seychelles. A land use/land cover analysis was realized in order to describe the various environments using SPOT-5 images by remote sensing (object-based image analysis). At each sampling site, landscape indices were calculated and combined with other geographical parameters together with rainfall records to be used in a multivariate statistical analysis. Several environmental factors were found to be associated with the carriage of leptospires in Rattus rattus and Rattus norvegicus, namely low elevations, fragmented landscapes, the proximity of urbanized areas, an increased distance from forests and, above all, increased precipitation in the three months preceding trapping. The analysis indicated that Leptospira renal carriage could be predicted using the species identification and a description of landscape fragmentation and rainfall, with infection prevalence being positively correlated with these two environmental variables. This model may help decision makers in implementing policies affecting urban landscapes and/or in balancing conservation efforts when designing pest control strategies that should also aim at reducing human contact with Leptospira-laden rats while limiting their impact on the autochthonous fauna.

Bats provide key ecosystem services such as crop pest regulation, pollination, seed dispersal, and soil fertilization. Bats are also major hosts for biological agents responsible for zoonoses, such as coronaviruses (CoVs). The islands of the Western Indian Ocean are identified as a major biodiversity hotspot, with more than 50 bat species. In this study, we tested 1,013 bats belonging to 36 species from Mozambique, Madagascar, Mauritius, Mayotte, Reunion Island and Seychelles, based on molecular screening and partial sequencing of the RNA-dependent RNA polymerase gene. In total, 88 bats (8.7%) tested positive for coronaviruses, with higher prevalence in Mozambican bats (20.5% ± 4.9%) as compared to those sampled on islands (4.5% ± 1.5%). Phylogenetic analyses revealed a large diversity of α- and β-CoVs and a strong signal of co-evolution between CoVs and their bat host species, with limited evidence for host-switching, except for bat species sharing day roost sites.ImportanceThis is the first study to report the presence of coronaviruses (CoVs) in bats in Mayotte, Mozambique and Reunion Island, and in insectivorous bats in Madagascar. Eight percent of the tested bats were positive for CoVs, with higher prevalence in continental Africa than on islands. A high genetic diversity of α- and β-CoVs was found, with strong association between bat host and virus phylogenies, supporting a long history of co-evolution between bats and their associated CoVs in the Western Indian Ocean. These results highlight that strong variation between islands does exist and is associated with the composition of the bat species community on each island. Future studies should investigate whether CoVs detected in these bats have a potential for spillover in other hosts.

International audience; Pteropus livingstonii and Pteropus seychellensis comorensis are endemic fruit bat species that are among the most threatened animals in the Comoros archipelago. Both species are pollinators and seed dispersers of native and cultivated plants and are thus of crucial importance for the regeneration of natural forests as well as for cultivated plantations. However, these species are subject to strong anthropogenic pressures and face one of the highest rates of natural habitat loss reported worldwide. Yet little is known about the population genetic structure of these two species, making it difficult to define relevant conservation strategies. In this study, we investigated for the two flying fox species (1) the level of genetic diversity within islands, as well as across the archipelago and (2) the genetic structure between the two islands (Anjouan and Moheli) for P. livingstonii and between the four islands of the archipelago (Anjouan, Moheli, Grande Comore and Mayotte) for P. s. comorensis using mitochondrial and microsatellite markers. The results revealed contrasting patterns of genetic structure, with P. s. comorensis showing low genetic structure between islands, whereas P. livingstonii exhibited high levels of inter-island genetic differentiation. Overall, the genetic analyses showed low genetic diversity for both species. These contrasting genetic patterns may be the result of different dispersal patterns and the populations' evolutionary histories. Our findings lead us to suggest that in terms of conservation strategy, the two populations of P. livingstonii (on Anjouan and Moheli islands) should be considered as two separate management units. We recommend focusing conservation efforts on the Anjouan population, which is the largest, exhibits the highest genetic diversity, and suffers the greatest anthropogenic pressure. As for P. s. comorensis, its four populations could be considered as a single unit for conservation management purposes. For this species, we recommend protecting roosting trees to reduce population disturbance.

Understanding the processes driving parasite assemblages is particularly important in the context of zoonotic infectious diseases. Leptospirosis is a widespread zoonotic bacterial infection caused by pathogenic species of the genus Leptospira. Despite a wide range of animal hosts, information is still lacking on the factors shaping Leptospira diversity in wild animal communities, especially in regions, such as tropical insular ecosystems, with high host species richness and complex biogeographical patterns. Using a large dataset (34 mammal species) and a multilocus approach at a regional scale, we analyzed the role of both host species diversity and geography in Leptospira genetic diversity in terrestrial small mammals (rodents, tenrecs, and shrews) and bats from 10 different islands/countries in the western Indian Ocean (WIO) and neighboring Africa. At least four Leptospira spp. (L. interrogans, L. borgpetersenii, L. kirschneri, and L. mayottensis) and several yet-unidentified genetic clades contributed to a remarkable regional Leptospira diversity, which was generally related to the local occurrence of the host species rather than the geography. In addition, the genetic structure patterns varied between Leptospira spp., suggesting different evolutionary histories in the region, which might reflect both in situ diversification of native mammals (for L. borgpetersenii) and the more recent introduction of non-native host species (for L. interrogans). Our data also suggested that host shifts occurred between bats and rodents, but further investigations are needed to determine how host ecology may influence these events.

Background Leptospirosis is a bacterial zoonosis caused by pathogenic Leptospira for which rats are considered as the main reservoir. Disease incidence is higher in tropical countries, especially in insular ecosystems. Our objectives were to determine the current burden of leptospirosis in Seychelles, a country ranking first worldwide according to historical data, to establish epidemiological links between animal reservoirs and human disease, and to identify drivers of transmission. Methods A total of 223 patients with acute febrile symptoms of unknown origin were enrolled in a 12-months prospective study and tested for leptospirosis through real-time PCR, IgM ELISA and MAT. In addition, 739 rats trapped throughout the main island were investigated for Leptospira renal carriage. All molecularly confirmed positive samples were further genotyped. Results A total of 51 patients fulfilled the biological criteria of acute leptospirosis, corresponding to an annual incidence of 54.6 (95% CI 40.7–71.8) per 100,000 inhabitants. Leptospira carriage in Rattus spp. was overall low (7.7%) but dramatically higher in Rattus norvegicus (52.9%) than in Rattus rattus (4.4%). Leptospira interrogans was the only detected species in both humans and rats, and was represented by three distinct Sequence Types (STs). Two were novel STs identified in two thirds of acute human cases while noteworthily absent from rats. Conclusions This study shows that human leptospirosis still represents a heavy disease burden in Seychelles. Genotype data suggests that rats are actually not the main reservoir for human disease. We highlight a rather limited efficacy of preventive measures so far implemented in Seychelles. This could result from ineffective control measures of excreting animal populations, possibly due to a misidentification of the main contaminating reservoir(s). Altogether, presented data stimulate the exploration of alternative reservoir animal hosts., Author summary Leptospirosis is an emerging environmental infectious disease caused by corkscrew shaped bacteria called Leptospira. Humans usually get infected during recreational or work-related outdoor activities through contact with urine excreted by animal reservoirs. As a zoonotic disease, leptospirosis is a good example of the One Health concept for it links humans, animals and ecosystems in a web of pathogen maintenance and transmission. This zoonosis is highly prevalent in the tropics and especially in tropical islands. Seychelles archipelago has been reported as the country with highest human incidence worldwide, although figures are based on dated studies and/or poorly specific tests. The presented investigation aimed at providing an updated information on human leptospirosis burden in Seychelles and exploring the transmission chains in their environmental aspects. Presented data confirms that the disease still heavily impacts the country. Genotyping of pathogenic Leptospira in human acute cases reveals that three distinct Sequence Types (STs) are involved in the disease. However, rats typically considered as the main reservoir in Seychelles, harbor only one of these STs, found only in a minority of human cases. Hence, it appears that rats are likely not the main reservoir of leptospirosis in Seychelles, which has important consequences in terms of preventive measures to be implemented for a better control of human leptospirosis.

Background Although leptospirosis is a zoonosis of major concern on tropical islands, the molecular epidemiology of the disease aiming at linking human cases to specific animal reservoirs has been rarely explored within these peculiar ecosystems. Methodology/Principal Findings Five species of wild small mammals (n = 995) as well as domestic animals (n = 101) were screened for Leptospira infection on Reunion Island; positive samples were subsequently genotyped and compared to Leptospira from clinical cases diagnosed in 2012–2013 (n = 66), using MLST analysis. We identified two pathogenic species in human cases, namely Leptospira interrogans and Leptospira borgpetersenii. Leptospira interrogans was by far dominant both in clinical samples (96.6%) and in infected animal samples (95.8%), with Rattus spp and dogs being its exclusive carriers. The genetic diversity within L. interrogans was apparently limited to two sequence types (STs): ST02, identified among most clinical samples and in all rats with complete MLST, and ST34, identified in six humans, but not in rats. Noteworthy, L. interrogans detected in two stray dogs partially matched with ST02 and ST34. Leptospira borgpetersenii was identified in two clinical samples only (3.4%), as well as in cows and mice; four haplotypes were identified, of which two seemingly identical in clinical and animal samples. Leptospira borgpetersenii haplotypes detected in human cases were clearly distinct from the lineage detected so far in the endemic bat species Mormopterus francoismoutoui, thus excluding a role for this volant mammal in the local human epidemiology of the disease. Conclusions/Significance Our data confirm rats as a major reservoir of Leptospira on Reunion Island, but also pinpoint a possible role of dogs, cows and mice in the local epidemiology of human leptospirosis. This study shows that a comprehensive molecular characterization of pathogenic Leptospira in both clinical and animal samples helps to gaining insight into leptospirosis epidemiology within a specific environmental setting., Author Summary Leptospirosis is a zoonosis caused by infection with pathogenic Leptospira species. A broad range of animals, including rodents, pets and livestock, are maintenance hosts for leptospires. However, assessing the relative importance of each host in the contamination of the environment and, in fine, in the infection of humans, has rarely been performed. In this study, we surveyed various wild and domestic animal species and their Leptospira carriage in Reunion Island, where human leptospirosis is endemic. We determined and compared the Leptospira genetic diversity at the species and infra-species levels in laboratory-confirmed human cases and in infected animals. The two Leptospira species infecting humans, Leptospira interrogans and Leptospira borgpetersenii, could be traced back to different animal species: rats and dogs for the former species, cows and mice for the latter. The Leptospira infecting the single bat species endemic to the island was not found to be involved in human leptospirosis. Aside from rats, which were expected to play a role in the local epidemiology of the disease, the putative role of dogs, cattle and mice in human epidemiology on Reunion Island, pinpointed by our data, deserves a specific investigation. These results have strong implications in terms of local control actions aimed at reducing the burden of human leptospirosis.

Salmonellosis is an economic burden to the livestock industry in Reunion Island. In this study, we wanted to improve our understanding of Salmonella epidemiology by studying the wild fauna of Reunion Island. We assessed Salmonella diversity in small non-flying mammals, birds and cockroaches in order to evaluate their potential role in the epidemiology of Salmonella. A total of 268 samples were collected from cockroaches, small mammals and birds. The bacteriological analyses revealed that 11.7% of non-flying mammals and 25% of cockroaches tested were Salmonella infected; two wild bird species were also detected positive. The 128 Salmonella isolates were distributed in fifteen serotypes and the most predominant were S. 4,[5],12:i:- (21.9% of positive samples) followed by S. Enteritidis (15.6%), S. Typhimurium (15.6%), S. Infantis (12.5%) and S. Weltevreden (12.5%). A total of 27 XbaI profiles were identified using pulsed-field gel electrophoresis. Comparison of these Salmonella strains with our collection of Salmonella isolated from pigs and pig farm environments at the same period revealed 14 strains in common between wild fauna and pigs, especially for cockroaches. Our results suggest that wild fauna of Reunion Island could be infected by strains of Salmonella also isolated from pigs or pig environment. They may play a role in both persistence and spreading of Salmonella and therefore, could be a source of infection in pig farms. Pest control against cockroaches could be a helpful tool in the reduction of Salmonella infection of pigs, limiting contacts between wild fauna and both pigs and pig environment. Special attention should be paid to S. 4,[5],12:i:- since it was predominant in Reunion Island’s wild fauna and pigs and was the third most frequently reported serotype in human salmonellosis in Europe.

International audience; Connaître la distribution des espèces vectrices de zoonoses est fondamentale pour comprendre les dynamiques épidémiologiques et caractériser les zones à risque. Cette distribution peut être appréhendée à différentes échelles : régionalement, l'aire de répartition de ces espèces définit les limites d'occurrence de la transmission d'agents pathogènes et, localement, leurs niches déterminent l'aléa. Cette dernière échelle requiert une connaissance très précise des milieux qui peut être acquise par l'analyse d'images satellite à très haute résolution spatiale. Dans le cadre d'un projet de recherche sur la leptospirose à La Réunion (LeptOI), cette étude propose de mesurer le potentiel d'utilisation d'une image Pléiades (produit rééchantillonné à 50 cm) pour caractériser la distribution locale de différents petits mammifères terrestres autour de l'Etang de Saint-Paul. Trois campagnes de piégeage ont permis la capture de 134 animaux appartenant à quatre des cinq espèces présentes à La Réunion (Rattus rattus, R. norvegicus, Suncus murinus et Tenrec ecaudatus). Une analyse orientée-objet de l'image Pléiades a permis de discriminer dix classes d'occupation des sols avec une précision totale de 83,6%. Des indices paysagers ont été calculés à partir de cette classification autour de chacun des pièges (distances les plus courtes aux classes d'occupation, densité de contours et pourcentages de surface des classes au sein de zones tampon de 50 et de 100 mètres). Trois zones (« humide », « anthropisée » et « sèche ») sont clairement discriminées par ces variables, qui pourront être utilisées comme déterminants écologiques de différents profils épidémiologiques : la zone humide étant propice à la leptospirose et la zone anthropisée étant un lieu de cooccurrence des quatre espèces et ainsi potentiellement une zone d'échanges de pathogènes. Abstract Knowing the distribution of vectors of zoonoses is fundamental to understand epidemiological dynamics and characterize risk areas. This distribution can be assessed at different scales: regionally, the range of these species defines the limits of occurrence of pathogen transmission and, locally, niches determine the hazard. This latter scale requires a very precise knowledge of the areas that can be gained through the analysis of very high spatial resolution satellite images. As part of a research project on leptospirosis in La Réunion (LeptOI), this study proposes to measure the potential of a Pleiades image (product resampled to to 50 cm) to characterize the local distribution of various small terrestrial mammals in Saint-Paul swamp area. Three trapping campaigns have led to the capture of 134 animals belonging to four of the five species occurring in Reunion Island (Rattus rattus, R. norvegicus, Suncus murinus and Tenrec ecaudatus). An object-based image analysis of the Pleiades image allowed discriminating ten land use classes with a total accuracy of 83.6%. Landscape indices were calculated from the classification around each trap (shortest distances to each land use class, edge densities and proportion of each patch within buffers defined by a 50 meters or a 100 meters radius). Three areas ("wet", "urbanized" and "dry") are clearly discriminated by these variables, which can be used as ecological determinants of different epidemiological profiles. The wetland area is known as suitable to leptospirosis whereas the urbanized area is a place of co-occurrence of the four species and thus potentially favorable to the exchange of pathogens.

The Paramyxoviridae form an increasingly diverse viral family, infecting a wide variety of different hosts. In recent years, they have been linked to disease emergence in many different animal populations and in humans. Bats and rodents have been identified as major animal populations capable of harboring paramyxoviruses, and host shifting between these animals is likely to be an important driving factor in the underlying evolutionary processes that eventually lead to disease emergence. Here, we have studied paramyxovirus circulation within populations of endemic and introduced wild small mammals of the southwestern Indian Ocean region and belonging to four taxonomic orders: Rodentia, Afrosoricida, Soricomorpha, and Chiroptera. We report elevated infection levels as well as widespread paramyxovirus dispersal and frequent host exchange of a newly emerging genus of the Paramyxoviridae , currently referred to as the unclassified morbillivirus-related viruses (UMRVs). In contrast to other genera of the Paramyxoviridae , where bats have been shown to be a key host species, we show that rodents (and, in particular, Rattus rattus ) are significant spreaders of UMRVs. We predict that the ecological particularities of the southwestern Indian Ocean, where small mammal species often live in densely packed, multispecies communities, in combination with the increasing invasion of R. rattus and perturbations of endemic animal communities by active anthropological development, will have a major influence on the dynamics of UMRV infection. IMPORTANCE Identification of the infectious agents that circulate within wild animal reservoirs is essential for several reasons: (i) infectious disease outbreaks often originate from wild fauna; (ii) anthropological expansion increases the risk of contact between human and animal populations and, as a result, the risk of disease emergence; (iii) evaluation of pathogen reservoirs helps in elaborating preventive measures to limit the risk of disease emergence. Many paramyxoviruses for which bats and rodents serve as major reservoirs have demonstrated their potential to cause disease in humans and animals. In the context of the biodiversity hot spot of southwestern Indian Ocean islands and their rich endemic fauna, we show that highly diverse UMRVs exchange between various endemic animal species, and their dissemination likely is facilitated by the introduced Rattus rattus . Hence, many members of the Paramyxoviridae appear well adapted for the study of the viral phylodynamics that may be associated with disease emergence.

Leptospirosis is a bacterial zoonosis of major concern on tropical islands. Human populations on western Indian Ocean islands are strongly affected by the disease although each archipelago shows contrasting epidemiology. For instance, Mayotte, part of the Comoros Archipelago, differs from the other neighbouring islands by a high diversity of Leptospira species infecting humans that includes Leptospira mayottensis, a species thought to be unique to this island. Using bacterial culture, molecular detection and typing, the present study explored the wild and domestic local mammalian fauna for renal carriage of leptospires and addressed the genetic relationships of the infecting strains with local isolates obtained from acute human cases and with Leptospira strains hosted by mammal species endemic to nearby Madagascar. Tenrec (Tenrec ecaudatus, Family Tenrecidae), a terrestrial mammal introduced from Madagascar, is identified as a reservoir of L. mayottensis. All isolated L. mayottensis sequence types form a monophyletic clade that includes Leptospira strains infecting humans and tenrecs on Mayotte, as well as two other Malagasy endemic tenrecid species of the genus Microgale. The lower diversity of L. mayottensis in tenrecs from Mayotte, compared to that occurring in Madagascar, suggests that L. mayottensis has indeed a Malagasy origin. This study also showed that introduced rats (Rattus rattus) and dogs are probably the main reservoirs of Leptospira borgpetersenii and Leptospira kirschneri, both bacteria being prevalent in local clinical cases. Data emphasize the epidemiological link between the two neighbouring islands and the role of introduced small mammals in shaping the local epidemiology of leptospirosis., Author Summary Islands are exceptionally prone to species introduction, including pathogens with detrimental public health consequences for which the invasive alien species could act as reservoirs. Our study shows how the local non-native mammal fauna of Mayotte Island is associated with the introduction and epidemiology of human leptospirosis, a zoonosis strongly affecting tropical islands. Data presented herein identify Tenrec ecaudatus (Family Tenrecidae), an omnivorous species introduced from neighbouring Madagascar, as a reservoir of the recently named Leptospira mayottensis. Further, we suggest a Malagasy origin of L. mayottensis, which occurs naturally within the Tenrecidae radiation endemic to Madagascar. Finally, we provide evidence that dogs and rats are reservoirs for other Leptospira lineages of medical importance on Mayotte. Altogether, our study highlights the impact of species introduction on human health and further suggests that the biogeography of microorganisms in insular ecosystems, including pathogenic endemic lineages, should be considered from evolutionary and medical perspectives.