Hyperparasitism in bat flies (Diptera: Streblidae): new records and interaction networks in the Neotropics.

Hyperparasitism is defined as the interaction where one parasite is infected by another parasite. In bat flies (Streblidae and Nycteribiidae), both hyperparasites and microparasites (bacteria, viruses, fungi, and arthropods such as mites) have been documented. Fungi belonging to the order Laboulbeniales are microscopic parasites of a wide diversity of arthropod hosts. Three genera exclusively target bat flies: Arthrorhynchus, which parasitizes species within Nycteribiidae in the Eastern Hemisphere, while genus Gloeandromyces and Nycteromyces parasitize Streblidae in the Western Hemisphere. Among the hyperparasitic arthropods, mites of family Neothrombidiidae, particularly the monospecific genus Monunguis, are known to parasitize bat flies. Here we present the first records of the hyperparasites Monunguis streblida and Gloeandromyces pageanus f. polymorphus parasitizing Streblidae bat flies in Colombia and a summary of these hyperparasitic interactions in the Neotropics. We detected fungi and mites parasitizing bat flies that were collected in the Magdalena River Basin, Colombia, in field expeditions in 2018, 2022, and 2023. We identified 17 bat flies and two species of hyperparasites, specifically M. streblida and the fungi Gloeandromyces. Our search for reports of these interactions in the Neotropics revealed that seven species of Trichobius (Streblidae) are parasitized by M. streblida, whereas Paratrichobius longicrus (Streblidae) is parasitized by Gloeandromyces pageanus f. polymorphus. These interactions have been reported in 11 countries, but our records are the first of M. streblida and Laboulbeniales fungi parasitizing bat flies in Colombia. So far, a total of 14 species of fungi and one species of mite have been associated with 19 species of bat flies, which in turn, are linked to 15 species of Neotropical bats.

Molecular prevalence of Bartonella spp. in bat flies in east coast Malaysia.

Bats are a significant reservoir for numerous pathogens, including Bartonella spp. It is one of the emerging zoonotic bacterial diseases that can be transmitted to humans and may cause various unspecific clinical manifestations. Thus, bartonellosis is rarely diagnosed and is regarded as a neglected vector-borne disease (VBD). Bat flies have been hypothesised to be a vector in the transmission of pathogens among bats. They are host-specific, which reduces the likelihood of pathogen transmission across bat species; however, they are likely to maintain high pathogen loads within their host species. To explore the presence of Bartonella spp. in bat flies from Peninsular Malaysia; bat fly samples collected from various sites at the east coast states were subjected to molecular detection for Bartonella spp. It was discovered that 38.7 % of bats from Terengganu and Kelantan were infested with bat flies; however, no bat fly was found in bats collected from Pahang. The collected bat flies belonged to the families Nycteribiidae (79.6 %) and Streblidae (20.4 %). The collected bat flies were pooled according to the locations and species into 39 pools. Out of these 39 pools, 66.7 % (n = 26) were positive for Bartonella spp. by PCR. Sequence analyses of five randomly selected PCR-positive pools revealed that pools from Kelantan (n = 3) have the closest sequence identities (99 %) to Bartonella spp. strain Lisso-Nig-922 from Nigeria. However, the other pools from Terengganu (n = 2) were closely related to Bartonella spp. strain KP277 from Thailand and Bartonella spp. strain Rhin-3 from the Republic of Georgia with 99 % and 100 % sequence identity, respectively. This suggests that the Bartonella spp. found in Malaysian bat flies are genetically diverse and can potentially serve as reservoirs for pathogenic Bartonella spp.

BatFly: A database of Neotropical bat-fly interactions.

Global changes have increased the risk of emerging infectious diseases, which can be prevented or mitigated by studying host-parasite interactions, among other measures. Bats and their ectoparasitic flies of the families Streblidae and Nycteribiidae are an excellent study model but, so far, our knowledge has been restricted to fragmented records at a local scale. To help boost research, we assembled a data set of bat-fly interactions from 174 studies published between 1904 and 2022 plus three original data sets. Altogether, these studies were carried out at 650 sites in the Neotropics, mainly distributed in Mexico, Brazil, Argentina, southern USA, and Colombia, among other countries. In total, our data set contains 3984 interaction records between 237 bat species and 255 fly species. The bat species with the largest number of recorded interactions were Carollia perspicillata (357), Artibeus jamaicensis (263), and Artibeus lituratus (228). The fly species with the largest number of recorded interactions were Trichobius joblingi (256), Megistopoda aranea (235), and Megistopoda proxima (215). The interaction data were extracted, filtered, taxonomically harmonized, and made available in a tidy format together with linked data on bat population, fly population, study reference, sampling methods and geographic information from the study sites. This interconnected structure enables the expansion of information for each interaction record, encompassing where and how each interaction occurred, as well as the number of bats and flies involved. We expect BatFly to open new avenues for research focused on different levels of ecological organization and spatial scales. It will help consolidate knowledge about ecological specialization, resource distribution, pathogen transmission, and the drivers of parasite prevalence over a broad spatial range. It may also help to answer key questions such as: Are there differences in fly prevalence or mean infestation across Neotropical ecoregions? What ecological drivers explain those differences? How do specialization patterns vary among fly species in the Neotropics? Furthermore, we expect BatFly to inspire research aimed at understanding how climate and land-use changes may impact host-parasite interactions and disease outbreaks. This kind of research may help us reach Sustainable Development Goal 3, Good Health and Wellbeing, outlined by the United Nations. The data are released under a Creative Commons Attribution 4.0 International License.

Characterization of the bacterial microbiome of non-hematophagous bats and associated ectoparasites from Brazil.

Introduction: Bats, along with their ectoparasites, harbor a wide diversity of symbiotic and potential pathogenic bacteria. Despite the enormous diversity of bats (181 species), few studies aimed to investigate the bacterial microbiome of Brazilian chiropterans and associated ectoparasites. This study aimed to characterize the bacterial microbiome of non-hematophagous bats and associated Streblidae flies and Macronyssidae and Spinturnicidae mites in the state of Mato Grosso do Sul, midwestern Brazil. Methods: Oral and rectal swabs were collected from 30 bats (Artibeus lituratus [n = 13], Artibeus planirostris [n = 9], Eptesicus furinalis [n = 5], Carollia perspicillata [n = 2], and Platyrrhinus lineatus [n = 1]). In addition, a total of 58 mites (15 Macronyssidae and 43 Spinturnicidae) and 48 Streblidae bat flies were collected from the captured bats. After DNA extraction and purification, each sample's bacterial composition was analyzed with metagenomic sequencing. Results: The microbiome composition of both oral and rectal bat swab samples showed that Gammaproteobacteria was the most abundant bacterial class. Spiroplasma, Wolbachia and Bartonella represented the most abundant genera in Streblidae flies. While Wolbachia (Alphaproteobacteria) was the most abundant genus found in Spinturnicidae, Arsenophonus (Gammaproteobacteria) was found in high abundance in Macronyssidae mites. In addition to characterizing the microbiome of each sample at the class and genus taxonomic levels, we identified medically significant bacteria able to infect both animals and humans in oral (Streptococcus and Anaplasma) and rectal swabs (Enterobacter, Klebsiella, Escherichia, Enterococcus, Streptococcus), Macronyssidae (Anaplasma, Bartonella, Ehrlichia) and Spinturnicidae (Anaplasma, Bartonella) mites as well as Streblidae flies (Spiroplasma, Bartonella). Discussion and conclusion: Besides expanding the knowledge on the bacterial microbiome of non-hematophagous bats and Streblidae flies from Brazil, the present work showed, for the first time, the bacterial community of bat-associated Macronyssidae and Spinturnicidae mites.

Virome characterization and identification of a putative parvovirus and poxvirus in bat ectoparasites of Yunnan Province, China.

Ectoparasites found on bats are known to contain important microbes. However, the viruses hosted by these obligate parasites are understudied. This has led to the near oversight of the potential role of these ectoparasites in virus maintenance and transmission from bats to other interacting species and the environment. Here, we sampled bat ectoparasites parasitizing a diverse selection of bat species in the families Rhinolophidae, Vespertilionidae, Megadermatidae, Hipposideridae and Pteropodidae in Yunnan Province, China. We show that the ectoparasite prevalence was generally higher in male compared to female bats. Most ectoparasites were found to fall within the Nycteribiidae, Spinturnicidae and Streblidae bat ectoparasite families. We subsequently applied a non-biased sequencing of libraries prepared from the pooled ectoparasites, followed by an in-silico virus-centric analysis of the resultant reads. We show that ectoparasites hosted by the sampled families of bats are found to carry, in addition to a diverse set of phages, vertebrate and insect viruses in the families Aliusviridae, Ascoviridae, Chuviridae, Circoviridae, Flaviviridae, Hepadnaviridae, Hepeviridae, Herpesviridae, Iridoviridae, Marseilleviridae, Nairoviridae, Orthomyxoviridae, Parvoviridae, Poxviridae, Reoviridae, Retroviridae, and Rhabdoviridae. We further report a partial Parvovirus VP1/VP2 gene and partial Poxvirus ubiquitin-like gene predicted by two independent next generation sequencing data analysis pipelines. This study describes the natural virome of bat ectoparasites, providing a platform for understanding the role these ectoparasites play in the maintenance and spread of viruses to other animals.

Exploring the relationship between bats (Mammalia, Chiroptera) and ectoparasitic flies (Diptera, Hippoboscoidea) of the Orinoquia Region in South America.

Bat flies (Nycteribiidae and Streblidae) have been used to study co-evolutionary patterns between ectoparasites and bats. In the world, Nycteribiidae and Streblidae are represented by approximately 276 and 237 species, respectively. In regions such as the Orinoquia located in the north of South America (Colombia and Venezuela), the richness of bats is high (more than 100 documented species), but studies on Nycteribiidae and Streblidae are scarce and discontinuous. To contribute to the knowledge of ectoparasitic flies in the Orinoquia, records of flies and their interactions with bats were reviewed, including new records and associations using interaction networks. We documented 124 species of Streblidae and only 12 of Nycteribiidae for the Orinoquia in approximately 102 bat species reported in Colombia and Venezuela. New records for six species of bat flies in Colombia were found (Mastopteraguimaraesi, Noctiliostreblamaai, Paradyschiriaparvuloides, Trichobiusjubatus, Trichobiusparasiticus, and Basiliaferrisi) associated with six species of bats (Cynomopsplanirostris, Desmodusrotundus, Myotishandleyi, Molossusrufus, Noctilioalbiventris, and Phyllostomushastatus). The bat-ectoparasite interaction networks in the Orinoquia revealed a pattern of antagonistic relationships, with high specialization, modularity, and low connectivity and nesting. The identified networks are between bat fly species belonging to different ecomorphological groups with unique host species. This supports the idea of ecological niche partitioning among ectoparasitic bat flies and hosts. Our study expanded the knowledge of the distribution of some fly species and the associations with bat hosts in Colombia, by presenting morphological descriptions and new observations, which are key to understanding the ecology, diversity, and distribution of these species.

Network and parasitological analyses reveal latitudinal gradient in bats-ectoparasitic fly interactions across the Neotropic.

Ecological interactions between parasites and their hosts play a fundamental role in evolutionary processes. Selection pressures are exerted on parasites and their hosts, usually resulting in high levels of specificity. Such is the case of ectoparasitic bat-flies, but how large-scale spatial gradients affect the dynamics of their interactions with their bat hosts is still unknown. In the present study, we investigated interaction patterns between bats and their ectoparasitic flies (Streblidae and Nycteribiidae), both presenting their peak of diversity in the Neotropical region, along a latitudinal gradient. Using network analyses and parasitic indices, grounded on the latitudinal diversity gradient pattern, we evaluated how spatial gradients affect species interactions and parasitic indices at the biogeopraphic scale, with increasing species richness in interaction networks closer to the tropics, leading to increases in network modularity, size, and specialization, and to a decrease in nesting and connectivity. We conducted a literature review, focusing on studies done in the Neotropical region, and own data. We obtained a bat richness of 97 species parasitized by 128 species of ectoparasitic flies, distributed into 57 interaction networks between latitudes 29° S and 19° N in the Neotropic. Network metrics and parasitic indices varied along the latitudinal gradient, with changes in the richness of bats and their ectoparasitic flies and in the structure of their interactions; network specialization, modularity, and connectance increase with latitude, while network size decreases with latitude. Regions closer to the equator had higher parasite loads. Our results show that interaction network metrics present a latitudinal gradient and that such interactions, when observed at a local scale, hide variations that only become perceptible at larger scales. In this way, ectoparasites such as bat flies are not only influenced by the ecology and biology of their hosts, but also by other environmental factors acting directly on their distribution and survival.

Effects of forest loss and fragmentation on bat-ectoparasite interactions.

Human land use causes habitat loss and fragmentation, influencing host-parasite associations through changes in infestation rates, host mortality and possibly local extinction. Bat-ectoparasite interactions are an important host-parasite model possibly affected by such changes, as this system acts as both reservoirs and vectors of several pathogens that can infect different wild and domestic species. This study aimed to assess how the prevalence and abundance of bat ectoparasites respond to forest loss, fragmentation, and edge length. Bats and ectoparasites were sampled at twenty sites, forming a gradient of forest cover, in southwestern Brazil during two wet (2015 and 2016) and two dry (2016 and 2017) seasons. Effects of landscape metrics on host abundance as well as parasite prevalence and abundance were assessed through structural equation models. Nine host-parasite associations provided sufficient data for analyses, including one tick and eight flies on four bat species. Forest cover positively influenced the prevalence or abundance of three fly species, but negatively influenced one fly and the tick species. Prevalence or abundance responded positively to edge length for three fly species, and negatively for the tick. In turn, number of fragments influenced the prevalence or abundance of four fly species, two positively and two negatively. Our results support species-specific responses of ectoparasites to landscape features, and a tendency of host-generalist ticks to benefit from deforestation while most host-specialist flies are disadvantaged. Differences in host traits and abundance, along with parasite life cycles and environmental conditions, are possible explanations to our findings.

Phylogenetic analyses reveal bat communities in Northwestern Mexico harbor a high diversity of novel cryptic ectoparasite species.

Parasites are integral parts of ecosystem function and important drivers of evolutionary processes. Characterizing ectoparasite diversity is fundamental to studies of host-parasite interactions, evolution, and conservation, and also for understanding emerging disease threats for some vector borne pathogens. With more than 1400 species, bats represent the second most speciose mammalian clade, but their ectoparasite fauna are poorly known for most species. We sequenced mitochondrial Cytochrome Oxidase C subunit I and nuclear 18S ribosomal gene fragments, and used Bayesian phylogenetic analyses to characterize ectoparasite taxon identity and diversity for 17 species of parasitized bats sampled along the Baja California peninsula and in Northwestern Mexico. The sequence data revealed multiple novel lineages of bat bugs (Cimicidae), flies (Nycteribiidae and Streblidae), and ticks (Argasidae). Within families, the new linages showed more than 10% sequence divergence, which is consistent with separation at least at the species level. Both families of bat flies showed host specificity, particularly on Myotis species. We also identified new records for the Baja peninsula of one tick (Carios kelleyi), and of five Streblid bat fly species. One Nycteribiid bat fly haplotype from Pallid bat (Antrozous pallidus) hosts was found throughout the peninsula, suggesting potential long distance co-dispersal with hosts. Different bat bug and tick communities were found in the north and south of the peninsula. This study is the first systematic survey of bat ectoparasites in the Baja California peninsula, revealing novel lineages that are highly genetically differentiated from other parts of North America. For some ectoparasite species, haplotype distributions may reflect patterns of bat migration. This work is a first step in characterizing ectoparasite diversity over the Baja California peninsula, and understanding how ecological and evolutionary interactions shape bat ectoparasite communities among host species in different parts of their ranges.

Parasitism on bats by bat flies in remnants of a humid forest enclave area in Northeastern Brazil.

Flies are the main bloodsucking ectoparasites of Neotropical bats. The aim of this study was to describe the community of flies on bats from a humid forest enclave, surrounded by the semiarid landscape of the Caatinga domain, in Northeastern Brazil. In addition, we tested the influence of dry and rainy periods on the parasitological indices. Fieldwork was carried out in three anthropized remnants of a humid forest enclave in the Agreste mesoregion of Pernambuco state, Northeastern Brazil. Bats were captured monthly, from August 2018 to July 2019, and the ectoparasites were collected. Overall, 807 bats of 14 species and 11 genera of the families Phyllostomidae (13 spp.) and Vespertilionidae (1 sp.) were captured. Only four bat species had >50 individuals captured, and Carollia perspicillata was the most abundant species (259/807; 32%). Five hundred and seventy-one bat flies from 13 species (11 Streblidae and 2 Nycteribiidae) were collected, of which Trichobius joblingi was the most prevalent fly (197/571; 34.5%). Only the parasitism by T. joblingi was influenced by seasonality, with higher prevalence (P = 0.01) and intensity of infestation (P = 0.02) in the dry season. No lesions associated with parasitism were observed in any of the captured bats. Our results suggest that the bat-fly relationship can be affected in different ways by environmental variables, depending on the species involved. The results of this work contribute to the knowledge about ectoparasites of bats in the Neotropical region, with emphasis on humid forest enclaves, where studies of the host-parasite-environment relationship are scarce.

Species richness of bat flies and their associations with host bats in a subtropical East Asian region.

BACKGROUND: Understanding the interactions between bat flies and host bats offer us fundamental insights into the coevolutionary and ecological processes in host-parasite relationships. Here, we investigated the identities, host specificity, and patterns of host association of bat flies in a subtropical region in East Asia, which is an understudied region for bat fly research. METHODS: We used both morphological characteristics and DNA barcoding to identify the bat fly species found on 11 cavernicolous bat species from five bat families inhabiting Hong Kong. We first determined the phylogenetic relationships among bat fly species. Then, we elucidated the patterns of bat-bat fly associations and calculated the host specificity of each bat fly species. Furthermore, we assembled the mitogenomes of three bat fly species from two families (Nycteribiidae and Streblidae) to contribute to the limited bat fly genetic resources available. RESULTS: We examined 641 individuals of bat flies and found 20 species, of which many appeared to be new to science. Species of Nycteribiidae included five Nycteribia spp., three Penicillidia spp., two Phthiridium spp., one Basilia sp., and one species from a hitherto unknown genus, whereas Streblidae included Brachytarsina amboinensis, three Raymondia spp., and four additional Brachytarsina spp. Our bat-bat fly association network shows that certain closely related bat flies within Nycteribiidae and Streblidae only parasitized host bat species that are phylogenetically more closely related. For example, congenerics of Raymondia only parasitized hosts in Rhinolophus and Hipposideros, which are in two closely related families in Rhinolophoidea, but not other distantly related co-roosting species. A wide spectrum of host specificity of these bat fly species was also revealed, with some bat fly species being strictly monoxenous, e.g. nycteribiid Nycteribia sp. A, Phthiridium sp. A, and streblid Raymondia sp. A, while streblid B. amboinensis is polyxenous. CONCLUSIONS: The bat fly diversity and specificity uncovered in this study have shed light on the complex bat-bat fly ecology in the region, but more bat-parasite association studies are still needed in East Asian regions like China as a huge number of unknown species likely exists. We highly recommend the use of DNA barcoding to support morphological identification to reveal accurate host-ectoparasite relationships for future studies.

Configuration and composition of human-dominated tropical landscapes affect the prevalence and average intensity of mite and fly infestation in Phyllostomidae bats.

The conversion of natural areas into agricultural landscapes results in different mosaics of land use types, modifying biodiversity and consequently altering the patterns of ecological interactions, such as between frugivorous bats and ectoparasites. Our objectives were to investigate whether variations in the configuration and composition of human-disturbed landscapes interfere with the prevalence and average intensity of ectoparasite infestation in the frugivorous bats Artibeus lituratus (Olfers, 1818), Carollia perspicillata (Linnaeus, 1758), and Sturnira lilium (É Geoffroy, 1810), in a region of the Brazilian Atlantic Forest. We also evaluated whether there is a response in the parasite load associated with the ectoparasite group (mite or fly). We found six species of flies and three mites. The proportion of infested hosts was more affected by the landscape than the mean infestation values. Land cover diversity influenced seven of the interactions studied. Forest cover affected eight of the interactions and was associated with a reduction in the parasite load in seven of them. The increase in the proportion of edges per area of each fragment presented a different influence related to the host species. Variations in parasite load did not show any typical response related to the mite or fly group. Our study indicates that landscape configuration and composition interfere with bat-ectoparasite interactions, which may be related to interference in encounter rates between hosts (for mites and flies) and between hosts and their ectoparasites in roosts (for flies). The taxonomic identity of the interacting species suggests that the relationship with the landscape is context-dependent.

Ectoparasitic flies of bats (Mammalia: Chiroptera) in urban green areas of northeastern Brazil.

In urban and degraded areas, ectoparasite abundance can be affected by increasing human population density and habitat fragmentation. This study aimed to characterize the ectoparasitic fly community associated with bats in the urban green areas of Sergipe, Brazil. Campaigns were conducted monthly, for two consecutive nights, between September 2019 and February 2021. To capture the bats, ten mist nets were set up inside and at the edge of the habitat fragments. All ectoparasites found were removed from the bats and stored in 70% alcohol. The specificity index, parasitological rates, and level of parasite aggregation were calculated, and the influence of host sex and seasonality on parasitological rates were verified for the most parasitized bats. The collected ectoparasites corresponded to the families Nycteribiidae (S = 1; n = 26) and Streblidae (S = 13; n = 849), with Trichobius costalimai and Medistopoda aranea being the most abundant species. For some interactions, there was an influence of host sex on the prevalence rates, with the highest number of parasites being found on females, which can be explained by their greater susceptibility to parasitism owing to their long stay in roosts. The seasonality influenced the parasitological rates, and opposing patterns (from what was expected) were observed for some interactions; this influence may be due to the biological differences between parasite species. This study provides relevant data on this interaction, especially for urban areas in northeastern Brazil, expanding the number of studies in the State of Sergipe and promoting future studies.

Review of studies about bat-fly interactions inside roosts, with observations on partnership patterns for publications.

Pressures from anthropogenic disturbances have triggered a wealth of studies focusing on the assessment and mitigation of the negative impacts of these disturbances on inter and intraspecific ecological interactions, including bats and bat flies in their roosts. The heterogeneity of research methods employed for these studies and the scientific imbalance between countries may constrain advances and the consolidation of the knowledge on this subject. We reviewed the literature regarding bat and bat-ectoparasite interactions in roosts assessing global research trends and patterns of author collaborative work to be able to identify key questions for future studies and potential initiatives to improve the knowledge on this subject. Current information available has mostly come from the Americas and is predominantly focused on the recognition and description of parasite-host interactions between bats and bat flies. Our findings suggest the value of increasing collaboration for future research, as several countries with largely diverse environments and high organismal richness are disconnected from the countries that produce the most publications in this area, and/or have low records of publications. These regions are in the Global South, mostly in South American and African countries. We suggest that more collaborative networks may increase scientific production in the area, and that investing in local research development and enhancing partnerships for publications may strengthen the field. These research programs and collaborations are key for the development of conservation strategies for bats and bat flies, for their roosts, and for understanding bat and bat-ectoparasite interactions.

Modularity and specialization in bat-fly interaction networks are remarkably consistent across patches within urbanized landscapes and spatial scales.

Patterns of specialization and the structure of interactions between bats and ectoparasitic flies have been studied mostly on non-urban environments and at local scales. Thus, how anthropogenic disturbances influence species interactions and network structure in this system remain poorly understood. Here, we investigated patterns of interaction between Phyllostomidae bats and ectoparasitic Streblidae flies, and variations in network specialization and structure across Cerrado patches within urbanized landscapes in Brazil and between local and regional scales. We found high similarity in the richness and composition of bat and fly species across communities, associated with low turnover of interactions between networks. The high specialization of bat-streblid interactions resulted in little connected and modular networks, with the emergence of modules containing subsets of species that interact exclusively or primarily with each other. Such similarities in species and interaction composition and network structure across communities and scales suggest that bat-fly interactions within Cerrado patches are little affected by the degree of human modification in the surrounding matrix. This remarkable consistency is likely promoted by specific behaviors, the tolerance of Phyllostomidae bats to surrounding urbanized landscapes as well as by the specificity of the streblid-bat interactions shaped over evolutionary time.

Bat-ectoparasitic fly relationships in a seasonally dry tropical forest in Brazil.

Bat ectoparasitic flies are hematophagous insects highly specialized to parasitize only bats. Knowledge about how biotic and abiotic factors can influence ecological relationships between parasites and hosts is in general incipient. Large information gaps are even worst in biodiversity-rich, but poorly sampled areas like Brazil's Caatinga, the largest tropical dry forest in South America. We used bats and their highly specialized ectoparasitic flies as a study model to clarify some aspects of this host-parasite system in this semiarid environment. We conducted fieldwork at 55 sites in the Caatinga, between April 2017 and March 2020 and collected 1300 flies (19 species) on 333 bats (15 species). Bat ectoparasitic flies were highly host-specific, had an aggregated distribution, frequently formed infracommunities with only one species, and had a male-biased sex ratio. Except for the prevalence of the streblid fly Strebla guajiro on Carollia perspicillata, which was significantly higher in the rainy season, bat flies showed no preference - expressed by frequency and intensity of infestation - for host sex, and their interspecific relationships were not mediated by rainfall. Other variables that could potentially be interfering in this host-parasite relationship deserve further attention, especially in environments such as the Caatinga where there is great seasonal variation. Furthermore, the existence of species-specific responses must be taken into account.

Bat Flies of the Family Streblidae (Diptera: Hippoboscoidea) Host Relatives of Medically and Agriculturally Important "Bat-Associated" Viruses.

Bat flies (Hippoboscoidea: Nycteribiidae and Streblidae) are obligate hematophagous ectoparasites of bats. We collected streblid bat flies from the New World (México) and the Old World (Uganda), and used metagenomics to identify their viruses. In México, we found méjal virus (Rhabdoviridae; Vesiculovirus), Amate virus (Reoviridae: Orbivirus), and two unclassified viruses of invertebrates. Méjal virus is related to emerging zoonotic encephalitis viruses and to the agriculturally important vesicular stomatitis viruses (VSV). Amate virus and its sister taxon from a bat are most closely related to mosquito- and tick-borne orbiviruses, suggesting a previously unrecognized orbivirus transmission cycle involving bats and bat flies. In Uganda, we found mamucuso virus (Peribunyaviridae: Orthobunyavirus) and two unclassified viruses (a rhabdovirus and an invertebrate virus). Mamucuso virus is related to encephalitic viruses of mammals and to viruses from nycteribiid bat flies and louse flies, suggesting a previously unrecognized orthobunyavirus transmission cycle involving hippoboscoid insects. Bat fly virus transmission may be neither strictly vector-borne nor strictly vertical, with opportunistic feeding by bat flies occasionally leading to zoonotic transmission. Many "bat-associated" viruses, which are ecologically and epidemiologically associated with bats but rarely or never found in bats themselves, may actually be viruses of bat flies or other bat ectoparasites.

Visual system characterization of the obligate bat ectoparasite Trichobius frequens (Diptera: Streblidae).

As an obligate ectoparasite of bats, the bat fly Trichobius frequens (Diptera: Streblidae) inhabits the same subterranean environment as their nocturnal bat hosts. In this study, we characterize the macromorphology, optical architecture, rhabdom anatomy, photoreceptor absorbance, and opsin expression of the significantly reduced visual system in T. frequens resulting from evolution in the dark. The eyes develop over a 21-22 day pupal developmental period, with pigmentation appearing on pupal day 11. After eclosion as an adult, T. frequens eyes consist of on average 8 facets, each overlying a fused rhabdom consisting of anywhere from 11 to 18 estimated retinula cells. The dimensions of the facets and fused rhabdoms are similar to those measured in other nocturnal insects. T. frequens eyes are functional as shown by expression of a Rh1 opsin forming a visual pigment with a peak sensitivity to 487 nm, similar to other dipteran Rh1 opsins. Future studies will evaluate how individuals with such reduced capabilities for spatial vision as well as sensitivity still capture enough visual information to use flight to maneuver through dark habitats.

Moscas de murciélagos en algunas localidades de la costa peruana / Bat flies in some localities of the Peruvian coast

Resumen En el Perú se registran al menos 180 especies de murciélagos. Sin embargo, son pocos los estudios acerca de sus ectoparásitos, dentro de los cuales destacan las denominadas moscas de murciélagos (Diptera: Streblidae y Nycteribiidae), en los cuales se estima un total de 158 especies en la región Neotropical. En el Perú, una de las regiones con menos información es la costa. En este estudio actualizamos el conocimiento de las asociaciones ectoparásito-hospedero en murciélagos de la costa peruana, una de las regiones ecologicas con más degradación y pérdida de hábitat. Se evaluaron cinco localidades en las regiones de Piura, Lima y Tacna. Se capturaron 85 murciélagos pertenecientes a 7 especies; 4 de Phyllostomidae, 2 de Molossidae y 1 de Vespertilionidae. Se encontraron seis especies de dípteros (Diptera: Streblidae) y un hemíptero (Hemiptera: Polyctenidae) y se reporta por primera vez algunas de estas asociaciones en las regiones de Lima y Piura. Se reporta también a Anoura peruana como hospedero de Anastrebla modestini (Streblidae) en una colonia reproductiva compartida con Platalina genovensium, y por primera vez para la región de Piura, el co-parasitismo de los estréblidos Megistopoda aranea, Aspidoptera phyllostomatis y Metelasmus pseudopterus en un mismo individuo de Artibeus fraterculus.

Abstract There are at least 180 species of bats in Peru. However, there are few studies about ectoparasites of bats, among which the so-called bat flies (Diptera: Streblidae and Nycteribiidae) stand out, with a total of 158 species estimated for the Neotropical region. In Peru, one of the ecological regions with the least information is the coast. In this study we updated the knowledge of the ectoparasite-host associations in bats from the Peruvian coast, one of the regions of the country with the most degradation and loss of habitat. Five locations in the Piura, Lima and Tacna regions were evaluated. 85 bats belonging to 7 species were captured: 4 from Phyllostomidae, 2 from Molossidae and 1 from Vespertilionidae. Six dipterous species (Diptera: Streblidae) and one Hemiptera species (Hemiptera: Polyctenidae) were found and some of these associations are reported for the first time in the Lima and Piura regions. Anoura peruana is also reported as the host of Anastrebla modestini (Streblidae) in a reproductive colony shared with Platalina genovensium, and for the first time for the Piura region, the co-parasitism of the Megistopoda aranea, Aspidoptera phyllostomatis and Metelasmus pseudopterus in a same individual of Artibeus fraterculus.

Host ecology moderates the specialization of Neotropical bat-fly interaction networks.

The transmission of diseases through parasites is a key mechanism in the regulation of plant and animal populations in ecosystems. Therefore, it is necessary to investigate the relative effect of the variables that can shape the specificity of host-parasite interactions. Previous studies have found that specialization of antagonistic interactions between fly ectoparasites and bats changes according to forest type, host richness, and roosting ecology of bats. In this study, we tested these hypotheses using data from 48 bat communities. In general, our results support previous findings that bat-fly interactions are specialized, resulting in lower niche overlap among bat flies species. In addition, we found that the specificity of bat-fly interactions is lower in tropical mountain forests and is positively related with the richness of bat host species of each study site. Finally, there was a higher bat flies niche overlap in smaller bat-fly interaction networks recorded in bat roosts in caves. We conclude that the roosting ecology of bats could be a key factor to understand the mechanisms related to the horizontal transmission of ectoparasitic flies among bats.