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Bats are the second most diverse order of mammals and play a central role in ecosystem dynamics. They are also important reservoirs of potentially zoonotic microorganisms, of which rabies virus is the most lethal among the bat-transmitted zoonotic pathogens. Importantly, recent outbreaks of human rabies have been reported from the Brazilian Amazon. Here we present a survey of bat species and rabies virus (RABV) circulation in a bat assemblage in the Marajó region, northern Brazil. Using data from mist-net captures and bioacoustic sampling, 56 bat species were recorded along the Jacundá River basin over a 10-day expedition in November 2022. For the investigation of RABV, we used the direct fluorescent antibody test (DFAT) and the rapid fluorescent focus inhibition test (RFFIT). In total, 159 bat individuals from 22 species were investigated for RABV. Five adults of the common vampire bat, Desmodus rotundus, showed RABV-specific antibodies in serum samples. Additionally, we report on local residents with injuries caused by D. rotundus bites and the occurrence of colonies of non-hematophagous bats from different species roosting inside human residences. This scenario raises concerns about the risks of new cases of human rabies and other zoonotic diseases associated with bats in the region and highlights the need for epidemiological surveillance and mitigation measures to prevent outbreaks of emerging infectious diseases.

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The causative agent of Chagas disease is Trypanosoma cruzi, which is widely distributed throughout the South American continent and extends into North America. Its occurrence in bats is poorly described and may impact the disease's maintenance and epidemiology. The aim of this study was to detect the agent by PCR assays targeting kDNA and nuclear DNA in the organs of 203 urban bats and rural vampire bats from the Brazilian Atlantic Forest, São Paulo state, during the pandemic period from 2020 to 2022. In total, 6 of the 203 bats (2.97%) were positive for T. cruzi. Infection was detected in 2% (2/101) of Desmodus rotundus, 33% (1/3) of Nyctinomops laticaudatus, 25% (1/4) of Artibeus lituratus, 4% (1/24) of Eumops glaucinus and in 2% (1/41) of Molossus molossus. The gene sequences obtained were assessed for quality and deposited in a public repository. Fruit bats were statistically associated with positivity for T. cruzi. To our knowledge, this study detected T. cruzi for the first time in bats from São Paulo state and in N. laticaudatus and E. glaucinus species.

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Social structure can emerge from hierarchically embedded scales of movement, where movement at one scale is constrained within a larger scale (e.g. among branches, trees, forests). In most studies of animal social networks, some scales of movement are not observed, and the relative importance of the observed scales of movement is unclear. Here, we asked: how does individual variation in movement, at multiple nested spatial scales, influence each individual's social connectedness? Using existing data from common vampire bats (Desmodus rotundus), we created an agent-based model of how three nested scales of movement-among roosts, clusters and grooming partners-each influence a bat's grooming network centrality. In each of 10 simulations, virtual bats lacking social and spatial preferences moved at each scale at empirically derived rates that were either fixed or individually variable and either independent or correlated across scales. We found that numbers of partners groomed per bat were driven more by within-roost movements than by roost switching, highlighting that co-roosting networks do not fully capture bat social structure. Simulations revealed how individual variation in movement at nested spatial scales can cause false discovery and misidentification of preferred social relationships. Our model provides several insights into how nonsocial factors shape social networks.

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In addition to zoonotic viral pathogens, bats can also harbor bacterial pathogens, including hemoplasmas (hemotropic mycoplasmas) and Coxiella burnetii. The present study aimed to investigate, using molecular techniques, the presence of hemoplasmas and C. burnetii in spleen samples from vampire bats in northern Brazil. For this purpose, between 2017 and 2019, spleen samples were collected from Desmodus rotundus (n = 228) and Diaemus youngii (n = 1) captured in the states of Pará (n = 207), Amazonas (n = 1), Roraima (n = 18) and Amapá (n = 3). DNA samples extracted from the bat spleen and positive in PCR for the endogenous gapdh gene were subjected to conventional PCR assays for the 16S rRNA, 23S rRNA and RNAse P genes from hemoplasmas and to qPCR based on the IS1111 gene element for C. burnetii. All spleen samples from vampire bats were negative in the qPCR for C. burnetii. Hemoplasmas were detected in 10 % (23/229) of spleen samples using a PCR based on the 16S rRNA gene. Of these, 21.73 % (5/23) were positive for the 23S rRNA gene and none for the RNAseP gene. The seven hemoplasma 16S rRNA sequences obtained were closely related to sequences previously identified in vampire bats from Belize, Peru and Brazil. The 23S rRNA sequence obtained revealed genetic proximity to hemoplasmas from non-hematophagous bats from Brazil and Belize. The analysis revealed different circulating genotypes among Brazilian vampire bats, in addition to a trend towards genera-specific hemoplasma genotypes. The present study contributes to the knowledge of the wide diversity of hemoplasmas in vampire bats.

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Bat-mediated human rabies is a viral zoonotic disease that poses a serious threat to the public health of traditional peoples, especially indigenous populations that maintain primitive cultural and social habits, such as the Maxakali ethnic group, located in the southeastern region of Brazil. The sociocultural habit of this population led to the emergence between April and May 2022 of the viral spillover of rabies transmitted by bats, which decimated the lives of four children from this population who maintained contact with this animal as a recreational practice. Because the vampire bats Desmodus rotundus have exceptional ecology and social characteristics that can have important effects on the dynamics of viral dispersion in this indigenous population, I present the dynamics of contact between native children and the bat and the meaning of this relationship, which involves ritualistic and recreational significance. As important as knowing the reasons for this practice is discussing some intrinsic and extrinsic factors that imply risks that intensify the vulnerability of this population to the transmission of the rabies virus at any time. In view of this, I warn of the need to adopt efficient strategies to mitigate the risks of a new emergency in this region. Although emergency containment measures were carried out during the critical period of the outbreak, such animal and environmental control actions must become routine programmatic and structuring interventions. Essential for rabies surveillance in this population is to develop culturally adapted interethnic health education campaigns to guarantee the accessibility of the Maxakali indigenous people to the content taught, so that any attempt at domestication, captivity and recreational practices with bats of any species is discouraged, thus avoiding a possible re-emergence of this anthropozoonosis that has impacted not only the epidemiological scenario in this region, but throughout Brazil, and also throughout Latin America.

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Desmodus rotundus, vampire bats, transmit dangerous infections, and brucellosis is a hazardous zoonotic disease, two adversities that coexist in the subtropical and tropical areas of the American continent. Here, we report a 47.89% Brucella infection prevalence in a colony of vampire bats inhabiting the tropical rainforest of Costa Rica. The bacterium induced placentitis and fetal death in bats. Wide-range phenotypic and genotypic characterization placed the Brucella organisms as a new pathogenic species named Brucella nosferati sp. nov., isolated from bat tissues, including the salivary glands, suggesting feeding behavior might favor transmission to their prey. Overall analyses placed B. nosferati as the etiological agent of a reported canine brucellosis case, demonstrating its potential for infecting other hosts. To assess the putative prey hosts, we analyzed the intestinal contents of 14 infected and 23 non-infected bats by proteomics. A total of 54,508 peptides sorted into 7,203 unique peptides corresponding to 1,521 proteins were identified. Twenty-three wildlife and domestic taxa, including humans, were foraged by B. nosferati-infected D. rotundus, suggesting contact of this bacterium with a broad range of hosts. Our approach is appropriate for detecting, in a single study, the prey preferences of vampire bats in a diverse area, demonstrating its suitability for control strategies where vampire bats thrive. IMPORTANCE The discovery that a high proportion of vampire bats in a tropical area is infected with pathogenic Brucella nosferati and that bats forage on humans and many wild and domestic animals is relevant from the perspective of emerging disease prevention. Indeed, bats harboring B. nosferati in their salivary glands may transmit this pathogenic bacterium to other hosts. This potential is not trivial since, besides the demonstrated pathogenicity, this bacterium possesses all the required virulent arsenal of dangerous Brucella organisms, including those that are zoonotic for humans. Our work has settled the basis for future surveillance actions in brucellosis control programs where these infected bats thrive. Moreover, our strategy to identify the foraging range of bats may be adapted for exploring the feeding habits of diverse animals, including arthropod vectors of infectious diseases, and therefore of interest to a broader audience besides experts on Brucella and bats.

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Spatial assortment can be both a cause and a consequence of cooperation. Proximity promotes cooperation when individuals preferentially help nearby partners, and conversely, cooperation drives proximity when individuals move towards more cooperative partners. However, these two causal directions are difficult to distinguish with observational data. Here, we experimentally test if forcing randomly selected pairs of equally familiar female common vampire bats (Desmodus rotundus) into close spatial proximity promotes the formation of enduring cooperative relationships. Over 114 days, we sampled 682 h of interactions among 21 females captured from three distant sites to track daily allogrooming rates over time. We compared these rates before, during and after a one-week period, during which we caged random triads of previously unfamiliar and unrelated vampire bats in proximity. After the week of proximity when all bats could again freely associate, the allogrooming rates of pairs forced into proximity increased more than those of the 126 control pairs. This work is the first to experimentally demonstrate the causal effect of repeated interactions on cooperative investments in vampire bats. Future work should determine the relative importance of mere association versus interactions (e.g. reciprocal allogrooming) in shaping social preferences.

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The control of vampire bat rabies (VBR) in Brazil is based on the culling of Desmodus rotundus and the surveillance of outbreaks caused by D. rotundus in cattle and humans in addition to vaccination of susceptible livestock. The detection of anti-rabies antibodies in vampire bats indicates exposure to the rabies virus, and several studies have reported an increase of these antibodies following experimental infection. However, the dynamics of anti-rabies antibodies in natural populations of D. rotundus remains poorly understood. In this study, we took advantage of recent outbreaks of VBR among livestock in the Sao Paulo region of Brazil to test whether seroprevalence in D. rotundus reflects the incidence of rabies in nearby livestock populations. Sixty-four D. rotundus were captured during and after outbreaks from roost located in municipalities belonging to three regions with different incidences of rabies in herbivores. Sixteen seropositive bats were then kept in captivity for up to 120 days, and their antibodies and virus levels were quantified at different time points using the rapid fluorescent focus inhibition test (RFFIT). Antibody titers were associated with the occurrence of ongoing outbreak, with a higher proportion of bats showing titer >0.5 IU/ml in the region with a recent outbreak. However, low titers were still detected in bats from regions reporting the last outbreak of rabies at least 3 years prior to sampling. This study suggests that serological surveillance of rabies in vampire bats can be used as a tool to evaluate risk of outbreaks in at risk populations of cattle and human.

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Outbreaks of human rabies transmitted by hematophagous bats occurred in 2018 in Pará state, Brazil, eastern Amazon, after 12 years of no record of the disease. Thus, it is necessary to understand the epidemiological characteristics of these attacks to protect the local population. This study aimed to characterize the bat bite populations in the municipality of São João da Ponta, Pará State, Brazil, in 2013-2015. The Notifiable Diseases Information System (SINAN) database was used to identify the five individuals who sought medical care during the study period (seeds). Other individuals who were attacked during the same period but did not seek medical care (n = 61) were reached by snowball sampling, and a descriptive analysis was performed based on information obtained from questionnaires. Majority of the interviewees were men (92.4%; 61/66) and adults aged 20-50 years (69.9%; 46/66) and had <4 years of formal school education (86.3%; 57/66). Additionally, most of them were rural residents (92.4%; 61/66) and crab fishermen (79.3%; 53/66). The interviewees (92.4%; 61/66) identified mangroves of the Mãe Grande de Curuçá extractive reserve, where groups of fishermen sometimes gather for several days for crab fishing, often living in improvised dwellings without walls and covered by tarps or straw (88.8%; 56/66), conducive to attacks by vampire bats. Overall, 42.4% (28/66) of the participants had been bitten more than four times throughout their life. The median number of attacks over the participant's lifetime was 3.11 (range, 1-23). Participants were unaware of the risk of contracting rabies from the bite (95.4%; 65/66). These results suggest that vampire bat attacks are essentially an occupational hazard in the study region. Moreover, for each reported attack, there were at least 12.2 unreported cases. Thus, the study highlights the need to develop strategies for prophylactic treatment of this population.

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Individual animals across many different species occasionally 'adopt' unrelated, orphaned offspring. Although adoption may be best explained as a by-product of adaptive traits that enhance parental care or promote the development of parental skills, one factor that is possibly important for the likelihood of adoption is the history of cooperative interactions between the mother, adopted offspring and adopter. Using 652 h of behavioural samples collected over four months, we describe patterns of allogrooming and food sharing before and after an instance of non-kin adoption between two adult female common vampire bats (Desmodus rotundus) that were captured from distant sites (340 km apart) and introduced to one another in captivity. The first female died from an illness 19 days after giving birth. The second female groomed and regurgitated food to the mother more often than any other group member, then groomed, nursed and regurgitated food to the orphaned, female pup. The substantial increase in alloparental care by this female after the mother's death was not observed among the 20 other adult females that were present in the colony. Our findings corroborate previous reports of non-kin adoption in common vampire bats and are consistent with the hypothesis that non-kin adoption can be motivated, in part, by a history of cooperative interactions.

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Some nonhuman animals form adaptive long-term cooperative relationships with nonkin that seem analogous in form and function to human friendship [1-4]. However, it remains unclear how these bonds initially form, especially when they entail investments of time and energy. Theory suggests individuals can reduce the risk of exploitation by initially spreading out smaller cooperative investments across time [e.g., 5] or partners [6], then gradually escalating investments in more cooperative partnerships [7]. Despite its intuitive appeal, this raising-the-stakes model [7] has gained surprisingly scarce empirical support. Although human strangers do "raise the stakes" when making bids in cooperation games [8], there has been no clear evidence for raising the stakes during formation of social bonds in nature. Existing studies are limited to cooperative interactions with severe power asymmetries (e.g., the cleaner-client fish mutualism [9]) or snapshots of a single behavior within established relationships (grooming in primates [10-13]). Raising the stakes during relationship formation might involve escalating to more costly behaviors. For example, individuals could "test the waters" by first clustering for warmth (no cost), then conditionally grooming (low cost), and eventually providing coalitionary support (high cost). Detecting such a pattern requires introducing random strangers and measuring the emergence of natural helping behaviors that vary in costs. We performed this test by tracking the emergence of social grooming and regurgitated food donations among previously unfamiliar captive vampire bats (Desmodus rotundus) over 15 months. We found compelling evidence that vampire bats selectively escalate low-cost grooming before developing higher-cost food-sharing relationships.

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The life history strategy of common vampire bats (Desmodus rotundus) suggests that learning might play a role in development of their foraging skills. We took advantage of 12 captive births in a study colony of vampire bats to test the role of past experience in two aspects of feeding. First, we compared preferences for blood temperature in 32 wild-born vampire bats versus 11 captive-born vampire bats that had only previously fed on blood of ambient temperature or colder. We found no evidence for a preference in either group for blood presented at 4 °C versus 37 °C. Second, we tested whether captive-born vampire bats with no previous experience of feeding on live animals could successfully feed on a live chicken. Five of 12 naïve captive-born bats were able to bite the chicken and draw blood, but only one bat gained more than 5% of body mass. We were unable to reasonably compare their feeding performance with that of wild-born bats because only two of three wild-born, short-term captive bats fed on the chicken and none of the seven wild-born, long-term captive mothers attempted to feed. This unexpected lack of feeding might be due to a previously reported age-dependent neophobia. When six of the captive-born bats were released in the wild, they appeared to feed successfully because they survived for more than three consecutive nights. We suggest further tests that would better clarify the role of learning in the development of foraging in vampire bats.

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Molecular phylogenetic studies have revealed the growing diversity of bat trypanosomes. Here, 14 isolates from blood samples of the vampire bat Desmodus rotundus (Phyllostomidae) from Rio de Janeiro, Southeast Brazil, were cultivated, and morphologically and molecularly characterized. All isolates represent a novel species named Trypanosoma madeirae n. sp. positioned in the Neobat lineage of the clade T. cruzi. The Neobat lineage also comprises closely related trypanosomes of clades Neotropic 1, 2 and 3 from diverse phyllostomid species. Trypanosomes of Neotropic 1, found in Trachops cirrhosus and Artibeus jamaicensis (phyllostomids), likely represent a different species or genotype closely related to T. madeirae. Consistent with its phylogenetic positioning, T. madeirae differs from Trypanosoma cruzi in morphology of both epimastigote and trypomastigote culture forms and does not infect Triatoma infestans. Similar to its closest relatives of Neobat lineage, T. madeirae was unable to develop within mammalian cells. To date, PCR-surveys on archived blood/liver samples unveiled T. madeirae exclusively in D. rotundus from Southern to Northern Brazil. The description of a new species of bat trypanosome associated with vampire bats increases the repertoire of trypanosomes infecting D. rotundus, currently comprised of Trypanosoma cruzi, T. cruzi marinkellei, Trypanosoma dionisii, Trypanosoma rangeli, Trypanosoma pessoai, and Trypanosoma madeirae.

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Bartonella spp. have been identified in many bat species worldwide, including the zoonotic species, Candidatus Bartonella mayotimonensis. The common vampire bat (Desmodus rotundus) preys preferentially on livestock in Latin America and is frequently infected with Bartonella spp. To determine the potential role of D. rotundus in transmitting Bartonella to livestock, common vampire bats and bat-bitten domestic ruminants from Mexico were tested for Bartonella infection by blood culture or conventional PCR. Furthermore, to explore the possibility of bite transmission during blood feeding, saliva swabs from 35 D. rotundus known to be either Bartonella bacteremic (N = 17) or blood culture negative (N = 18) were tested by PCR to detect the presence of Bartonella DNA. Twenty (17.1%) of 117 sheep and 16 (34.8%) of 46 cattle were Bartonella bacteremic by PCR testing. However, none of them were infected with Bartonella strains previously isolated from vampire bats and none of the 35 D. rotundus saliva swabs tested were PCR positive for Bartonella. All but two animals among those which were Bartonella culture and/or PCR positive, were infected with either B. bovis (cattle) or B. melophagi (sheep). Two sheep were infected by a possible new species, Candidatus Bartonella ovis, being phylogenetically closer to B. bovis than B. melophagi. This study does not support the role of D. rotundus as a reservoir of Bartonella species infecting livestock, which could be transmitted via bite and blood feeding and therefore suggest limited risk of zoonotic transmission of Bartonella from common vampire bats to humans.

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Designing experiments on social learning using an untested behaviour or species requires baseline knowledge of how the animals will perform. We conducted a pilot study of a procedure for rapidly testing social learning in the highly social common vampire bat (Desmodus rotundus) using a simple maze. To create demonstrators, we allowed captive bats to learn to exit a three-dimensional maze, which reunited them with their colony as a reward. We then filmed naive bats in the same maze, comparing their ability to exit the maze before, during and after the addition of a trained demonstrator. The presence of a demonstrator increased the exit rates of naive bats, presumably by attracting the attention of the naive bats to the maze exit. Four of the five naive bats that exited in the presence of a demonstrator retained the ability to exit without the demonstrator. No naive bat exited during trials without a potential demonstrator present. This experimental procedure appears to be a promising approach for efficient tests of social learning in vampire bats because maze difficulty can be manipulated to adjust learning rates and each trial requires only 15 min.

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Vampire bats are the only mammals known to feed exclusively on blood from other animals, often from domestic cattle. We tested the hypothesis that the adaptation of vampire bats to hematophagy would have resulted in shared viral communities among vampire bats and cattle, as a direct result of historic spillover events occurring due to hematophagy. We analyzed the presence of different viruses in sample populations of sympatric bat and prey populations and searched for shared viruses between taxa. A limited number of DNA viral groups were detected within each species. However, there was no evidence for a shared viral community among the vampire bat and cattle populations tested.

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Common vampire bats (Desmodus rotundus) are a key rabies vector in South America. Improved management of this species requires long-term, region-specific information. To investigate patterns of demography and dispersal, we analysed 13 642 captures of common vampire bats in Northern Argentina from the period 1969-2004. In contrast with findings from more tropical regions, we found reproductive seasonality with peak pregnancy in September and peak lactation in February. Curiously, sex ratios were consistently male-biased both in maternity roosts and at foraging sites. Males comprised 57% of 9509 adults caught at night, 57% of 1078 juveniles caught at night, 57% of 603 juveniles caught in roosts during the day, and 55% of 103 newborns and mature fetuses. Most observed roosts were in man-made structures. Movements of 1.5-54 km were most frequent in adult males, followed by young males, adult females and young females. At night, males visited maternity roosts, and non-pregnant, non-lactating females visited bachelor roosts. Males fed earlier in the night. Finally, we report new longevity records for free-ranging vampire bats: 16 and 17 years of age for a female and male, respectively. Our results are consistent with model predictions that sex-biased movements might play a key role in rabies transmission between vampire bat populations.

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Many bats are extremely social. In some cases, individuals remain together for years or even decades and engage in mutually beneficial behaviours among non-related individuals. Here, we summarize ways in which unrelated bats cooperate while roosting, foraging, feeding or caring for offspring. For each situation, we ask if cooperation involves an investment, and if so, what mechanisms might ensure a return. While some cooperative outcomes are likely a by-product of selfish behaviour as they are in many other vertebrates, we explain how cooperative investments can occur in several situations and are particularly evident in food sharing among common vampire bats (Desmodus rotundus) and alloparental care by greater spear-nosed bats (Phyllostomus hastatus). Fieldwork and experiments on vampire bats indicate that sharing blood with non-kin expands the number of possible donors beyond kin and promotes reciprocal help by strengthening long-term social bonds. Similarly, more than 25 years of recapture data and field observations of greater spear-nosed bats reveal multiple cooperative investments occurring within stable groups of non-kin. These studies illustrate how bats can serve as models for understanding how cooperation is regulated in social vertebrates.

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Vampire bat rabies is a public and animal health concern throughout Latin America. As part of an ecological study of vampire bat depredation on cattle in southern Guatemala, we conducted a vaccine seroconversion study among three dairy farms. The main objectives of this cross sectional and cohort study were to understand factors associated with bat bites among cattle, to determine whether unvaccinated cattle had evidence of rabies virus exposure and evaluate whether exposure was related to bat bite prevalence, and to assess whether cattle demonstrate adequate seroconversion to two commercial vaccines used in Guatemala. In 2012, baseline blood samples were collected immediately prior to intramuscular inoculation of cattle with one of two modified live rabies vaccines. Post vaccination blood samples were collected 13 and 393 days later. Sera were tested for rabies virus neutralizing antibodies (rVNA) by the rapid fluorescent focus inhibition test (RFFIT). Across two years of study, 36% (254/702) of inspected cattle presented gross evidence of vampire bat bites. Individual cattle with a bat bite in 2012 were more likely have a bat bite in 2013. Prior to vaccination, 12% (42/350) of cattle sera demonstrated rVNA, but bite status in 2012 was not associated with presence of rVNA. Vaccine brand was the only factor associated with adequate rVNA response of cattle by day 13. However, vaccine brand and rVNA status at day 13 were associated with an adequate rVNA titer on day 393, with animals demonstrating an adequate titer at day 13 more likely to have an adequate titer at day 393. Our findings support stable levels of vampire bat depredation and evidence of rVNA in unvaccinated cattle. Brand of vaccine may be an important consideration impacting adequate rVNA response and long-term maintenance of rVNA in cattle. Further, the results demonstrate that initial response to vaccination is associated with rVNA status over one year following vaccination.

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A limited number of studies have focused on the population genetic structure of vampire bats (Desmodus rotundus) in America. This medium-sized bat is distributed in tropical areas of the continent with high prevalence in forested livestock areas. The aim of this work was to characterize the vampire population structure and their genetic differentiation. For this, we followed standard methods by which live vampires (caught by mist-netting) and preserved material from scientific collections, were obtained for a total of 15 different locations, ranging from Chihuahua (North) to Quintana Roo (Southeast). Tissue samples were obtained from both live and collected animals, and the genetic differentiation, within and among localities, was assessed by the use of seven microsatellite loci. Our results showed that all loci were polymorphic and no private alleles were detected. High levels of heterozygosis were detected when the proportion of alleles in each locus were compared. Pairwise F ST and R ST detected significant genetic differentiation among individuals from different localities. Our population structure results indicate the presence of eleven clusters, with a high percentage of assigned individuals to some specific collecting site. Rev. Biol. Trop. 62 (2): 659-669. Epub 2014 June 01.

Muy pocos trabajos se han enfocado en el estudio genético de las poblaciones de vampiro (Desmodus rotundus) en América. Este murciélago de tamaño mediano se encuentra distribuido en las áreas tropicales de América, con una gran prevalencia en zonas de ganadería. La recolecta de tejidos se realizó mediante redes de niebla y en con ejemplares de colecciones, estas dan un total de 15 localidades. Mediante el uso de siete microsatellites, nosotros estudiamos la diferenciación genética dentro y entre localidades muestreadas, estas fueron desde Chihuahua en el norte, hasta Quintana Roo en el sureste. Todos los loci fueron polimórficos y no se encontraron alelos privados. Se encontraron altos niveles de heterócigos cuando se compararon la proporción de alelos en cada locus. Comparaciones pareadas de F ST y R ST mostraron una diferenciación genética entre los individuos de las diferentes localidades. Los resultados de estructura genética indican la presencia de once clusters, con un alto porcentaje de asignación de los individuos a las localidades en donde fueron recolectados.

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