during 20072012 , we sampled 1,220 bats representing at least 48 species from multiple locations in selected countries in africa ( table 1 ) . bats were anesthetized with the use of ketamine ( 0.050.1 mg / g body mass ) and exsanguinated by cardiac puncture . voucher specimens were identified through morphologic characterization ( 7 ) or , alternatively , through genetic barcoding . approximately 30100 mg of renal tissue was used for rna extraction . a heminested primer set targeting the conserved polymerase ( large ) gene of respirovirus , morbillivirus , and henipavirus was used for sample screening through reverse transcription pcr ( 8) . a total of 103 samples ( 8.4% ) tested positive , and the obtained amplicons of 490 bp were sequenced ( technical appendix table 1 ) . for phylogenetic analysis , representative paramyxovirus sequences available from genbank were included ( technical appendix table 2 ) , and bayesian analysis was performed by using beast version 1.7.4 software ( http://beast.bio.ed.ac.uk/ ) ( figure ) . * the sampling protocol was approved by the institutional animal care and use committee of the centers for disease control and prevention ; protocol 2096framulx - a3 and the university of pretoria animal ethics committee ( ec05414 ) . maximum clade credibility tree based on partial polymerase ( large ) gene sequences ( 439 bp ) of paramyxoviruses built in beast version 1.7.4 software ( http://beast.bio.ed.ac.uk/ ) , applying the general time reversible plus invariant sites plus gamma model inferred by jmodeltest version 0.1.1 ( 10 ) . genus - specific clusters are indicated on the right and show possible opportunistic infections in other species grouping within these clusters . several samples from bat species not previously implicated as paramyxovirus reservoirs tested positive in our study . sequence analysis of paramyxovirus sequences showed a clear bifurcation of the phylogenetic tree , segregating paramyxoviruses detected in pteropodid bats ( pteropodidae ) from paramyxoviruses detected in bats of other families ( figure ) . two viral sequences detected in rousettus aegyptiacus bats grouped within this cluster as part of a sister clade to the henipaviruses . some of these sequences grouped with the sequences from the morbillivirus and proposed jeilongvirus genera , whereas others could not be included in any of the other paramyxovirus genera . we observed a strong association of several viral lineages to particular bat genera for paramyxoviruses identified in hipposideros , miniopterus , coleura , myotis , and pipistrellus bats , although the bats were sampled from geographically distant locations . in contrast to the sequences of european and south american origin , for which geographic clustering was observed , no such clustering was found among the sequences from african bats . the incidence and diversity of viral sequences varied according to bat species . sampled from a single colony in the democratic republic of the congo ( n = 40 ) . in other cases , several distinct viral sequences were detected in different individual bats of 1 species , such as miniopterus minor bats sampled from a single colony in kenya ( n = 53 ) , which harbored 6 distinct viral sequences . some of the sequences were found more frequently than others . in contrast to a previous study which did not identify paramyxoviruses in coleura afra bats sampled in ghana ( n = 71 ) ( 4 ) , we detected a substantial paramyxovirus incidence ( 37% , n = 27 ) in the same bat species sampled in kenya ( table 2 ) . tissue type stated for positive samples only and may not indicate all tissues sampled . central africa refers to gabon / republic of congo / drc / republic of central africa . the henipaviruses were the first bat paramyxoviruses directly linked to human disease ; however , most aspects of pathogenicity and the host ranges of the increasingly detected novel bat paramyxoviruses remain to be investigated . here we report information regarding paramyxovirus distribution through molecular evidence of bat - associated paramyxoviruses in cameroon , nigeria , and south africa , as well as evidence of paramyxoviruses in nonpteropodid bats from the democratic republic of the congo . our results suggest that 2 separate lineages were established during the evolution of bat - associated paramyxoviruses : the pteropodid bats potentially harbor 1 lineage , and the nonpteropodid bats potentially harbor the other . in contrast to the proposed chiropteran classification , which supports a sister - taxon relationship between rhinolophoidae and pteropodidae on the suborder level , paramyxovirus divergence appears to correlate with traditional bat taxonomy . the evolution behind this divergence might be a result of multiple evolutionary origins or a single origin with subsequent divergence . as with the evolution of echolocation , this question remains to be answered ( 11 ) . more extensive bat sampling and molecular dating of the paramyxovirus phylogeny may help resolve this question . intensified anthropogenic transformations have facilitated closer contact between humans , domestic animal populations , and wildlife . our study demonstrates that some bat species , adapted to peridomestic roosting , can have a substantial incidence of diverse paramyxoviruses . the variation in incidence and viral diversity observed in several bat species may suggest that some species are the true reservoirs , whereas others are mere incidental hosts . given the observed virus diversity , implications for public health and veterinary medicine should be taken into account , especially considering the known likelihood of direct bat - to - human and human - to - human transmission of nipah virus ( 12 ) . enhanced surveillance in bats and other animals clearly , systematic longitudinal studies are needed to elucidate critical factors of paramyxovirus circulation within bat communities ( 13 ) , and further research is needed to clarify the pathobiology , tissue tropism , and excretion pathways of these novel paramyxoviruses because these factors can be directly related to their zoonotic potential .
as part of a larger survey for detection of pathogens among wildlife in sub - saharan africa conducted during 20072012 , multiple diverse paramyxovirus sequences were detected in renal tissues of bats . 
 phylogenetic analysis supports the presence of at least 2 major viral lineages and suggests that paramyxoviruses are strongly associated with several bat genera .