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Background  Malaria is a top cause of mortality on the island nation of Madagascar, where many rural communities rely on subsistence agriculture and livestock production. Understanding feeding behaviours of  Anopheles  in this landscape is crucial for optimizing malaria control and prevention strategies. Previous studies in southeastern Madagascar have shown that  Anopheles  mosquitoes are more frequently captured within 50 m of livestock. However, it remains unknown whether these mosquitoes preferentially feed on livestock. Here, mosquito blood meal sources and  Plasmodium  sporozoite rates were determined to evaluate patterns of feeding behaviour in  Anopheles  spp. and malaria transmission in southeastern Madagascar.    Methods  Across a habitat gradient in southeastern Madagascar 7762 female  Anopheles  spp. mosquitoes were collected. Of the captured mosquitoes, 492 were visibly blood fed and morphologically identifiable, and a direct enzyme-linked immunosorbent assay (ELISA) was used to test for swine, cattle, chicken, human, and dog blood among these specimens. Host species identification was confirmed for multiple blood meals using PCR along with Sanger sequencing. Additionally, 1,607  Anopheles  spp. were screened for the presence of  Plasmodium falciparum ,  P. vivax -210, and  P. vivax  247 circumsporozoites (cs) by ELISA.    Results  Cattle and swine accounted, respectively, for 51% and 41% of all blood meals, with the remaining 8% split between domesticated animals and humans. Of the 1,607  Anopheles  spp. screened for  Plasmodium falciparum, Plasmodium vivax  210 ,  and  Plasmodium vivax  247 cs-protein, 45 tested positive, the most prevalent being  P. vivax  247, followed by  P. vivax  210 and  P. falciparum . Both variants of  P. vivax  were observed in secondary vectors, including  Anopheles squamosus/cydippis ,  Anopheles coustani , and unknown  Anopheles  spp. Furthermore, evidence of coinfection of  P. falciparum  and  P. vivax  210 in  Anopheles gambiae  sensu lato ( s.l. ) was found.    Conclusions  Here, feeding behaviour of  Anopheles  spp. mosquitoes in southeastern Madagascar was evaluated, in a livestock rich landscape. These findings suggest largely zoophagic feeding behaviors of  Anopheles  spp., including  An. gambiae s.l.  and presence of both  P. vivax and P. falciparum  sporozoites in  Anopheles  spp. A discordance between  P. vivax  reports in mosquitoes and humans exists, suggesting high prevalence of  P. vivax  circulating in vectors in the ecosystem despite low reports of clinical vivax malaria in humans in Madagascar. Vector surveillance of  P. vivax  may be relevant to malaria control and elimination efforts in Madagascar. At present, the high proportion of livestock blood meals in Madagascar may play a role in buffering (zooprophylaxis) or amplifying (zoopotentiation) the impacts of malaria. With malaria vector control efforts focused on indoor feeding behaviours, complementary approaches, such as endectocide-aided vector control in livestock may be an effective strategy for malaria reduction in Madagascar.

Widespread zoophagy and detection of Plasmodium spp. in Anopheles mosquitoes in southeastern Madagascar