PLoS PathogensPathologyPeer-reviewed journal article
Stomoxys flies (Diptera, Muscidae) are competent vectors of multiple livestock hemopathogens
Julia W. Muita, Joel Ltilitan BarguliD, JohnMark O. MakwattaiD, E Ngatia, Simon K. TawichiD, Daniel K. MasigaiD, Merid Negash GetahuniD
Abstract
Stomoxys flies are widely distributed and economically significant vectors of various livestock pathogens of veterinary importance. However, the role of Stomoxys spp. in pathogen transmission is poorly understood. Therefore, we studied the feeding patterns of these blood feeders collected from specific locations in Kenya, to identify various vertebrate hosts they fed on, and the livestock hemopathogens they carried, to elucidate their role in pathogens transmission. Our findings show that field-collected Stomoxys flies carried several pathogens including Trypanosoma spp., Anaplasma spp., and Theileria spp. that were also found in the blood of sampled livestock, namely camels and cattle. The findings on blood meal analysis show that Stomoxys flies fed on a variety of domestic and wild vertebrate hosts. We further determined whether Stomoxys spp. are vectors of hemopathogens they harbored by studying the vector competence of S. calcitrans, S. niger niger, and S. boueti species complex, through laboratory and natural experimental in vivo studies. We show that in the process of blood feeding Stomoxys spp. complexes can transmit T. evansi (8.3%) and T. vivax (30%) to Swiss white mice. In addition, field-collected Stomoxy spp. were exposed to healthy mice for blood meal acquisition, and in the process of feeding, they transmitted Theileria mutans and Anaplasma spp. to Swiss white mice (100% infection in the test mice group). All mice infected with both trypanosomes via stomoxys bite died while those infected with Theileria and Anaplasma species did not, demonstrating virulence difference between pathogens. The key finding of this study showing broad feeding host range, cosmopolitan, plethora of pathogens harboured, and efficient vector competence in spreading multiple pathogens suggests profound role of Stomoxys on pathogen transmission and infection prevalence in livestock. Author summaryStomoxys flies are highly adaptable to several ecological settings, including metropolitan areas. In contrast, tsetse flies (genus Glossina), the main biological vectors of African trypanosomes, have a limited distribution to parks and other conservation areas. Stomoxys flies could play a significant role in the spread of animal African trypanosomes, among other hemopathogens, particularly in areas with or without tsetse infestation. Although there have been speculations about the potential role of Stomoxys flies in the transmission of various pathogens, there is lack of data to link hemopathogens occurring in both bloodmeal hosts of Stomoxys and in the flies, and further in vivo experimental studies to confirm the vector competence of Stomoxyine flies. Here, we explored a host and pathogens network, and investigated species diversity at various ecologies, and demonstrated that Stomoxys flies feed on diverse vertebrate hosts and are infected with a plethora of pathogens. We also showed experimentally that they could transmit some of these hemopathogens to mice, for instance, T. vivax, T. evansi, Theileria mutans, and Anaplasma spp. with varying infection success rates. Stomoxys flies could play a significant role in transmitting and spreading various hemopathogens of veterinary importance and possibly maintaining their circulation in livestock, which could explain the occurrence of animal African trypanosomes in the regions outside the tsetse belts.
- organism: multiple species
- assay: proteomics
- assay: metagenomics
- research focus: host-pathogen & immune response
- approach: experimental assay
In-depth summary
Claude Sonnet 4.6 · xhigh effort · 8 Nov 2026 · from the full text of version 1 (8 Oct 2024) · CC BY 4.0
Across nine Kenyan counties, Stomoxys flies fed on 15 vertebrate hosts, carried Anaplasma, Theileria/Babesia and Trypanosoma DNA, and transmitted T. vivax, T. evansi and Theileria mutans to mice.
The paper says data are lacking that link hemopathogens found in the blood-meal hosts of Stomoxys to those found in the flies, and that in vivo experiments confirming vector competence are also lacking. It combines field trapping, blood-meal and pathogen screening, and mouse transmission assays to address this.
Key results
ShownSix Stomoxys species were found across the nine counties. S. calcitrans was the most abundant, and species diversity varied strongly by ecology.
11,323 flies collected; S. calcitrans n=5,547 (49%), S. niger niger 2,938, S. boueti 1,471, S. niger bilineatus 778, S. sitiens 495, S. taeniatus 94. Shannon index 1.36 in Kwale vs 0.29 in Meru. · 3.1 Stomoxys species diversity relative…
ShownStomoxys flies fed on a wide range of wild and domestic hosts, including humans. Cattle were the most frequently detected host, and some flies carried blood from more than one host.
Hosts were identified in 225 blood-fed flies, out of 3,451 flies screened, by PCR-high-resolution melting. 15 host species were found. Double melt peaks, mostly cattle with goats, sheep or camels, indicated multiple hosts. · 3.2 Stomoxys flies blood meal host network…; 4. Discussion
ShownThe same hemopathogen genera (Anaplasma, Theileria/Babesia, Trypanosoma) occurred in livestock and Stomoxys. Anaplasma was most common in camels (64.7%) and Stomoxys (49.1%); in cattle Theileria/Babesia (56.6%) slightly exceeded Anaplasma (54.1%). Ehrlichia and Rickettsia occurred in livestock, not flies.
Camels (n=452): Anaplasma 64.7%, Trypanosoma 12.3%. Cattle (n=124): Theileria/Babesia 56.6%, Anaplasma 54.1%. Stomoxys (n=3,451): Anaplasma 49.1%, Theileria/Babesia 19.1%, Trypanosoma 9.1%. Glossina pallidipes (n=1000): Trypanosoma 7.5%, Anaplasma 4%, Theileria/Babesia 11%. · 3.3 Stomoxys flies and domestic animals harbor…; 4. Discussion
SuggestedLaboratory Stomoxys transmitted T. evansi and T. vivax to mice with interrupted feeding, though success was modest and based on few trials. T. vivax survived much longer in the fly gut than T. evansi.
T. evansi: 8.3% (2/24) mice infected; live parasites in gut up to 5 h. T. vivax: 30% (3/10) transmission; live parasites at 16 h. G. pallidipes transmitted T. evansi in 5 of 5 trials. · 3.4 Stomoxys flies are competent mechanical…
SuggestedField-caught Stomoxys given to healthy mice after delayed feeding transmitted Theileria mutans. Anaplasma spp. infection in mice was seen by microscopy only.
T. mutans sequence GenBank PP918990 from recipient mice; mice showed no clinical signs and no mortality for over 120 days. The number of mice and trials is not given in the text. · 3.5 Natural pathogen transmission assays through…
Methods, data and reproducibility
Flies trapped in nine Kenyan counties were identified (morphology, CO1) and screened by PCR and Sanger sequencing; blood meals were typed by PCR-high-resolution melting. Vector competence: lab-colony flies fed on infected donor mice, were interrupted, and moved to healthy mice; field-trapped flies were placed with restrained healthy mice for 30 minutes; mice were checked by microscopy and PCR.
- Field-collected Stomoxys flies, Kenya, Oct 2021 to Nov 2023; CO1 sequences deposited for S. boueti and S. taeniatus · Stomoxys spp. · n = 11,323 collected; 3,451 screened for pathogens; 225 blood-fed identified · PP587243; PQ203543
- Camel and cattle blood screened for hemopathogens · Camelus dromedarius; Bos taurus · n = 452 camels; 124 cattle
- Glossina pallidipes co-occurring with Stomoxys, screened for pathogens for comparison · Glossina pallidipes · n = 1000
- In vivo transmission assays in Swiss White mice with lab-colony Stomoxys (T. evansi from a Marsabit camel; T. vivax IL 2136) · Mus musculus · n = 24 mice for T. evansi; 10 for T. vivax
- Field bioassay with wild-caught flies; T. mutans sequence from recipient mice · Mus musculus · PP918990
- Methods & tools
- PCR with Sanger sequencing, PCR-high-resolution melting, BLAST / GenBank, PhyML v. 3.0, FigTree v. 1.4.4, R 4.4.1, bipartite R package, GraphPad, QGIS v.3
- Data & accessions
- https://www.ncbi.nlm.nih.gov/genbank/
- Reagents & environment
- R 4.4.1; PhyML v. 3.0; FigTree v. 1.4.4; QGIS v.3. No container or requirements file stated.
- Reproducibility gaps
- No analysis code is given. Data are said to be in the manuscript and supplementary tables, with sequences deposited in NCBI. Primer details are in a supplementary table.
Limitations (4)
- Vector competence was tested on the mixed natural Stomoxys species complex (mainly S. calcitrans, S. niger niger and S. boueti). Species-specific competence and tests in relevant livestock hosts remain to be done. · stated by the authors
- Only 225 of 3,451 flies yielded an identifiable blood meal. The authors suggest host DNA degradation during digestion and the small blood volume taken as explanations. · stated by the authors
- Transmission rates rest on few mouse trials (2/24 for T. evansi, 3/10 for T. vivax). Mice, not livestock, were the recipients. The authors also report that transmission depended on feeding time and parasitemia. · noted in this summary
- Pathogen detection in flies is by genus-level PCR on whole-fly DNA, so positives may reflect ingested blood rather than established infection. Anaplasma in recipient mice was seen by microscopy only. · noted in this summary
Read this if you work on vector-borne livestock disease, mechanical transmission of trypanosomes, or fly blood-meal and pathogen network surveys outside tsetse areas.
- Citations
- 16
- FWCI
- 9.74
- Field percentile
- top 1%
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