DOI: 10.5061/dryad.18931zd70
title: Data from: Diversity and plasticity in mosquito feeding patterns: a meta-analysis of ‘universal’ DNA diet studies

description: Although mosquitoes can have innate preferences for particular blood-meal
 hosts, their realised feeding patterns on different host species can be
 modified under climate and land use change, with implications for disease
 spread. It is therefore important to understand the niche breadth of
 vectors and to what extent shifts in feeding patterns are predictable. We
 investigated global shifts in feeding patterns among different functional
 and taxonomic groups of host species for six prominent disease-vectoring
 mosquitoes.Time period: 2000–2019. Major taxa studied: six
 disease-vectoring mosquito species: Aedes aegypti, Ae. albopictus,
 Anopheles funestus, An. gambiae, Culex pipiens, and Cx. quinquefasciatus.
 Focusing on blood meal studies that used universal molecular methods, we
 compiled evidence from &gt;15,600 blood-meals for the six mosquito
 species. We estimated mosquito’s host niche breadth, and we used
 hierarchical Dirichlet regression models to investigate shifts in feeding
 patterns in relation to human and livestock density, land use, and climate
 gradients. We estimated host ranges of 179–321 species for each of the two
 Culex mosquitoes and 24 - 65 species for Aedes mosquitoes, comprising
 considerably broader host niche breadths than previously anticipated. We
 found some evidence that shifts in feeding patterns among different host
 functional and taxonomic groups were associated with environmental
 conditions such as temperature and livestock density, while our results
 also demonstrate that, with the currently available evidence, global
 predictions of shifts in mosquito feeding patterns are challenged by
 considerable uncertainty. Our global metaanalysis afforded first insights
 into the shifts of feeding patterns in variable environments, suggesting
 that host choice is not a simple function of host availability, but
 contingent on other environmental drivers. Improving resolution and
 consistency of data gathering and reporting will improve the precision of
 how blood-meal studies can inform us of present and potential risks of
 pathogen transmission events.
description: We compiled blood meal data from the literature for six focal
 mosquito species: <em>Aedes aegypti</em>, <em>Aedes
 albopictus</em>, <em>Anopheles gambiae</em>,
 <em>Anopheles funestus</em>, <em>Culex pipiens
 </em>s.l., and <em>Culex quinquefasciatus</em>. These
 are of considerable interest as vectors of infectious diseases and feed to
 various extents on humans<em>.</em> We conducted a systematic
 literature search that reported counts of mosquitos feeding on different
 blood-meal hosts. Our initial search (August 2022) was broad and afforded
 us a “bird’s eye view” of the literature. For this, we used Web of
 Science, Pubmed, Scopus, GoogleScholar online databases with the search
 terms ‘bloodmeal*’, ‘blood meal’, blood-meal’, ‘feeding’, ‘habit’,
 ‘pattern*’, ‘preference*’, ‘interaction*’, combined with the terms
 ‘mosquito*’, ‘vector*’, ‘vector-host’, ‘host*’, ‘vertebrate*’, ‘animal*’.
 The asterisk (‘*’) was used to search for possible variations of terms.
 Our subsequent systematic search of the literature (July-September 2023)
 focused on Web of Science and involved two steps. The first used the
 phrase ‘bloodmeal*’ OR ‘blood meal’ OR 'blood-meal’” and the names
 (including synonyms) of the focal mosquito species. If the search returned
 too many irrelevant hits (&gt;200), then additional ‘methods’ search
 terms were used (refer to Table S1 in associated publication). Another
 search used publications of popular PCR primer sets as a search term
 (refer to Table S1 in associated publication). If the search returned too
 many irrelevant hits, then the number of linked articles was refined using
 the search term ‘mosquito*’ (refer to Table S1 in associated publication).
 These searches resulted in 1,495 published articles that we subsequently
 filtered with the following criteria: 1) The arthropod
 vector was one of six focal species 2) PCR primers used
 to identify blood-meal hosts were ‘universal’ (for
 vertebrates/mammals/birds) and not targeted to particular
 species 3) Either Sanger (dideoxy) sequencing or
 high-throughput DNA sequencing is used at some point to confirm host
 identity 4) Spatial information was provided as either
 geographical coordinates or salient location names and sampling sites were
 not pooled over spatial extents &gt;50km in diameter
 5) Host species identity data were available
 6) For any given mosquito species and sampling site, there must
 be ≥7 bloodmeals (corresponding to an ~80% chance of detecting an uncommon
 host present in ~20% of samples). This resulted in 86
 studies that covered 121 different study sites (some studies provided data
 for multiple sites, refer to Figure S1 in associated publication for
 location map, refer to Table S4<strong> </strong>in associated
 publication for references) with data collected between 2000 and 2019.
 From these, we generated a database of geographical coordinates, counts of
 blood-meals, and the host species (resolved to smallest taxonomic unit)
 for each mosquito species. Counts of blood-meal origins were assigned to
 both hosts in the sixteen studies that recorded mosquitos feeding on
 multiple hosts (e.g., ‘human + cow’). All extracted scientific species
 names were checked and aligned to the Integrated Taxonomic Information
 System (ITIS) database, using the <em>taxize</em> 0.9.1 R
 package. For studies with missing geographical
 coordinates, we identified coordinates for location names at smallest
 available administrative units using various online maps and search
 engines. For studies with multiple sites/coordinates from a single region,
 we computed the average of coordinates for use in analyses.
description: # Data from: Diversity and plasticity in mosquito feeding patterns: a
 meta-analysis of ‘universal’ DNA diet studies
 [https://doi.org/10.5061/dryad.18931zd70](https://doi.org/10.5061/dryad.18931zd70) ## Description of the data and file structure We compiled blood meal data from the literature (see methods) for six focal mosquito species: *Aedes aegypti*, *Aedes albopictus*, *Anopheles gambiae*, *Anopheles funestus*, *Culex pipiens* s.l., and *Culex quinquefasciatus*. For detailed method description, references, and description of the analysis, refer to the published paper: Lee, M., R. O'Rorke, N. J. Clark, T. U. Webster, and K. Wells. (2025) Diversity and Plasticity in Mosquito Feeding Patterns: A Meta-Analysis of ‘Universal’ DNA Diet Studies.” Global Ecology and Biogeography 34, no. 6: e70077. [https://doi.org/10.1111/geb.70077](https://doi.org/10.1111/geb.70077). The provided data set consists of three files with computer code in the programming language R ([https://cran.r-project.org/](https://cran.r-project.org/)), six data files and one file with metadata descriptions. The three file with R computer code are as follows: R1) "RCode_Data_formatting.txt": R Code for basic data preparation and formatting steps (refer to methods descriptions for details) . R2) "RCode_predictor_variables.txt": R code to gather remotely sensed predictor variables and match to site-level coordinates (refer to methods descriptions for details). R3) "RCode_Dirichlet_regression_fit_to_data.txt": R code to run the analysis (a Dirichlet regression analysis in the computer programming language Stan via an R interface; refer to methods descriptions for details). The six data files contain details of the number of blood meals for any given mosquito species at any given study site from different functional or taxonomic host species groups and the associated environmental data. Details of the data are described in the method description and the file “metadata_bloodmeal.studies.xlsx”. Note that missing data/ empty cells are indicated with “NA” (if loaded into the R computer software, ‘NA’ are categorised as missing data). D1) “Data_Genera.all_taxonomical.host.groups.csv”: contains data on the feeding patterns of the focal mosquitoes species, whereby all data for the two focal *Culex* species (*Culex pipiens* s.l. and *Culex quinquefasciatus*) are aggregated to ‘*Culex pipiens complex*’. Feeding patterns on different host species groups distinguishes in this data set among taxonomical host groups. See metadata and method descriptions for details. D2) “Data_Genera.all_functional.host.groups.csv”: contains data on the feeding patterns of the focal mosquitoes species, whereby all data for the two focal *Culex* species (*Culex pipiens* s.l. and *Culex quinquefasciatus*) are aggregated to ‘*Culex pipiens complex*’. Feeding patterns on different host species groups distinguishes in this data set among functional host groups. See metadata and method descriptions for details. D3) “Data_Culex.species_taxonomical.host.groups.csv”: contains data on the feeding patterns of the focal mosquitoes species, whereby data for the *Culex* species are given for *Culex pipiens* s.l. and *Culex quinquefasciatus* Feeding patterns on different host species groups distinguishes in this data set among taxonomical host groups. See metadata and method descriptions for details. D4) “Data_Culex.species_functional.host.groups.csv”: contains data on the feeding patterns of the focal mosquitoes species, whereby data for the *Culex* species are given for *Culex pipiens* s.l. and *Culex quinquefasciatus* Feeding patterns on different host species groups distinguishes in this data set among functional host groups. See metadata and method descriptions for details. D5) “Data_Culex.subspecies_taxonomical.host.groups.csv”: contains data on the feeding patterns of the focal mosquitoes species, whereby data for the *Culex* species *Culex pipiens* s.l. are distinguished among the tow biotypes *Culex pipiens f. pipiens* biotype and *Culex pipiens f. molestus biotype*. Feeding patterns on different host species groups distinguishes in this data set among taxonomical host groups. See metadata and method descriptions for details. D6) “Data_Culex.subspecies_functional.host.groups.csv”: ”: contains data on the feeding patterns of the focal mosquitoes species, whereby data for the *Culex* species *Culex pipiens* s.l. are distinguished among the tow biotypes *Culex pipiens f. pipiens* biotype and *Culex pipiens f. molestus biotype*. Feeding patterns on different host species groups distinguishes in this data set among functional host groups. See metadata and method descriptions for details. ### Files and variables #### Files: Data for functional/taxonomical host groups (applies to all six data files): **Description:**  ##### Variables * Study_ID: unique ID (identification number) for the studies, from which data have been extracted * Site_ID: unique ID (identification number) of the site for which data have been extracted; there might be multiple sites reported for any study * Study_Code: unique study code (comprised of first author name, year of publishing and first words from title) * Study_title: Study title * Country: Country where a study reported data from * Site: Name of study site/region/location as mentioned in studies * Vector: Binomial name of vector species * Vector_subtype: Name of vector subspecies (as relevant for Culex mosquitoes) * Latitude: Latitude value of the geographic coordinate fo the study site (WGS84) * Longitude: Longitude value of the geographic coordinate fo the study site (WGS84) * Indoors_Outside: Characterisation if study reported samples from indoor or outdoor sampling or both * Year: Year in which data have been collected as reported in a study * Habitat_local: Habitat characterisation based on site descriptions in study * Spatial.extent_m: Spatial extent of the local study region based on information provided in a study * n_total: Total number of blood meal reported for a given focal mosquito species * n_human: Number of blood meals from humans reported for a given focal mosquito species (for functional host species grouping) * n_mammPet: Number of blood meals from mammalian pet species reported for a given focal mosquito species (for functional host species grouping) * n_mammFarmed: Number of blood meals from mammalian farmed/domestic species reported for a given focal mosquito species (for functional host species grouping) * n_mammWild: Number of blood meals from mammalian wildlife species reported for a given focal mosquito species (for functional host species grouping) * n_birdDomest: Number of blood meals from domesticated bird species reported for a given focal mosquito species (for functional host species grouping) * n_birdWild: Number of blood meals from wild bird species reported for a given focal mosquito species (for functional host species grouping) * n_others: Number of blood meals from host species that did not match any of the groups used in this study o reported for a given focal mosquito species (for functional host species grouping) * n_birdZoo: Number of blood meals from bird species kept in a zoo reported for a given focal mosquito species (for functional host species grouping) * n_mammZoo: Number of blood meals from mammal species kept in a zoo reported for a given focal mosquito species (for functional host species grouping) * n_unidentified: Number of blood meals for which the host species were not identified (for functional host species grouping) * n_Homo: Number of blood meals from humans reported for a given focal mosquito species (for taxonomical host species grouping) * n_Gallus: Number of blood meals from chicken (Gallus sp.) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Canidae: Number of blood meals from Canidae reported for a given focal mosquito species (for taxonomical host species grouping) * n_Felidae: Number of blood meals from Felidae reported for a given focal mosquito species (for taxonomical host species grouping) * n_Bovidae: Number of blood meals from Bovidae reported for a given focal mosquito species (for taxonomical host species grouping) * n_Mammalia: Number of blood meals from Mammalia (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Cardinalidae: Number of blood meals from Cardinalidae (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Columbidae: Number of blood meals from Columbidae (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Corvidae: Number of blood meals from Corvidae (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Turdidae: Number of blood meals from Turdidae (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Aves: Number of blood meals from Aves (others than those considered with higher taxonomic resolution) reported for a given focal mosquito species (for taxonomical host species grouping) * n_Reptilia.Amphibia: Number of blood meals from reptiles and amphibians reported for a given focal mosquito species (for taxonomical host species grouping) * n_unidentified: Number of blood meals for which the host species were not identified or did not match any of the groups used in this study