The distribution of triatomine (Hemiptera: Reduviidae) vectors of Trypanosoma cruzi (Kinetoplastida: Trypanosomatidae) in Illinois and Missouri: historical records and specimen submissions from community science programs

Abstract

Triatomine species (kissing bugs) infected with Trypanosoma cruzi are found across the southern United States. The northern limits of Trypanosoma cruzi infected kissing bugs are less understood. The objective of this work was to describe the locations of kissing bugs from Illinois and Missouri based on historical records, submissions to Texas A&M University’s (TAMU) Kissing Bug Community Science Program and the Centers for Disease Control and Prevention (CDC), and records from online platforms (iNaturalist, BugGuide, and GBIF) up to and including 2022. A total of 228 records were discovered, including 186 from historical or observation platforms and 42 specimens submitted to TAMU or CDC. Species included Triatoma sanguisuga (221 total records, 9 nymphs) and Triatoma lecticularia (7 records). Notably, nearly all (24/26) records submitted to TAMU were collected indoors. Twelve of the 30 (40%) specimens tested were positive for the presence of T. cruzi, including parasite discrete taxonomic units TcI and TcIV. One triatomine sample had been found in a bed feeding on the submitter; this bug was positive for T. cruzi and had evidence of human blood in its gut. Records suggest a ubiquitous distribution in Missouri and potentially to the northernmost border in Illinois. Further investigations into triatomine distribution and infection status are needed within states assumed to be northern limits in order to create public health and veterinary health messaging and baseline distributional maps from which to measure future range shifts in relation to a changing climate.

Introduction

Commonly known as “kissing bugs”, triatomine insects (Order Hemiptera, Family Reduviidae, Subfamily Triatominae) are vectors of the parasite Trypanosoma cruzi, the causative agent of Chagas disease. Vector-borne transmission of T. cruzi to humans and animals occurs when the parasite is introduced to the bloodstream, typically when an infected kissing bug defecates near a bite or wound or is ingested. Chagas disease, also known as American trypanosomiasis, is estimated to affect nearly 6 million people throughout Latin America (Bern et al. 2019) and nearly 300,000 people in the United States (Manne-Goehler et al. 2016, Irish et al. 2022). Although the risk of vector-borne transmission to humans in the United States is potentially low, mounting evidence from bloodmeal analyses indicates that human-kissing bug interactions may be more common than previously thought. This includes, for example, an autochthonous (locally-acquired) case of Chagas disease reported in Missouri in 2018 (Turabelidze et al. 2020). Across Texas, bloodmeal analysis of kissing bugs in one study revealed that 40 (65%) of the collected insects had fed on humans (Gorchakov et al. 2016). In Arizona, human blood was found among all of 8 randomly selected free-roaming kissing bugs at a popular outdoor attraction (Klotz et al. 2014a).

Due to the sylvatic nature of the 11 triatomine species found in the United States, the geographical ranges of these vectors in the United States have been difficult to elucidate in an efficient and comprehensive way. Historical literature from as early as 1855 (Le Conte 1855) and subsequent work by many researchers and community scientists have slowly pieced together likely ranges of the species found in the United States (Lent and Wygodzinsky 1979, Bern et al. 2011, Zeledón et al. 2012, Curtis-Robles et al. 2018a). The 2 species reported in Illinois and Missouri are Triatoma sanguisuga (the Eastern Bloodsucking Conenose bug) and Triatoma lecticularia (also known as Paratriatoma lecticularia (de Paiva et al. 2021, Bern et al. 2011, McPherson 2017). Triatoma sanguisuga and T. lecticularia likely have northern range limits in the midwestern US states of Illinois and Missouri (Bern et al. 2011) (Fig. 1). However, historical literature reveals relatively little about the vector or T. cruzi in these states. T. sanguisuga appears to be first recorded in Illinois in the late-1800s, with mention in a Good Housekeeping article authored by entomologist C.V. Riley (Riley 1889,, 1890). In addition, various developmental stages of T. sanguisuga in sylvatic habitats and house infestations in southern and western Illinois were reported in the 1960s (Porter 1965). T. lecticularia has reportedly been collected at lights in Missouri (Bern et al. 2011).

Sylvatic transmission spillover may be a risk to people and domestic dogs living in Illinois and Missouri. Evidence of T. cruzi infection in raccoons sampled from Illinois and Missouri in the early 2000s (Vandermark et al. 2018) suggests the maintenance of a sylvatic transmission cycle. Understanding the current northern limits of triatomine distribution would allow for the detection of a possible northern expansion, for example, as a result of climate change.

While museum collections are a critical and time-tested means of documenting and studying biological occurrences, the internet has expanded and expedited the ability of laypeople to contribute to species documentation. There is an expanding appreciation for the role community science programs can play in collecting and identifying insect vectors most pertinent for human health (Hamer et al. 2018). Citizen and community science programs are used for a wide range of disease vectors including kissing bugs (Dumonteil et al. 2009, Curtis-Robles et al. 2015, Ceccarelli et al. 2020, Dye-Braumuller et al. 2021, Cochero et al. 2022, Delgado-Noguera et al. 2022), ticks (Nieto et al. 2018, Eisen and Eisen 2021, Lyons and Tuten 2021, Porter et al. 2021), and mosquitoes (Jordan et al. 2017, Tarter et al. 2019, Pernat et al. 2021). Community science enlists nonexpert members of the public to help in various aspects of data collection. For vector surveillance activities, this typically involves the identification and collection of insect vector species that can then be tested for the presence of pathogens. In addition to community science programs, online identification and observation sites such as iNaturalist (iNaturalist.org) and BugGuide (BugGuide.net) provide platforms for contemporary records. Over the past 2 decades, the Global Biodiversity Information Facility (GBIF) (GBIF.org) has gained traction as a repository for a myriad of types of biological records.

To date, no systematic investigations of the geographic limits and distribution of triatomine insects have been conducted in Illinois and Missouri. The objective of this work was to describe triatomine occurrences from Illinois and Missouri to lay the groundwork for future investigations into triatomine distribution and parasite prevalence within these 2 states that are assumed to be northern boundaries of T. cruzi infected triatomines.

Materials and Methods

Historical Records

Records from the Illinois Natural History Survey (INHS), the Illinois State Museum, and Enns Entomology Museum of the University of Missouri were compiled. Using the INHS search function of the Prairie Research Institute’s Insect Collection, all records of the genera Triatoma and Paratriatoma were searched on 18 October 2022. All records of T. sanguisuga, T. lecticularia, and P. lecticularia with a recorded location in Illinois or Missouri were documented (McElrath 2022). Records from the Illinois State Museum Collection were shared by coauthors MM and JS, and records from Enns Entomology Museum were shared by the Collections Manager, K. Simpson.

Submissions to Texas A&M University and Centers for Disease Control and Prevention

We compiled all reports of kissing bugs to the Texas A&M University (TAMU) Kissing Bug Community Science Program (Curtis-Robles et al. 2015, TAMU 2022) and submissions to Centers for Disease Control and Prevention (CDC) from Illinois and Missouri. Reports to TAMU were initiated by community scientists via an online form to submit a photograph and information about the location and behavior of the sample. Our team screened all reports to confirm that the insect was a triatomine and shared instructions for submitting to our laboratory. For bugs found in bedrooms (and occasionally other indoor locations), submitters were encouraged to submit to their respective state health departments for routing to CDC for identification and testing. This step allowed for appropriate public health response to findings of T. cruzi infected triatomines. For bugs found elsewhere indoors or outdoors, individuals were encouraged to submit the specimen to the TAMU program for processing.

Triatomines submitted to TAMU were stored at 4 °C until identified (Lent and Wygodzinsky 1979) and dissected as previously described (Curtis-Robles et al. 2018b). Evidence of a recent bloodmeal was scored based on examination of guts (1 = no blood, desiccated guts; 2 = no blood, guts visible; 3 = traces of blood in gut; 4 = blood present, but either not much or not fresh [dried]; 5 = large amount of fresh blood) (Curtis-Robles 2018b). After dissection, guts were stored at −20 °C and/or −80 °C until extraction. DNA from hindgut tissue was extracted using the Omega E.Z.N.A Tissue DNA kit (Omega Bio-Tek, Norcross, GA). Samples were subjected to multiple PCRs for detection (Cruzi 1/2/3 probe-based qPCR (Duffy et al. 2013, Cura et al. 2015) and strain-typing (SL-IR qPCR, [Cura et al. 2015]) of T. cruzi DNA, as previously described (Curtis-Robles et al. 2018c). Triatomines submitted to CDC were stored at room temperature in 70% ethanol until genomic DNA extraction. Prior to extraction, the abdomen was severed from the rest of the body with a scalpel and allowed to dry overnight. DNA extraction was performed using the QIAamp DNA Blood Mini Kit (Qiagen, Valencia, CA). Extracted DNA was subjected to 2 conventional PCRs for detection of T. cruzi (Kinetoplast Minicircle and Nuclear TCZ) (Virreira et al. 2003). Samples positive for T. cruzi underwent a subsequent hemi-nested PCR to detect the presence of a mammalian bloodmeal (Roellig et al. 2013). Sanger sequencing was performed on samples positive for the presence of a mammalian bloodmeal to determine the exact source. Analyzed sequences were compared to known sequences via Basic Local Alignment Search Tool (BLAST) search with National Center for Biotechnology Information (NCBI) GenBank (Clark et al. 2016).

Records from Community-Driven Platforms

To gain a more complete understanding of triatomine distribution in Illinois and Missouri, we compiled reports from other community-driven platforms (iNaturalist and BugGuide) (iNaturalist 2022, BugGuide 2022) and GBIF (GBIF 2022). Since data are shared among these occurrence databases, we carefully reviewed all records to avoid counting duplicate records. First, iNaturalist was searched for all research grade submissions of T. sanguisuga and P. or T. lecticularia as of 31 August 2022; research grade submissions, as per iNaturalist data quality definitions, were those with a photo, observation date and geographic coordinates, and a taxon for which at least 2/3 identifiers agree (iNaturalist Contributors 2022). Next, BugGuide was searched for pages containing taxon identifier numbers 4789 (Triatoma), 478629 (T. lecticularia), and 59564 (T. sanguisuga) as of 1 September 2022, then updated on 21 October 2022. For all records, author E.S. visually confirmed that images matched morphological characteristics of the submitted species using the Lent and Wygodzinsky keys (Lent and Wygodzinsky 1979). Finally, GBIF was searched for entries for Triatoma sanguisuga (LeConte, 1856) and Paratriatoma or Triatoma lecticularia (Stȧl, 1859) on 1 September 2022. Each record from the historical collections data and the online databases were compared, and only unique records were counted. GBIF occurrences were also excluded if no date was listed and there were already existing county records from other sources. These exclusions decisions were made to allow for greater certainty in occurrence distribution by county, but specimen counts within counties may not be as accurately reflected. The database of county locations and specimen sources is included as a Supplementary File.

Results

Historical and Community-Driven Platforms

A total of 228 records (223 T. sanguisuga, 9 nymphs, 5 T. lecticularia) from Illinois and Missouri were revealed in historical databases and from community-driven online platforms (Fig. 2). The INHS Insect Database contained 29 known T. sanguisuga samples (McElrath 2022), 4 of which were cross-posted in GBIF. Two additional adult T. sanguisuga samples were found as recently as April (on a nature trail near a state office building) and July (inside a home being played with by cats) of 2022 in Sangamon County by authors JS and MM. One of the bugs (July 2022) has been submitted to authors ES and CS for testing and to be entered into a new local tracking system, and the other (April 2022) is in the Illinois State Museum Collections. Historical records from the Illinois State Museum contain 2 additional specimens from Sangamon (August 2012) and Union (July 2000) Counties (Fig. 2). There are 88 specimens with known Missouri collection locations in the Enns Entomology Museum Collections; 3 of these are T. lecticularia and the remaining 85 are T. sanguisuga.

There were 9 unique records from Bug Guide (7 MO, 2 IL) (BugGuide 2022), 16 unique records from iNaturalist (13 MO, 3 IL) (iNaturalist 2022), and 17 unique records from GBIF (14 MO, 3 IL) (GBIF 2022, A.J. Cook Arthropod Research Collection 2022, Ceccarelli et al. 2022). The remaining 23 records (16 MO, 7 IL) from these databases were recorded in both iNaturalist and GBIF.

Submissions to TAMU Community Science Program and CDC

The earliest submission of a kissing bug from Illinois or Missouri to TAMU or CDC was in 2016, and the most recent submission was in 2022. A total of 42 kissing bug records were submitted by 39 individuals to TAMU or CDC from Illinois and Missouri (3 and 39 bugs, respectively). In Illinois, bugs were collected in 3 counties: Greene, Jersey, and Union (Fig. 2). In Missouri, bugs were found in at least 15 counties: Adair, Bates, Boone, Caldwell, Christian, Franklin, Howell, Jackson, Jefferson, Lewis, Newton, Ralls, Randolph, St. Francois, St. Louis, and 1 submission from “northwestern Missouri” (Fig. 2).

With the exception of 2 female adults and 2 unrecorded sex T. lecticularia from Missouri, all specimens submitted to TAMU or CDC were identified as adult T. sanguisuga (14 female, 10 male, 14 not recorded). For the 3 samples from Illinois and 7 samples from Missouri that were not submitted for processing, identifications were made based on photos the community scientists had submitted. Of the 24 submissions that indicated whether the triatomine was alive or dead when found, 19 had been found alive. Live specimens were collected from April through November, although more frequently in July (6) and August (7). Of the 19 live-found bugs, 15 submitters had indicated the time of discovery: 10 had been discovered between evening (~7:00 pm) and early morning (~7:00 am), while 5 had been discovered mid-day.

A variety of collection locations were reported, although almost exclusively indoors (92.3%; Table 1). Locations indoors included bedrooms, dining room, bathroom, kitchen, near an indoor dog kennel, a washing machine (after a load of bedding was run), and near windows. Eight (30.8%) had been found in beds or bedrooms, of which 6 had been found alive (2 submissions did not indicate whether the bug had been found dead or alive). Three of the bugs found live in bedrooms were reported as having likely bitten a person. An additional 4 bugs had been reported as likely having bitten someone, although they were not found in beds or bedrooms; these bugs were found in a washing machine (after a load of bedding), a front entry way, and 2 were reportedly as generally “indoors”. Two of the 7 samples that had been associated with potential bites had been submitted to TAMU, and dissection revealed blood in the guts (blood meal scores of ‘4’ for both bugs).

Of the 26 TAMU records, 7 specimens were sent to the state health department for routing to the CDC, 9 specimens were submitted to TAMU, and 10 specimens had been discarded by their submitters and were unavailable for further processing. Of the 30 samples processed and tested by TAMU or CDC, 12 (40%) were positive for T. cruzi; these 12 positive specimens included 2 that had been found in a bedroom, 1 of which had been reported as biting a person. The 6 positive samples were from the following counties: Bates, Boone, Howell, Jefferson, Lewis, and Ralls; the other 6 did not have a county recorded. Three of the positive samples that had been processed at TAMU were further processed to determine T. cruzi DTU—1 sample was TcI (adult T. sanguisuga female from Lewis County), 1 was TcIV (adult T. sanguisuga female from Ralls County), and 1 was not successfully typed. Three samples that had been routed to the CDC and had tested positive for T. cruzi had additional processing to determine whether human blood was present in the samples. One of these samples—a T. sanguisuga from Jefferson County, Missouri—had evidence of human blood, and the submitter had mentioned that the bug had been feeding on her. One T. lecticularia sample from Boone County, Missouri had no evidence of human blood, but did have evidence of hog (Sus scrofa) and domestic dog (Canis lupus familiaris) blood; this bug was found in a bedroom.

Records From All Sources

In total, we documented 228 kissing bug records in Illinois and Missouri. Most bugs were found during the summer months (Fig. 3). Among the 215 records with known date of encounter or capture, 124 (57.7%) were found in July or August and 42 (19.5%) were found in June or September (Fig. 3). Four of the specimens were noted as dead upon collection. The specimen collected in February was found dead in a washing machine with bedding, but likely bit a household member prior to washing. Three others were found dead indoors (found in June, October, and December), and 1 was found dead outdoors in a woodpile (September).

Discussion

Although Illinois and Missouri likely harbor the current northernmost limits of triatomines (Bern et al. 2019), few observations exist in the historical literature. The 26 TAMU/CDC specimens collected by community scientists described here represent an important addition to our knowledge of current distributions of T. sanguisuga and T. lecticularia in these states. Though recent work has suggested re-assigning Triatoma lecticularia to Paratriatoma lecticularia (de Paiva et al. 2021), the original genus name was kept throughout as the purpose was to assess historical records and to maintain consistency across search methods. Community scientists played a key role in collaborating with academic and public health institutions to generate these important data. In conjunction with preserved collections and other community-driven platforms such as iNaturalist and BugGuide, we were able to piece together the most comprehensive description of triatomines in both states to date.

In all likelihood kissing bugs are distributed ubiquitously across Missouri, as shown by the scattered pattern in all regions of the state (Fig. 2). The Illinois distribution (Fig. 1) may be at least the entire southern half of the state, if not further north. There was one historical record from the INHS database of T. sanguisuga found in a bed in May, 1968 in Stephenson County (northcentral Illinois) (McElrath 2022). However, no other records have been recorded north of Peoria and McLean counties in Illinois. T. sanguisuga was recently declared as established in southeastern Nebraska (which borders northwestern Missouri), including 1 sample positive for T. cruzi (Nielsen et al. 2021). Immediately to the west of Missouri lies the state of Kansas, where both T. sanguisuga and T. lecticularia have been reported (Kimball 1893, Schmidt 2011). Additionally, T. sanguisuga has been found in Wyoming, situated northwest of Kansas (Reeves and Miller 2020). When considering the historical records in Illinois and Missouri (Bern et al. 2011, Porter 1965), it may be important to keep in mind that prior to 1944, T. lecticularia was described as a subspecies of T. sanguisuga (Bern et al. 2011, Usinger 1944, Ryckman 1984), and so earlier records of T. sanguisuga may include some T. lecticularia records. It is clear that kissing bugs are established in these states, although less frequently reported than kissing bugs in more southern states such as Texas.

The finding of more samples in the summer months than other times of year (Fig. 3) corroborates data from the larger community science dataset from Texas, which found T. sanguisuga activity peaking in August (Curtis-Robles et al. 2018a). The typical nocturnal habits of T. sanguisuga make the evening-morning findings of samples expected, although the findings of a few samples around mid-day were interesting. It may have been that people found the bugs in their daytime resting locations. For example, 1 specimen found recently in Sangamon County, IL (April, 2022) was found in a tree stump along a nature trail.

More than 90% of TAMU samples were found indoors, which is expected for a community science program, but also higher than findings in Texas a few years ago, where 49% (202/414) of T. sanguisuga had been found indoors (Curtis-Robles et al. 2018a). Notably, 8 (31%) specimens were found in bedrooms, indicating triatomines enter homes and have opportunity for human blood meals and potential for T. cruzi transmission in this area. Triatomines in houses, and humans experiencing adverse reactions to bites, were previously reported in Illinois (Porter 1965). In our study, 7 bugs were reportedly associated with bites, including 3 that had been found live in beds or bedrooms and 1 found in a washing machine after a load of bedding had been washed. One bug that had been found feeding on the submitter in bed was positive for both T. cruzi and for human blood. Although nighttime feeding presents an ideal uninterrupted feeding opportunity for kissing bugs, there is evidence that even free-roaming kissing bugs feed on humans outside of homes (Klotz et al. 2014a). It remains important to investigate the distribution of triatomines in Illinois and Missouri, given this evidence of parasite positivity and human interaction. We emphasize that our collaborative academic-based community science initiative includes direct communication with state health departments and the CDC when submitters disclose that insects were found in a bedroom and/or associated with a human bite so that submitters are in contact with human health professionals to guide any medical steps.

T. cruzi infection prevalence in the small set tested at TAMU and CDC was 40% (12/30). T. cruzi has been detected from T. sanguisuga specimens in every state from which specimens have been tested (Bern et al. 2011). Previous examinations of T. sanguisuga from the 1960s in Illinois showed no evidence of T. cruzi (Porter 1965), although it is important to note that current PCR-based methods are much more sensitive. Along with triatomine distribution and T. cruzi infection prevalence, determining T. cruzi prevalence among animal reservoir species is important to also elucidating regions of public health importance (Kruse et al. 2019). In the United States, T. cruzi is frequently detected in canines (Curtis-Robles et al. 2018b, Meyers et al. 2020, Barr et al. 1995). It currently appears that raccoons may be the primary sylvatic reservoir species in Illinois and Missouri (Hamer et al. 2018, Brown et al. 2010), with many other mammalian reservoirs in the United States, particularly woodrats, packrats, raccoons, and wild canine species (John and Hoppe 1986, Klotz et al. 2014b, Gunter et al. 2017, Hodo and Hamer 2017). Recently, of 36 raccoons in Jackson and Williamson counties in southern Illinois, 6 (16.7%) were positive for T. cruzi, suggesting a sylvatic cycle of T. cruzi transmission occurs in the state (Vandermark et al. 2018). In Missouri, T. cruzi antibodies were found in 74 (68%) sampled raccoons (Brown et al. 2010). It is apparent that the T. cruzi transmission cycle is active in the sylvatic environment of this area, although a more thorough understanding of nidicolous habitats could lead to improved vector control efforts.

Autochthonous (locally-acquired) T. cruzi infections are assumed to be rare in the United States, likely owing to a variety of factors such as the sylvatic nature of many triatomine species found in the United States, typically well-sealed housing, inefficient transmission, etc., that could ultimately limit consistent human contact with the vectors and potential for infection with T. cruzi. (Klotz et al. 2014b, Nouvellet et al. 2013, Kribs-Zaleta 2010). Also, inefficient transmission could account for the possible rarity of cases in the United States so far. Nouvellet et al. estimated that the probability of human transmission of T. cruzi from triatomine vectors is probably very small, depending on the vector species, location, and abundance (Nouvellet et al. 2013). However, it is possible that autochthonous cases are occurring without detection, as Chagas disease is not a notifiable disease in most U.S. states, including Illinois and Missouri (Bennett et al. 2018). There have been at least 78 recorded autochthonous cases in the United States (Beatty and Klotz 2020), and among many of these there is no clear link to a triatomine bite or recollection of seeing kissing bugs around the home, suggesting either low density or elusive/cryptic vector populations and/or possible nonvector-borne transmission (Lynn et al. 2020). In fact, the autochthonous case reported in Missouri in 2018 reported no clear link to a bit and little to no recollection of finding kissing bugs (Turabelidze et al. 2020). There were no triatomines positive for T. cruzi within our study sample from the same county as the autochthonous case; however, our results suggest a high likelihood of T. cruzi infected triatomines across the state of Missouri.

There is little understanding of the dynamics of vector-borne transmission in the United States, and routes including vertical transmission, transmission through blood transfusions or organ donations, and oral transmission need additional study (Beatty and Klotz 2020).

In cases of nocturnal bites of unknown origin, cleaning the bedroom can be an important step, as evidenced by the submitter who found a triatomine in the washing machine after laundering sheets from a bed where a person had been bitten. Detecting and destroying kissing bugs can be aided by an intensive cleaning of the room, including laundering bedding, clearing under the bed, carefully checking the mattress, and thoroughly vacuuming the room. In order to keep triatomines from entering homes, screens should be kept in good repair, cracks, and crevices that may serve as entry points or hiding places should be reduced, and outdoor lights should be turned off in evening and night as much as possible to reduce attraction of kissing bugs to residences. Insecticide treatments of domestic or peridomestic areas can be considered if there is evidence of infestation (i.e., finding immatures insect stages, or repeated encounters of adults). However, the sylvatic nature of triatomines found in the United States challenges the potential efficacy of insecticides for suppressing populations. Further, no insecticide is currently specifically labeled for triatomines in the United States, and working with pest control professionals using integrated pest management strategies is advised.

Limitations

Due to the low number of triatomines submitted to TAMU/CDC from Illinois, analyses were not separated by state. The Supplementary data file may be used to separate records by state. One limitation of community-driven submissions is that submissions are only from individuals who sought out the program. Additionally, a complicating factor of using online databases is their “live” nature, such that the numbers may frequently change. To address this, we conducted searches of the online databases on a single day and carefully recorded the search days.

Conclusions

There is a need for a more complete understanding of the distribution of kissing bugs among states along the northern limits of triatomine species in the United States, particularly in Illinois and Missouri. As vectors of T. cruzi, their distribution has implications for public health importance in the United States. Increasing awareness of kissing bugs and the TAMU Community Science Program is one strategy that would allow for passive insect collection to improve our distribution understanding in these states. As illustrated in Fig. 2, community-derived contributions from online observation and data sharing platforms play a unique role in contemporary species documentation. Though care must be taken in compiling reports, and an understanding of data quality for each platform is important to avoid misclassification and duplication, these platforms are useful particularly for species of public health importance where little is known about distribution.

Supplementary Material

Supplementary material is available at Journal of Medical Entomology online.

Acknowledgments

This manuscript would have not been possible without the invaluable contributions of community scientists, including J.M. Balzer, M. Bishop, M.C., R. Cox, C. Dempsey Dotson, R.L. Daniels, K. Freborg, C. Farrell, B. Gravlin, S. Greathouse and J. Longaker, O. Hamer, S. Latham and E. Beck, D. Litherland, K.L. Morris, L. Morris, R. Steinkamp, D.L. Stewart Sr., K. Usher, K. Vinson, and D.K. Wilson. We are deeply grateful for the time and resources each submitter dedicated to submitting their observation(s) and answering questions about their sample(s). Community scientists contributing to online platforms (iNaturalist and BugGuide) played an important role in generating data for this work. We are grateful for their curiosity and documentation. We are also grateful for the support and interest of colleagues in Illinois and Missouri; especially to Dr. Holly Tuten of INHS for introducing ES to experts across the state, Dr. Tommy McElrath for INHS collections data, and Kristin Simpson at the Enns Entomology Museum for providing Collections data.

Funding

Funding from the Texas EcoLab Program supported the development and sustainment of the Texas A&M University Kissing Bug Community Science Program. Graduate student support for RC-R was provided by the National Science Foundation Graduate Research Fellowship Program under Grant No. 1252521. Finally, we would like to acknowledge the Southern Illinois University Edwardsville’s Emeriti Foundation grant that supported visiting scholars, which helped initiate this large collaborative effort. The findings and conclusions in this paper are those of the authors and do not necessarily represent the official position of the CDC.

Author Contributions

Ellen Santos (Conceptualization [Equal], Data curation [Equal], Formal analysis [Equal], Writing – original draft [Lead], Writing – review & editing [Equal]), Catherine Santanello (Conceptualization [Equal], Formal analysis [Equal], Investigation [Equal], Writing – original draft [Equal], Writing – review & editing [Equal]), Rachel Curtis-Robles (Data curation [Equal], Investigation [Equal], Methodology [Equal], Supervision [Equal], Validation [Lead], Writing – review & editing [Lead]), Keswick Killets (Data curation [Lead], Resources [Lead], Validation [Equal], Writing – review & editing [Equal]), Gena Lawrence (Methodology [Equal], Resources [Equal], Validation [Equal], Writing – review & editing [Equal]), Jet Sevenshadows (Data curation [Equal], Resources [Equal], Writing – review & editing [Equal]), Meredith Mahoney (Data curation [Equal], Resources [Equal], Writing – review & editing [Equal]), Molly Baker (Investigation [Equal], Validation [Equal], Writing – review & editing [Equal]), and Sarah Hamer (Conceptualization [Equal], Data curation [Equal], Resources [Equal], Supervision [Equal], Writing – review & editing [Equal])

References