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Review
Unraveling the Importance of Triatomine (Hemiptera:
Reduviidae: Triatominae) Feeding Sources in the Chagas
DiseaseContext
AlbertoAntonio-Campos, RicardoAlejandre-Aguilar, and NancyRivas1,
Laboratory of Medical Entomology, Department of Parasitology, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico
Nacional. Unidad Profesional Lázaro Cárdenas, Prolongación de Carpio y Plan de Ayala s/n, Col. Santo Tómas C.P. 11340 Alcaldía
Miguel Hidalgo, CDMX, Mexico and 1Corresponding author, e-mail: caliope_rivas@yahoo.com.mx
Subject Editor: CarlosBlanco
Received 17 July 2020; Editorial decision 24 October 2020
Abstract
The triatomines are vectors of Trypanosoma cruzi (Chagas, 1909), the etiologic agent of Chagas disease. All
species are strictly hematophagous, and the hosts used by vector species are important to understand the
transmission dynamics of T. cruzi, and eventually, for the development of effective control strategies in en-
demic countries. In the current review, we gather a comprehensively number of literature reporting triatomine
feeding sources, using rigorous targeted search of scientific publications, which includes research papers and
reviews to put together the most recent findings of the feeding behavior in triatomines and their applications
for vector control of Chagas disease. Our main findings suggest that the main feeding source in triatomines is
the human blood (22.75%), T.dimidiata (Latreille, 1811) (Hemiptera: Reduviidae) is the most frequent (13.68%)
triatomine species in this type of study, and most of the studies on feeding sources (47.5%) are conducted in
the domestic and peri-domestic environment.
Key words: Chagas disease vector, feeding source, behavior, Chagas disease, host choice
Chagas disease, also known as American trypanosomiasis, is caused
by the parasite Trypanosoma cruzi (Chagas, 1909)(Kinetoplastida:
Trypanosomatidae). This disease affects about 6 million to 7 million
people around the world, mostly in Mexico, Central, South America
and it has recently been reported in the southern part of the United
States (WHO 2019). To date, there is no vaccine and treatments are
ineffective and with severe side effects (Rassi etal. 2010). According
to paleo-parasitology ndings, this disease already existed 9,000 yr
ago in the Americas, and today, it continues to be a relevant public
health problem, mainly in endemic countries (Aufderheide et al.
2004, Mathers etal. 2007, Moncayo and Silveira 2009).
This disease is caused by the hemophlagelated intracellular para-
site T. cruzi, which belongs to the Trypanosomatidae family. This
parasite can infect any type of nucleated cell, mainly macrophages,
broblasts, and epithelial cells. During its life cycle, the parasite trans-
forms into four well-dened stages: metacyclic trypomastigote (in-
fective form present in the feces of the triatomine vector), amastigote
(replicative form in the mammalian host), blood trypomastigote
(infective form in the mammalian host) and, epimastigote (replica-
tive form in the triatomine vector) (Teixeira etal. 2012, Lidani etal.
2019). Trypanosoma cruzi is transmitted in 80% of cases, through
contaminated feces of triatomine insects belonging to the Hemiptera
order, Heteroptera suborder, Reduviidae family and Triatominae
subfamily, secondly, infection by transfusion with units of blood in-
fected with the parasite (10–20%); thirdly, congenital transmission
(5%), and to a lesser extent, infections due to ingestion of contam-
inated food (~1%) (Rassi etal. 2010). Most of the time triatomines
bite on an exposed area of the skin and tend to defecate near this site.
The parasite penetrates the host when it scratches as a result of the
pruritic effect of triatomine saliva and disperses the feces to the area
of sting, eyes, mouth or any exposed skin lesion (Rassi etal. 2010,
Teixeira etal. 2012).
A total of 154 species of triatomines, representing 19 genera,
have been described around the world (Justi and Galvão 2017,
Oliveira and Alevi 2017, Dorn etal. 2018, Oliveira etal. 2018, Lima-
Cordón et al. 2019, Nascimento et al. 2019, Poinar 2019). Some
endemic countries concentrate a wide variety of triatomine species.
In Mexico, the presence of 32 triatomine species has been reported:
Triatoma Laporte, 1832 (19), Meccus Stål, 1859 (6), Panstrongylus
Berg, 1879 (2), Belminus Stål, 1859 (1), Dipetalogaster Usinger,
1939 (1), Eratyrus Stål, 1859 (1), Paratriatoma Barber, 1938 (1) and
Rhodnius Stål, 1859 (1) (Cruz-Reyes and Pickering-López 2006,
Ramsey etal. 2015). Twenty-one of these insects have been reported
naturally infected with T. cruzi. It is also known that the genera
Annals of the Entomological Society of America, 114(1), 2021, 48–58
doi: 10.1093/aesa/saaa045
Advance Access Publication Date: 4 December 2020
Review
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49Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
Dipetalogaster, Meccus and eight species of the genus Triatoma
are endemic from Mexico. Furthermore, T. dimidiata (Latreille,
1811)(Antonio-Campos et al. 2019) and T. barberi Usinger, 1939
(Salazar-Schettino etal. 2010, Rivas et al. 2018) are considered of
medical importance since they inhabit human dwellings (Cruz-Reyes
and Pickering-López 2006, Ramsey etal. 2015). In Colombia, there
are 26 listed triatomine species, of which 15 have been reported as
naturally infected with T.cruzi (Guhl etal. 2007). Even developed
countries like the United States have been affected by insect trans-
mission of Chagas disease. In this country, T.cruzi was rst iden-
tied in a T.protracta (Uhler, 1894) bug collected in California in
1916. Furthermore, T.protracta (Uhler, 1894) was reported in this
country as a human pest in California in the 1860s, due to allergic
reactions associated with triatomine bites. Since these early reports,
11 species of triatomines have been documented in the United States
and all of them, except one (T.incrassata Usinger, 1939)have had
demonstrated infection with T.cruzi (Montgomery etal. 2016).
Feeding behavior of triatomines has been little studied compared
to other vectors of medical importance such as ticks and mosquitoes
(Rabinovich etal. 2011, Otálora-Luna etal. 2015). For triatomines,
blood is their only source of energy and nutrients, and unlike
other hematophagous insects such as mosquitoes, for which, their
larvae and males usually feed on other feeding sources, triatomines
are strictly hematophagous throughout their life cycle (Lent and
Wygodzinsky 1979). Therefore, elucidate triatomine’s feeding habits
is very useful to estimate the amplitude of their niche and therefore
its vector dynamics. Feeding sources of triatomines have been studied
mostly in domestic and peri-domestic habitats, and to a lesser extent
in the wild environment (sylvatic) (Rabinovich etal. 2011). In this
sense, little is known about the importance that the type of habitat
can play in the choice of triatomine feeding source, since the ‘choice’
could be limited by the availability of hosts, as previously shown
in other insects such as mosquitoes (Lefèvre et al. 2009). In add-
ition, the distribution of triatomine species could be overrepresented;
this means that the species that inhabit a certain habitat could be a
subset of species from another habitat that migrated due to food
availability (Rabinovich etal. 2011). Hence, knowledge of the habits
and feeding sources of triatomines is key for understanding the eco-
epidemiology of T.cruzi and the interactions of these insects with its
mammalianhosts.
Insect vector host choice is recognized as an important compo-
nent inuencing the dynamics of vector-borne disease transmission
and has shown effects for the efciency in vector control strategies
(Lazzari and Lorenzo 2009, Lazzari etal. 2013, Lima-Cordón etal.
2018, Zahid and Kribs 2020).
Triatomine Feeding Behavior
Hematophagy has been a milestone in the natural history of
triatomines, since, due to this need, it was that the triatomines got
involved into the life cycle of T.cruzi and later with the humans.
The exact mechanisms by which triatomines became hematopha-
gous remain unknown. However, it is proposed that the genetic
traits inherited by their non-hematophagous ancestors and the dif-
ferent evolutionary mechanisms caused by their physical environ-
ment were the architects of their feeding adaptations (Otálora-Luna
etal. 2015).
Triatomine insects depend on a wide range of sensory signals to
nd hosts. These consist of CO2 gradients, odors, humidity, and heat
(Lazzari and Lorenzo 2009, Lazzari etal. 2013). In general, these
insects carry out their activities mainly at night, and during the day
they remain hidden in their dens, although sometimes, they may go
out to feed during the day under food stress (Lazzari and Lorenzo
2009). An interesting aspect to consider is that in colonies reared in
laboratory, the triatomines change their habits and tend to feed both
at day and night (Lazzari etal. 2013). According to the triatomine
species, the amount of ingested blood varies, although generally, the
4th and 5th stages are the most voracious; however, it is possible that
different factors modulate this behavior (Jurberg and Galvão 2006).
Heat is one of the most used sensory signals by triatomines, since
in addition to its main role as an orientation signal; heat is used to
detect the host’s blood vessels (Ferreira etal. 2007, Lazzari et al.
2013). By analyzing the feeding behavior of bugs on a live host under
laboratory conditions, it has been demonstrated that triatomines do
not bite randomly. Instead, they extend their proboscises directly to-
wards host blood vessels according to different sensory cues (Ferreira
etal. 2007). In laboratory experiments, it has been shown that when
the host skin is replaced by a warm surface, triatomines can detect
the surface with great precision since it exhibits the temperature that
they consider more attractive, indicating that these insects can use
these variations in the skin of its hosts to determine the exact site of
its bite (Ferreira etal. 2007).
Likewise, in studies conducted in laboratory-raised chickens
chopped by triatomines or articially injected with triatomine saliva,
it was noted that when these chickens were used as a food source,
triatomines preferred these groups inoculated with saliva than
chickens that had not been previously bitten or articially inoculated
with triatomine saliva. This suggests the possibility that animals
previously bitten by triatomines would be more likely to continue
being bitten, and therefore increase the risk of infection with T.cruzi
(Hecht etal. 2006).
Tools for Detecting Triatomine FeedingSources
Throughout the evolution of the study of the eco-epidemiology of
Chagas disease and triatomine vectors, different approaches have
been used to study their feeding sources (Fig.1). Each of these ap-
proaches has advantages and disadvantages; however, each of these
provides valuable information that can be used to determine dif-
ferent aspects of the ecology of triatomines, as well as the transmis-
sion dynamics of T.cruzi.
The techniques used so far for feeding source determination are
mainly divided in two: immunological and molecular methods. The
immunological techniques include precipitins (Freitas et al. 1960),
immunodiffusion (Quintal and Polanco 1977), complement x-
ation (Staak etal. 1981), and ELISA (Villalobos etal. 2011). These
techniques have the advantage that ‘specialized’ equipment is not
required to be functional; however, its most notable disadvantage
is that it is unlikely to obtain serum from wild animals. Likewise,
its detection capacity is limited to the different serum samples that
are available. For example, if we had a sample with different blood
sources and there are no sera against all of them, valuable informa-
tion would be lost, and incomplete conclusions could be drawn. In
this sense, these techniques are partially limited for the study of the
sylvaticcycle.
Recently, the use of molecular techniques based on DNA analysis
have been used to determine triatomine feeding sources, such as:
polymerase chain reaction (PCR) with specie-specic oligonucleo-
tides (Pizarro etal. 2007, Pizarro and Stevens 2008), multiplex PCR
(Mota etal. 2007), partial sequencing of cytochrome oxidase I(COI)
and cytochrome b genes (Cyt-b) (Townzen etal. 2008), heteroduplex
of Cyt-b assay (Lee etal. 2002, Bosseno etal. 2006, Torres-Montero
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50 Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
etal. 2012), high-resolution melting (HRM) of the Cyt-b gene (Peña
etal. 2012, Hernández et al. 2016), PCR-RFLP of 16S mitochon-
drial rDNA (Roellig etal. 2013), PCR-RFLP of Cyt-b (Chena etal.
2014, Fraenkel etal. 2020, Sánchez etal. 2020), direct sequencing of
Cyt-b gene and BLAST (Valença-Barbosa etal. 2015, Almeida etal.
2016, Lilioso etal. 2020), amplication by qPCR a fragment of the
mitochondrial 12S ribosomal gene (Ibañez-Cervantes et al. 2013),
amplication of 12S rRNA locus and BLAST (Gottdenker et al.
2012, Stevens etal. 2012, Klotz etal. 2014, Gorchakov etal. 2016,
Bezerra etal. 2018, Velásquez-Ortiz etal. 2019), protein mass spec-
trometry using liquid chromatography tandem mass spectrometry
(LC-MS/MS) (Keller etal. 2017), RAD seq-based analysis (Orantes
et al. 2018) and metabarcoding approach (12S rRNA) based on
next-generation sequencing (Dumonteil et al. 2018, Arias-Giraldo
etal. 2020).
Moreover, the molecular techniques can be subdivided into three
main groups: the rst group covers all those techniques based on
PCR using species-specic oligonucleotides. These techniques have
the same disadvantage as immunological, since, when species-
specic oligonucleotides are used, only the feeding sources for which
these oligonucleotides have been designed can be detected (Pizarro
and Stevens 2008). Therefore, the study of the triatomine sylvatic
cycle feeding sources is partially limited.
The second group techniques are those in which a PCR is per-
formed plus an extra step, such as PCR-RFLP (Roellig etal. 2013,
Chena etal. 2014, Fraenkel et al. 2020, Sánchez et al. 2020), and
some of these methods have the quality of being quantitative like the
qPCR (Peña etal. 2012, Ibañez-Cervantes etal. 2013).
The third group are all those in which highly specialized equip-
ment is needed, such as mass spectrometers and next-generation se-
quencers. The main disadvantage of these techniques is their high
cost and specialized equipment. However, the amount of information
that can be obtained from them is massive. In this way, it is possible
to determine all the components that are contained in the intestinal
contents of the triatomines, for example, DNA from T.cruzi, other
parasites, bacteria, viruses, etc. Therefore, it is possible to obtain
information on the different epidemiological cycles of triatomines,
mixed populations of T.cruzi within the same triatomine, as well
as the intraspecic interactions that occur between T.cruzi and the
triatomine microbiota (Dumonteil etal. 2018, Arias-Giraldo etal.
2020).
Host-Parasite Relationship and
FeedingSources
In the American continent, there is a great variety of triatomine
species; some of them stand out for their vector capacity like
R.prolixus Stål, 1859, T.infestans (Klug, 1834) and T. dimidiata;
therefore, vector control strategies are directed against this type of
species (Rassi etal. 2012). Recently, some species, such as T.barberi,
T. rubida (Uhler, 1894), E.mucronatus Stål, 1859, P. geniculatus
(Latreille, 1811), R.pallescens Barber, 1932, have begun to be con-
sidered relevant in the transmission cycle of T.cruzi because to its
ease of adaptation to domestic environment (Salazar-Schettino etal.
2010, Rabinovich etal. 2011).
Some processes such as urbanization, deforestation, and
defaunation produced by anthropogenic process, directly im-
pact ecology interactions that disturbs triatomine natural habitat.
Therefore, if the aforementioned events propitiate the migration and
extinction of hosts that the triatomines use as their main feeding
source, this spill would force the triatomines to look for other hosts
or even other habitats, otherwise, they would perish (Flores-Ferrer
et al. 2018). Furthermore, these human-caused activities generate
a greater interaction between the different components of the epi-
demiological cycle of Chagas disease: triatomines, the different
strains of T.cruzi, as well as their hosts (Lewis etal. 2011, Ramsey
etal. 2012, Oliveira etal. 2018a)(Fig.2).
Fig. 1. Most used techniques for the detection of feeding sources in triatomines. At the top, the molecular techniques are shown and at the bottom the
immunological techniques.
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51Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
Mixed infections (coinfections) with T.cruzi in a single host ra-
ther than the exception are the rule in nature, both in the mamma-
lian and triatomine hosts (Perez etal. 2014), this event are the result
of sequential feeding process of the triatomines on different hosts
infected with different T.cruzi genotypes. Likewise, it is during this
process that the triatomino defecate a mixture of different genotypes
of the parasite and the epidemiological cycle of T.cruzi continues
(Bosseno et al. 1996). In this sense, it has been reported that the
intraspecic interactions of T.cruzi and its different discrete typing
units (DTU’s) generate natural adaptation to selection pressures
(Lewis etal. 2011, Zingales etal. 2012). These features have been
described as elements that affect the virulence, ecology, and evolu-
tion of the parasite (Gower and Webster 2005, Araújo etal. 2014).
Regarding the direct associations of feeding sources of triatomines
and T.cruzi, it is known that most triatomines are eclectic in their
feeding habits; however, according to what is reported, it is suspected
that the different lineages of T.cruzi are conned or restricted to the
environment to which their reservoirs belong (Zingales etal. 2012,
Flores-Ferrer etal. 2018). However, this connement is being broken
with the introgression of humans’ activities into sylvatic environ-
ment. This is how triatomines move and colonize new habitats by ac-
tive and passive means, where they will nd new feeding sources and
establish new parasite-vector-host associations (Klotz etal. 2014, de
Fuentes-Vicente etal. 2018).
It has been postulated that when triatomines select a host they
base their choice on its abundance. However, under ideal circum-
stances, it is possible for a triatomine to have feeding preference for
one host over another. Therefore, it is possible that these traits dic-
tate adaptive behavior of the triatomine to domestic or wild habitats
(Rabinovich etal. 2011, Martínez-Ibarra etal. 2019). These behav-
ioral traits may be leading by evolutionary process and consequently
have a genetic heritage like mosquitoes (Takken and Verhulst 2013).
The genetic factors of the natural host choice can be studied in la-
boratory setups by crossbreeding and selection experiments (Hecht
et al. 2006, Martínez-Hernandez et al. 2010, Díaz-Albiter et al.
2016, Martínez-Ibarra etal. 2019).
In this review, feeding sources were analyzed in 57 pub-
lished studies (Quintal and Polanco 1977, Christensen et al.
1981, Wisnivesky-Colli et al. 1982, Wisnivesky-Colli et al. 1987,
Christensen etal. 1988, Marcondes etal. 1991, Salvatella etal. 1994,
Costa etal. 1998, Gonçalves etal. 2000, Calderón-Arguedas etal.
2001, Sasaki etal. 2003, Brenière etal. 2004, Sandoval etal. 2004,
Freitas etal. 2005, Bosseno etal. 2006, Caranha etal. 2006, Farfán
etal. 2007, Mota et al. 2007, Pineda etal. 2008, Pinto et al. 2008,
Pizarro and Stevens, 2008, Bosseno etal. 2009, Sandoval etal. 2010,
Sarquis et al. 2010, Villela et al. 2010, Farfán-García etal. 2011,
Souza et al. 2011, Villalobos et al. 2011, Gottdenker et al. 2012,
Peña et al. 2012, Ramsey et al. 2012, Stevens et al. 2012, Torres-
Montero etal. 2012, Buitrago et al. 2013, Ibañez-Cervantes etal.
2013, Kjos et al. 2013, Monteon etal. 2013, Roellig et al. 2013,
Gürtler etal. 2014, Klotz etal. 2014, Nattero etal. 2015, Valença-
Barbosa etal. 2015, Almeida etal 2016, Buitrago etal. 2016, Cecere
etal. 2016, Chacón etal. 2016, Gorchakov etal. 2016, Hernández
etal. 2016, Keller etal. 2017, Bezerra etal. 2018, Dumonteil etal.
2018, Minuzzi-Souza et al. 2018, Orantes et al. 2018, Velásquez-
Ortiz etal. 2019, Fraenkel et al. 2020, Lilioso etal. 2020, Sánchez
etal. 2020), and the main blood meals in all of them were analyzed
according to, the main host, triatomine specie, as well as the habitat
where the specimens were collected: domestic (Dom), peri-domestic
(Per), sylvatic(Syl).
Interestingly, for species with a variable degree of anthropophily
there is reported that biological factors are more important, such as,
vicinity of triatomines to feeding sources, abundance of host spe-
cies, and type of environment (Dom, Per or Syl) in which the study
was conducted (Rabinovich etal. 2011). The studies revised show a
high presence of blood meals from human hosts (Fig.3). However,
this observation does not mean that triatomines prefer human
blood over other species, but as previously mentioned, humans
Fig. 2. Epidemiological cycle of Trypanosoma cruzi. Main interactions between the different habitats: domestic, peri-domestic, and sylvatic.
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52 Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
increasingly invade the native habitat of these insects, which in-
creases the risk of contact (Rabinovich etal. 2011, Minuzzi-Souza
etal. 2018).
Curious, one of the studies noted that, when mouse, dog, and
chicken blood meals were identied in T.dimidiata, human blood
meals were likewise found (Monteon etal. 2013). This observation
suggests that the triatomines can disperse around domestic and peri-
domestic environment to search better host availability and accessi-
bility (Gürtler etal. 2009).
Birds were the second most frequently bloodmeal reported after
human (Fig.3). This data suggests that either in the peri-domestic
environment, but mainly in the wild, birds are in close contact with
triatomines, mainly those who live in palms and other types of trees
(Velásquez-Ortiz etal. 2019). Moreover, two of the most studied
ecological associations in the domestic cycle of Chagas disease is the
presence of chickens and dogs. In this regard, it has previously been
described that birds, including chickens, are immune to infection
with T. cruzi (Kierszenbaum et al. 1976, 1981; Minter-Goedbloed
and Croon 1981). Based on this fact, it would be logical to suppose
that those households that had chickens around them would be pro-
tected from triatomines and T. cruzi infection, but that assumption
seems to be incorrect considering recent studies (Dumonteil et al.
2013, Gürtler etal. 2014, Gürtler and Yadon 2015, Koyoc-Cardeña
etal. 2015, Zahid and Kribs 2020).
Although it has been proposed that biodiversity reduces the risk
of transmission of vector-borne diseases, this statement depends on
many factors and one of the most signicant is the competency of the
new hosts (LoGiudice etal. 2003, 2008; Saul 2003). In accordance
with the competency of new hosts added in the current habitat, the
effect of distraction of vectors from their tting host (e.g., the hu-
mans) can be divided in two types: alternative or incompetent hosts’
effect and decoy effect. Alternative hosts are capable of transmitting
pathogens, but not as signicantly as the main host (e.g., the dogs),
whereas decoy effect involves the presence of any incompetent host
(incapable of transmitting the pathogen), e.g., the chicken in the
Chagas disease context (Zahid and Kribs 2020).
In other models of vector-borne diseases, the use of decoys (e.g.,
livestock) to distract mosquitoes far away from human dwellings
may decrease infections in short-term, but the rise in successful
blood meals has the potential to cause long-term increases in mos-
quito populations and paradoxically boost the possibility of infec-
tion to humans (Saul 2003, Dobson etal. 2006).
Hence, a possible conclusion in this issue is that domestic animals
and chicken coops in domestic areas affect the host feeding choices,
human vector contacts rates and parasite transmission; however, the
presence of these animals should be considered as zoopotentiation
rather than zooprophylaxis (Dumonteil et al. 2013, Gürtler et al.
2014, Koyoc-Cardeña etal. 2015).
Mixed blood meals were detected third in frequency (Fig.3); they
mainly belong to animals that usually cohabit in the same niche,
suggesting the eclectic behavior of triatomines. Rodent blood meals
were detected fourth (Fig.3), and although the mouse model is one
of the most widely used in the world to assess infection with T.cruzi,
little is known about how this animal naturally interacts with
T. cruzi and humans. However, rodents are known to have great
synanthropic capacity, making them ideally suited for transmission
of T.cruzi through the different habitats that make up the epidemio-
logical cycle of Chagas disease (Ibarra-Cerdeña etal. 2017).
Dog blood meals were detected fth (Fig.3), and it is noteworthy,
since they have been considered hosts of medical importance for
T. cruzi, because they are in close contact with humans and peri-
domestic environment. Recent studies indicate that in environments
endemic for Chagas disease, the older a dog is, the more likely it is to
be infected with T.cruzi (Curtis-Robles etal. 2017).
On the other hand, atypical blood meals were reported such as
reptiles (Lilioso etal. 2020), amphibians (Gottdenker et al. 2012),
and insects (Sandoval etal. 2010). In part, these feeding sources in-
dicate what has been previously described, since during nutritional
stress triatomines usually feed on any suitable host, including fruits
(Díaz-Albiter etal. 2016).
Regarding the triatomine species most studied in terms of feeding
sources, the most important would be T.dimidiata, T.infestans, and
T.brasiliensis (Neiva, 1911), respectively (Fig.4). These triatomine
species are mainly South American, except for T.dimidiata, which
ranges from the southern part of Mexico to Colombia (Guhl etal.
2007, Flores-Ferrer etal. 2018). This information as well as the
country of origin of the research (Supp Table S1 [online only]) of
these reports indicates that the investigations are mainly generated
by South American countries (Rabinovich etal. 2011, Buitrago etal.
2016, Bezerra etal. 2018). Therefore, the information obtained is
biased towards a subset of triatomine species.
The results reported in each of the research works in this re-
view, highlight the different associations between triatomines and
their different hosts. However, if this information is intended to be
Fig. 3. Feeding sources. Percentage of different kind of feeding sources reported in scientific publications.
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53Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
used to design ecological networks, as well as vector control strat-
egies, it is necessary to consider that the data are biased. Comparable
studies have not been conducted between the different geographic
regions, and different epidemiological habitats, because studies in
the domestic and peri-domestic environment predominate over a few
in the sylvatic environment (Fig.5).
Discussion
This review illustrates that some triatomine species display an adapt-
able behavior for the selection of their blood meals, but also that,
some triatomines are host specic, mainly in the sylvatic environ-
ment. The feeding sources of most triatomines are frequently deter-
mined by the host species that are most abundant or easily available.
An example of this is the variable host preference of T. dimidiata,
which varies from high degree of anthropophily (Torres-Montero
etal. 2012), with several blood meals taken on humans, to a high
degree of blood meals taken from another mammals, birds and even
reptiles and other insects (Durán etal. 2016, Dumonteil etal. 2018,
Minuzzi-Souza etal. 2018).
The high degree of plasticity in host preference of T.dimidiata
(Torres-Montero et al. 2012), T. brasiliensis Neiva, 1911 (Bezerra
etal. 2018), T.infestans (Nattero etal. 2015), M.pallidipennis (Stål,
1872)(Ramsey etal. 2012) and P.arthuri (Pinto, 1926)(Velásquez-
Ortiz etal. 2019); indicates that, although there may be particular
preferences for a host, these triatomine species are adapted to feed
under different circumstances, therefore, the most abundant host is
more likely to be itsprey.
On the other hand, T. barberi has a mainly anthropophilic be-
havior, probably due to its intra-domestic habits and its frequent
contact with humans. Although changes in this behavior are de-
scribed, has been probably under conditions of food stress (Zárate
etal. 1980, Salazar-Schettino etal. 2010, de Fuentes-Vicente etal.
2018).
Fig. 5. Predominant triatomine habitat reported in feeding sources studies.
Fig. 4. Major triatomine species reported in feeding sources studies.
Downloaded from https://academic.oup.com/aesa/article/114/1/48/6020064 by guest on 16 January 2021
54 Annals of the Entomological Society of America, 2021, Vol. 114, No. 1
Most of the studies in which high levels of anthropophilia are
reported are carried out in the intra-domestic environment, where
the probability that a triatomine feeds on humans is greater.
Consequently, there is an inherent bias when feeding sources are
limited, because triatomines will feed on any available host in their
quest for survival. If what is intended is to analyze host preferences
and not only feeding sources, then it will be necessary to perform
analyzes with live hosts (different from each other), hosts previously
stung by triatomines (Hecht etal. 2006), or host-derived odor sam-
ples as in the case of mosquito tests (Pates etal. 2001) and even carry
out prospective studies in thewild.
The possible association of anthropophilic behavior of the trans-
mitting insects of Chagas disease requires the use of new approaches
like behavioral ecology and the new discipline of Eco-Evo-Devo.
Anthropophilia is very likely an evolutionary trait exploited by
some pathogens that require humans to complete their life cycle as
Plasmodium falciparum and dengue virus. Therefore, it is possible
that triatomine host choice is an evolutionary adaptation caused by
parasite–host interactions, and that such adaptation is benecial to
the parasite (Koella and Boëte 2003, Cohuet etal. 2010).
Issues discussed in this review show that, unlike other vector-
borne diseases, there is little information on the feeding habits of
triatomines. However, these studies are important because they can
help us to understand how triatomines select some host over others
(Lazzari and Lorenzo 2009). Above all, the underlying mechanisms
and the parasite–host interactions, that govern host choice in both
the triatomine and T.cruzi (Lazzari and Lorenzo 2009, Lazzari etal.
2013). Moreover, a better understanding of feeding sources, as well
as the choice of the host, is necessary to understand the factors as
well as the relationships that rule the current eco-epidemiology of
Chagas disease and last but not least important, for the design of
effective strategies for the control of triatomines in endemic areas
(Lazzari and Lorenzo 2009, Lazzari etal. 2013, Lima-Cordón etal.
2018).
One of the most widely used approaches to vector control is
Eco-health (Waleckx etal. 2015, Lima-Cordón etal. 2018). This
approach focuses on households’ improvements like wall plas-
tering and cement ooring, backyard cleaning, as well as insecti-
cide spraying. Considering the information regarding the feeding
habits, main hosts, and habitat of the triatomines, it would be
possible to focus the efforts of this type of approach. In add-
ition, to reduce the environmental impact, spraying insecticides
exclusively in the areas where triatomines interact with humans,
for example in specic areas of the domestic inhabitant. (Lima-
Cordón etal. 2018).
Conclusion
According to the scientic literature analyzed, the triatomine feeding
sources have been studied using different approaches, ranging from
the oldest such as precipitins to the more modern such as next-
generation sequencing (metabarcoding). Using these and other tools,
it has been possible to elucidate some of the relationships between
T.cruzi, triatomines, and their hosts. In addition, it is shown that the
main feeding source of triatomines is the human blood, and studies
on the feeding sources of triatomines cover a wide range of species,
of which the most recurrent is T.dimidiata.
From the eco-epidemiology point of view, this review highlights
the importance of the feeding behavior of triatomines, the inter-
actions between these insect vectors of T. cruzi and their hosts, as
well as the application of this knowledge to design effective strat-
egies for the vector control of Chagas disease.
SupplementaryData
Supplementary data are available at Annals of the Entomological Society of
Americaonline.
Table S1. Database of the 57 references analyzed.
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