DAILY SURVIVAL RATES AND DISPERSAL OF AEDES AEGYPTI FEMALES IN RIO
DE JANEIRO, BRAZIL
RAFAEL MACIEL-DE-FREITAS,* CLAUDIA TORRES CODEÇO, AND RICARDO LOURENÇO-DE-OLIVEIRA
Laboratório de Transmissores de Hematozoários, Departamento de Entomologia, Instituto Oswaldo Cruz, Fiocruz, Rio de Janeiro,
Brazil; Programa de Computação Científica, Fiocruz, Rio de Janiero, Brazil
of Aedes aegypti. These parameters were estimated for mosquito populations from a slum and a suburban district in Rio
de Janeiro, during the wet and dry seasons in 2005. In each mark-release-recapture experiment, three cohorts of
dust-marked Ae. aegypti females were released. Recaptures were carried out daily in randomly selected houses, using
backpack aspirators, adult traps, and sticky ovitraps. Recapture varied between 6.81% and 14.26%. Daily survival was
estimated by fitting two alternative models: exponential and nonlinear models with correction for the removal of
individuals. Slum area presented higher survival and parity rates (68.5%). Dispersal rates were higher in the suburban
area, where a maximum dispersal of 363 m was observed. Results suggest intense risk of dengue epidemic, particularly
in the urban area.
Daily survival rates, life expectancy, dispersal, and parity are important components of vectorial capacity
A precise estimation of life history parameters of Aedes
aegypti is essential for the development of dengue transmis-
sion models.1Among these parameters, the daily survival
probability of adult females is one of the most important,
because small increases in survival may exponentially in-
crease the vectorial capacity of mosquitoes.2As a rule, vec-
tors must survive longer than the sum of the initial non-
feeding period plus the extrinsic incubation period to be able
to infect another human. For dengue transmission, that means
a lifespan of least 12 days (2 days of nonfeeding and 10 days
Besides survival, another important component of vectorial
potential is dispersal. A very mobile infected vector has
greater chance of finding susceptible humans than one with
low dispersal. Ae. aegypti is not a very mobile species, gener-
ally flying 50–300 m during its lifetime5. Reports of longer
distance flights exist and suggest that Ae. aegypti is capable of
covering considerable distances in few days if necessary.6,7
Variation in mosquito displacement may be explained by het-
erogeneity in the availability of breeding sites and blood op-
In Rio de Janeiro city, dengue fever has become endemic
since its reintroduction in the 1980s. Ae. aegypti is specially
abundant in urbanized and densely populated neighbor-
hoods.11–13To provide parameters for the development of
models of dengue transmission in Rio de Janeiro, we used a
mark-release-recapture (MRR) study design to evaluate the
variation in the probability of daily survival, average life ex-
pectancy, and dispersal of Ae. aegypti females in two areas of
contrasting urbanization patterns in Rio de Janeiro during the
dry and wet seasons. We also evaluated the ovarian develop-
ment and parity rates of natural populations of Ae. aegypti
living in the above conditions. The ultimate goal of this study
is to increase the understanding on dengue transmission in
Rio de Janeiro.
MATERIALS AND METHODS
Study area. MRR studies were conducted in two neighbor-
hoods at Rio de Janeiro city—Favela do Amorim and Tubia-
canga—characterized by contrasting urbanization patterns
and potential isolation from the surroundings, which is ex-
pected to minimize mosquito losses by emigration during
Favela do Amorim (22°52?30? S; 43°14?53? W) was chosen
to represent a densely populated (901.2 hab/ha) low-income
urban area, characterized by disordered human occupation
and scarce vegetation coverage (a typical Brazilian slum).
There, 2,992 people live in 897 substandard houses, in an
estimated area of 2.32 ha. Houses are very small—rarely with
more than one room—and without yards or any kind of peri-
domestic area. The neighborhood is surrounded by large
highways and the Oswaldo Cruz Foundation campus, which is
a largely vegetated, non-residential area, which probably does
not encourage Ae. aegypti females to emigrate from the slum.
Tubiacanga (22°47?08? S; 43°13?36? W) was chosen to rep-
resent a planned, suburban area. There, 2,902 residents live in
867 houses, which resulted in a human density of 337.4 hab/ha
(an estimated area of 8.6 ha). This neighborhood is located in
a lowland coastal area, partially surrounded by the Guana-
bara Bay shores and a 3-m wall of the Tom Jobim Interna-
tional Airport of Rio de Janeiro and its numerous landing
strips. There is a single way to get into Tubiacanga by car: a
2.1-km paved road, which connects the area to the nearest
neighborhood. Thus, mosquito emigration is not expected to
happen in large scale. The area has extensive unpaved streets
and moderate vegetation coverage. Most houses have a large
peridomestic environment and at least two bedrooms.
Climate and MRR periodicity. Climate in Rio de Janeiro is
characterized by a dry winter (May–August) and a wet sum-
mer season (November–March).14During the 1930–1990 pe-
riod, dry and wet seasons in Rio de Janeiro had mean tem-
peratures of 25.1°C and 28.8°C and mean total rainfall of 46.4
and 132 mm, respectively. MRR experiments were performed
during both seasons in each area: Tubiacanga in February
2005 (wet) and July 2005 (dry) and Favela do Amorim in June
2005 (dry) and December 2005 (wet season). Air temperature
and precipitation data for these months were obtained from a
meteorological station located 5 km away from the two study
* Address correspondence to Rafael Maciel-de-Freitas, Laborato ´rio
de Transmissores de Hematozoa ´rios, Pavilhao ´ Carlos Chagas, 4° an-
dar sala 04, Departamento de Entomologia, Rio de Janeiro, Brazil
21040-360. E-mail: email@example.com
Am. J. Trop. Med. Hyg., 76(4), 2007, pp. 659–665
Copyright © 2007 by The American Society of Tropical Medicine and Hygiene
Mosquitoes. Aedes aegypti used in MRR experiments came
from a laboratory colony that is constantly renewed with eggs
collected in Rio de Janeiro. Larvae were fed with fish food
(Tetramin, Tetra Sales, Blacksburg, VA) and reared accord-
ing to Consoli and Lourenço-de-Oliveira.15After emergence,
females were kept together at 25 ± 3°C and 65 ± 5% relative
humidity (RH) and fed with sucrose solution until the time to
Marking and releasing. Before each experiment, eggs were
split into three groups and allowed to hatch on 3 consecutive
days, producing three female adult cohorts. Each cohort was
marked with a different color of fluorescent dust (Day-Glo
Color Corp., Cleveland, OH) in small cylindrical cups (12 × 10
cm). Mated and unfed females were released outdoor at each
area in the morning of their fourth day after emergence (be-
tween 8:00 AM and 9:00 AM), ∼1 hour after dust marking.
One mosquito cohort was released each day for 3 consecutive
days in each area, both in dry and in wet seasons, totalizing 12
field experiments. Each cohort was released at different
points of the study areas. Around 4–5 months elapsed be-
tween releases of the cohorts in each area.
Capturing. Dust-marked females were captured with CDC
backpack aspirators (John W. Hock, Gainesville, FL), BG-
Sentinels adult trap (BioSantos GmbH, Rejemsburg, Ger-
many), and sticky ovitraps.16–19Captures started 1 day after
the release of the first cohort. Fifteen houses were randomly
selected per day for aspiration, which was done in 15–20 min-
utes per house. The whole house was aspirated, including the
peridomestic area. BG-Sentinel traps (BGS-Trap) were in-
stalled in 15 houses, and remained there during the whole
extension of the MRR study, being daily monitored for the
presence of dust-marked females. Occasionally, aspiration
was done in the same house where a BGS-Trap was installed.
Results concerning species specificity, capture efficiency, and
a more suitable description of BGS-Trap can be found else-
where.17,18Finally, 20 sticky ovitraps were placed at the pe-
rimeter of the study areas, as an attempt to capture emigrat-
ing insects from study areas. Sticky ovitraps were placed on
the ground, in the peridomestic environment, 30–40 m apart
from each other in Tubiacanga and 50–60 m apart in Favela
do Amorim. Because we assumed mosquito females would
not fly toward the sea, we installed sticky ovitraps in only one
half of the perimeter in Tubiacanga, which resulted in a more
dense coverage in Tubiacanga than in Favela do Amorim. A
prior report attested that oviposition traps (OTs) placed in
the peridomestic area were more efficient at capturing Ae.
aegypti mosquitoes in comparison with the OTs placed in-
doors.20Adhesive papers were applied to the internal section
of ovitraps baited with hay infusion, as used in previous re-
ports.10Sticky ovitraps were checked only once at the end of
experimental period. Daily capture stopped when no dust-
marked females were collected by any method for 3 consecu-
tive days. Captured mosquitoes were examined under UV
light to check for the presence of fluorescent dust.
Survival analysis. Daily probability of survival (PDS) was
estimated by fitting two models: the exponential model21and
the nonlinear model by Buonaccorsi.22The exponential
model has been traditionally used for Ae. aegypti23but has
two fundamental drawbacks. It assumes a priori that mos-
quito mortality does not vary with increasing age and does not
consider removal of individuals by the capturing procedure.
Recently, a non-linear survival model was proposed, which
allows for the correction of estimates caused by the removal
C?t? = NStc?1 − c?t−1
where C(t) in the number of marked individuals captured on
day t; c is the daily capture probability; and S is daily survival
probability. Buonaccorsi22compared the two models using
Ae. aegypti data from MRR experiments conducted in Thai-
land24and found that the new model had a better fit to the
data. We fit both models to our data, using linear and non-
linear least squares standard procedures available in the soft-
ware R 2.0.25
From the lower and upper 95% limits of the confidence
interval for PDS (estimated by the nonlinear model), we de-
rived two quantities: the average life expectancy (ALE), de-
fined as 1/–logePDS26, and longevity, defined as PDS10,
(where 10 is the duration of the extrinsic incubation period
for dengue), which gives the expected proportion of mosqui-
toes surviving long enough to transmit dengue virus.
Dispersal analysis. The locations of all release and positive
capture points were geo-referenced using a Global Position
System (GPS; Garmin eTrex personal navigator, Gavmin In-
ternational, Olathe, KS) to calculate distance between release
and capture points. The flight behavior of Ae. aegypti females
in the two study areas was summarized by a set of dispersal
measures: mean distance traveled (MDT), maximum distance
traveled (MAX), and flight ranges (FR).27,28Frequency dis-
tributions of the numbers of marked mosquitoes recaptured
that had traveled < 100 and > 200 m from the release point
were constructed to further characterize the flight range of
Ae. aegypti females.
Ovarian development stage and parity rate. To evaluate the
evolution of ovarian development in marked females, all
dust-marked females captured with backpack battery-
powered aspirator had their ovaries removed in saline solu-
tion, and skeins of the ovarian tracheal system were evaluated
under a microscope. Ovarian of dissected females were clas-
sified according to Christophers,29with stages 1, 1–2, and 2
grouped as initial stages of development; stages 3 and 4
grouped as intermediary stages; and stage 5 classified as final
stage (gravid females).
All non-marked females captured (i.e., wild individuals)
were dissected for determination of parity based on the con-
dition of the tracheal system, as described by Detinova.30
Parity of natural population of Ae. aegypti for each season
and area was calculated as the number of parous divided by
the total number of females captured.
Comparisons between sites and seasons. Differences in sur-
vival rates between study areas and seasons were evaluated
by comparing the point estimates of survival rates by two-
sample t test.31To evaluate the effect of study area (categori-
cal variable: Tubiacanga/Amorim), season (categorical vari-
able: dry/wet season), and days since release (continuous vari-
able) on mosquito dispersal, we used generalized estimating
equations (GEEs), which are linear regression models with a
correction of variance caused by the blocking design (co-
horts). The GEE model was fitted, assuming an exchangeable
correlation structure.32In all analyzes, dispersal distances
were log-transformed, because this variable presented non-
normal distribution. All model fitting was performed using
the R 2.0 program (package geepack, for GEE fitting).25
Parity rates observed in field populations were compared
MACIEL-DE-FREITAS AND OTHERS
with ?2tests. The aim was to evaluate whether parity rates
observed during MRR experiments were different regarding
the study areas and seasons.
Ethical considerations. MRR experimental protocols were
submitted to and approved by Fiocruz Ethical Committee
(CEP/Fiocruz protocol no. 11591-2005).
Climate. In Tubiacanga, during the wet season, tempera-
ture varied from 23.3°C to 29.6°C, and month rainfall was
161.9 mm; during the dry season experiment, temperature
varied from 18.3°C to 26.5°C, with 44.1 mm of rainfall. In
Favela do Amorim, temperatures ranged from 19.8°C to
27.5°C during the dry season and from 23.8°C to 30.1°C dur-
ing the wet season, and precipitation was 58.3 and 125.9 mm,
Release and recapture. During the whole study, 8,792
marked Ae. aegypti females were released. Capturing lasted
8–13 days, and the proportion captured in each experiment
varied from 6.81% to 14.26% (Table 1).
Daily survival rates and longevity. Overall, the Buonaccorsi
model provided higher estimates of daily survival (PDS) than
the exponential model. This is expected because it corrects for
the removal of individuals. Within each site/season, cohorts
showed good agreement (Table 2). Mosquitoes released in
Tubiacanga showed lower survival than those released in
Favela do Amorim. This happened both in the dry and wet
seasons (dry season: t ? 17.58, P < 0.001; wet season: t ? 9.52,
P < 0.001). Survival differences between sites were found to
be as great as 10% during the wet season and 13% during the
dry season. Within each area, survival was higher during the
dry season (Tubiacanga: t ? 2.28, P < 0.05; Favela do
Amorim: t ? 17.4, P < 0.001).
Variation in PDS led to large variation in the ALE, which
varied from 3 to 12 days in Tubiacanga and 4 to 16 in Amorim
Dispersal. In Tubiacanga, 60% of females were captured
within a radius of 100 m from the releasing point in the wet
season (93% in the dry season). The average distance trav-
eled was 81–86 m, depending on the season, and the maxi-
mum distance traveled was 363 m. In Favela do Amorim, on
the other hand, 96% (dry season) and 100% (wet season) of
captured mosquitoes was found within 100 m from their re-
lease point, having traveled an average distance of 40 m in the
wet summer and 53 m in the dry winter; no female was found
beyond 200 m (Table 3). No significant differences were ob-
served between distances traveled in the dry and wet seasons
in both areas (t ? 0.16, P ? 0.87).
In Tubiacanga during the wet season, two marked (0.65%
from the total of collections) and five unmarked females were
captured, whereas in the dry season no marked and three
unmarked females were collected in sticky ovitraps. In Favela
do Amorim during the dry season, 5 marked (2.08%) and 1
unmarked females were collected, whereas in the wet season,
12 (4.41%) marked and 6 unmarked females were captured in
stick ovitraps. Low numbers of captures in the surroundings
of both areas suggests low emigration of mosquitoes. How-
ever, this information should be reviewed carefully, because
spacing between sticky ovitraps could be considered too high,
particularly in the Favela do Amorim area.
Number of dust-marked Ae. aegypti females released and captured per day in the two neighbors in Rio de Janeiro, using two capturing methods:
backpack battery-power aspirator and BGS-Traps
Tubiacanga wet season Tubiacanga dry seasonFavela do Amorim dry season Favela do Amorim wet season
Cohort 1Cohort 2 Cohort 3 Cohort 1Cohort 2 Cohort 3Cohort 1Cohort 2 Cohort 3Cohort 1 Cohort 2Cohort 3
11.38Recapture rate (%) 6.81 7.449.85 9.59
Survival analysis for three Ae. aegypti female cohorts released in
Tubiacanga and Favela do Amorim, Rio de Janeiro city, during wet
and dry seasons, and captured with backpack battery-powered as-
pirators and BGS-Traps
Daily probability survival (95% CI)
Favela do Amorim
Favela do Amorim
Daily probability of survival (DPS) was estimated by the exponential and Buonaccorsi
ALE, average life expectancy.
SURVIVAL AND DISPERSAL OF AEDES AEGYPTI
When a multivariate linear model was fit to the data (Table
4), “time since release” and “study area” had significant ef-
fects on dispersal. In Tubiacanga, marked mosquitoes were
found significantly far away from the release point than in
Amorim. No significant effect of season was found. Within-
cohort correlation was estimated as 0.1, which is marginally
non-significant (P ? 0.065). Assuming that cohort effect is
negligible, we may use R2to measure goodness-of-fit. We
found R2? 0.34.
Ovarian development stages. In Tubiacanga, during the
wet season, almost 60% of recovered mosquitoes in the first
day after release were in stage N. This proportion tended to
decrease in the following days (Figure 1A). Stages 3–4 and
gravid females started to appear at days 3 and 4 after release,
respectively. A similar pattern of ovarian development was
observed during the dry season (Figure 1B).
In Favela do Amorim, during the dry season, 40% were at
stage N at the first day, and we observed a steep decrease in
this frequency in subsequent collections (no females at stage
N were found after day 4). Females at stages 3–4 first
appeared in the second day after release, whereas gravid
females were collected on day 3 (Figure 2A). Similar patterns
of ovarian development were observed in the wet season (Fig-
Parity. In Tubiacanga, parity rates of 53.8% (N ? 405) and
48.7% (N ? 316) were observed in the captured wild popu-
lation during the wet and dry seasons, respectively. In Favela
do Amorim, during the dry season, the parity rate was 62.6%
(N ? 297) among unmarked captured females and 68.5%
(N ? 476) during the wet season. In both areas, parity rate
did not vary significantly between seasons (Tubiacanga: ?2?
2.13, P > 0.05; Amorim:?2? 2.51, P > 0.05). Parity rate ob-
served in Favela do Amorim was significantly higher than in
Tubiacanga in both seasons (wet season: ?2? 16.83, P < 0.05;
dry season: ?2? 6.02, P < 0.05).
This report provides substantial information about Ae. ae-
gypti ecology and vectorial potential under the natural con-
ditions of Rio de Janeiro city. It was the first time that survi-
val rate of dengue vector was evaluated locally, in a city
with intense history of dengue transmission during the last
20 years. Besides survival, longevity, dispersal, ovarian
development, and parity of the natural population were stud-
ied in two areas in different seasons. Our findings suggest that
both localities have appropriate conditions for elevated
dengue transmission. In Favela do Amorim, a poor commu-
nity with extremely high human density and disordered oc-
cupation, ecological conditions showed to be more adequate
for mosquito survival, and consequently, dengue transmis-
sion. High daily survival rates increase the chance of a mos-
quito to blood feed in a viremic person, become infective, and
transmit the virus. Indeed, almost one half of living females
could live for periods longer than the extrinsic incubation
Generalized estimating equations (GEE) model for log (dispersal) of
marked Ae. aegypti females released in Tubiacanga and Favela do
Amorim in both dry and wet seasons and captured with BGS-Traps
and backpack aspirators
Variable Effect SE Wald
Time after release
* Correlation structure used was the “exchangeable.”
FIGURE 1. Ovarian development stages of dust-marked Ae. aegypti
females released in the suburban area (Tubiacanga) and recaptured
with backpack aspirators. (A) MRR conducted in the wet season,
where 210 females were dissected. (B) MRR performed in the dry
season, where 118 females were collected.
Distances traveled by dust-marked Ae. aegypti females captured with backpack aspirators and BGS-Traps in a suburban neighbor (Tubiacanga)
and a slum in the urban area (Favela do Amorim) in Rio de Janeiro during the dry and wet seasons
Tubiacanga wet season Tubiacanga dry season Favela do Amorim dry seasonFavela do Amorim wet season
Females flying up to 100 m (%)
Females flying beyond 200 m (%)
MACIEL-DE-FREITAS AND OTHERS
period of dengue virus in the urban area of Favela do
Amorim during the dry season and 33.3% during the wet
season. Life expectancy estimated for Favela do Amorim is
long enough to allow four or even five gonotrophic cycles.
Thus, because Ae. aegypti females usually takes multiple
blood feedings during a single gonotrophic cycle and rarely
feed on sugar,33,34pathogen transmission could be enhanced
there. Even with a lower life expectancy, Tubiacanga also has
suitable conditions for dengue transmission, because ∼10% of
females could live longer than the extrinsic incubation period
(i.e., 10 days). However, because of its higher survival rates,
dengue transmission seems to be more feasible in Favela do
Amorim. Therefore, everything else been equal, it would be
reasonable to expect a higher number of dengue cases in
Favela do Amorim than in Tubiacanga.
During 2005, 983 dengue cases were reported in Rio de
Janeiro, 7 in Favela do Amorim, and 3 in Tubiacanga. Up to
the 18th entomological week of 2006, 9,408 dengue cases were
confirmed in Rio de Janeiro city, with 66 cases in Favela do
Amorim and 21 in Tubiacanga.35,36Thus, differences in mos-
quito survival are concordant with dengue incidence in both
areas, although the incidence of a disease in an area is depen-
dent of several factors (e.g., vector and host densities and
number of susceptible individuals).
Estimates of daily survival probabilities agree with previous
studies, which show values ranging between 0.70 and
0.90.24,37,38We found a tendency toward longer survival
during the dry winter. Possibly, weather conditions during
the wet and hot season were less favorable to adult mosqui-
toes; the mean temperature during tropical summer can be
∼30°C and reach a maximum temperature of > 40°C at
midday. On the other hand, the dry season generally presents
more suitable climatic conditions, with mean tempera-
tures rarely exceeding 28°C in a single day. Therefore, the
unfavorable weather conditions during the wet season
possibly constrained Ae. aegypti survival rate. The assumption
that adult daily survival is constant over the year, despite
seasonal trends in rainfall and temperature, was used in simu-
lation models describing vector population dynamics and the
epidemiology of dengue viruses in urban environments.39–41
The seasonal variation in survival rates observed in this report
can add accuracy to current models.39–41
Regarding dispersal, we found differences between areas
but not seasons. This result agrees with previous studies.5
Females released in Tubiacanga dispersed more than those
released in Favela do Amorim. Tubiacanga and Amorim
share approximately the same density of water-holding con-
tainers (R. Maciel-de-Freitas, unpublished data), but human
density in Favela do Amorim is almost three times greater. It
is possible that the diversity of obstacles posed by the irregu-
lar and dense constructions in Favela do Amorim, associated
with high availability of blood sources, constrained mosquito
dispersal in this area, where very few or none mosquito flew
beyond 100 m from the release site. Because mosquito cap-
turing was performed in houses randomly selected rather than
in concentric circles from the release point, dispersal rate
estimations may be biased. However, dispersal distances
found in this study agrees with similar studies performed by in
Dispersal has important consequences for dengue control.
Usually, the first action after the identification of a new
dengue case is the implementation of source reduction activi-
ties and larvicides applications within a ring area centered at
the identified case. Our results suggest that a ring with radius
? 200 m would be appropriate in both Tubiacanga and
Favela do Amorim areas. Indeed, a smaller ring could be
established in the slum area, because mosquitoes displaced
Data on ovarian development of marked females in Tubia-
canga and Favela do Amorim showed important differences.
Mainly, the continuous appearance of females at stage N in
Tubiacanga and the appearance of gravid females 3 days
after release in Favela do Amorim (in Tubiacanga gravid
females were collected just 4 days after release) reinforce the
idea that females can complete a gonotrophic cycle earlier in
more densely human populated areas, where the probability
of a blood-seeking female encounter a host is higher.
Parity rate in Favela do Amorim was higher than in Tubia-
canga in both dry and wet seasons. However, with the study
design reported in this paper, the authors are not confident to
state if higher parity was a consequence of higher human
density or higher mosquito survivorship, which was also
higher in Favela do Amorim.
Recapture rates varied between the MRR experiments.
Lower recapture was observed during dry season (mean of
7.88% and 11.23% for Tubiacanga and Favela do Amorim,
respectively), when the highest survival rates were observed.
Low rates can lead to a bias in results concerning mortality
rate and dispersal, although observed recapture rates were
similar or even higher than several prior reports.9,37
In summary, this study assesses several aspects of mosquito
ecology in two distinct neighbors in Rio de Janeiro, a city
where vector density and dengue transmission have shown
to be highly heterogeneous1and mosquito control faces
FIGURE 2. Ovarian development stages of dust-marked Ae. aegypti
females released in the urban area (Favela do Amorim) and recap-
tured with backpack aspirators. (A) MRR conducted in the dry sea-
son, where 107 females were dissected. (B) MRR performed in the
wet season, where 201 females were collected.
SURVIVAL AND DISPERSAL OF AEDES AEGYPTI
several drawback to be surpassed, such as insecticide
resistance and reduced effectiveness in mosquito control be-
cause of the inaccessibility to some areas such as slums be-
cause of criminality.43–45Other field evaluations concerning
daily survival rates, longevity, dispersal, ovarian develop-
ment, and parity in different areas and seasons would give
more light for the understanding of dengue transmission pat-
tern in Rio de Janeiro and improve mosquito control.
Received January 26, 2006. Accepted for publication June 24, 2006.
Acknowledgments: We thank Gláuber Rocha, Kleber Soares,
Marcelo Celestino dos Santos, Marcelo Quintela Gomes, Mauro
Menezes Muniz, Reginaldo Rego, Renato Carvalho, Roberto Costa
Peres, and Sérgio Cunha for field assistance. We also thank Dr
Álvaro Eiras for allowing the use of BG-Sentinels and the com-
ments of two anonymous referees that improved this manuscript
Financial support: This study was supported by FAPERJ, CNPq, and
Authors’ addresses: R. Maciel-de-Freitas and R. Lourenc ¸o-de-
Oliveira, Laborato ´rio de Transmissores de Hematozoa ´rios, Pav. Car-
los Chajas, 4° andar, Sala 04, Departamento de Entomologia. CEP:
21040-360. Fundac ¸ao ´ Oswaldo Cruz, Fiocruz, Rio de Janiero, Brazil.
E-mails: firstname.lastname@example.org, email@example.com. C. T. Co-
dec ¸o, Programa de Computaco ´ Cientifica, Fundac ¸ao Oswaldo Cruz,
Fiocruz, Rio de Janiero, Brazil. CEP: 21040-360. E-mail: codeco@
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