Human and Environmental Factors Affecting Aedes aegypti

Distribution in an Arid Urban Environment

Author(s): Kathleen R. Walker, Teresa K. Joy, Christa Ellers-Kirk, and Frank B.
Ramberg
Source: Journal of the American Mosquito Control Association, 27(2):135-141.
2011.
Published By: The American Mosquito Control Association
DOI: http://dx.doi.org/10.2987/10-6078.1
URL: http://www.bioone.org/doi/full/10.2987/10-6078.1

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the

biological, ecological, and environmental sciences. BioOne provides a sustainable online
platform for over 170 journals and books published by nonprofit societies, associations,
museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content
indicates your acceptance of BioOnes Terms of Use, available at www.bioone.org/page/
terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial
use. Commercial inquiries or rights and permissions requests should be directed to the
individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit
publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to
critical research.

Journal of the American Mosquito Control Association, 27(2):135141, 2011

Copyright E 2011 by The American Mosquito Control Association, Inc.

HUMAN AND ENVIRONMENTAL FACTORS AFFECTING AEDES

AEGYPTI DISTRIBUTION IN AN ARID URBAN ENVIRONMENT
KATHLEEN R. WALKER, TERESA K. JOY, CHRISTA ELLERS-KIRK

AND

FRANK B. RAMBERG

Center for Insect Science, University of Arizona, Tucson, AZ 85721

ABSTRACT. Aedes aegypti has reappeared in urban communities in the southwestern USA in the 1990s
after a 40-year absence. In 2003 and 2004, a systematic survey was conducted throughout metropolitan
Tucson, AZ, to identify human and environmental factors associated with Ae. aegypti distribution within an
arid urban area. Aedes aegypti presence and abundance were measured monthly using the Centers for Disease
Control and Prevention enhanced oviposition traps at sampling sites established in a grid at 3- to 4-km
intervals across the city. Sampling occurred in the summer rainy season (July through September), the peak
of mosquito activity in the region. Multiple regression analyses were conducted to determine relationships
between mosquito density and factors that could influence mosquito distribution. House age was the only
factor that showed a consistent significant association with Ae. aegypti abundance in both years: older houses
had more mosquito eggs. This is the 1st study of Ae. aegypti distribution at a local level to identify house age
as an explanatory factor independent of other human demographic factors. Further research into the reasons
why mosquitoes were more abundant around older homes may help inform and refine future vector
surveillance and control efforts in the event of a dengue outbreak in the region.
KEY WORDS

Aedes aegypti, urban, house age, mosquito ecology

INTRODUCTION
The dengue vector Aedes aegypti (L.) (Diptera:
Culicidae), originally an African species, is now
widely distributed throughout most tropical and
subtropical regions of the world (Tabachnick and
Powell 1979). Several factors may explain the
cosmopolitan distribution of this mosquito, including its close association with human habitations, its tendency to oviposit in man-made
containers, and the ability of the eggs to withstand
long periods of desiccation (Christophers 1960,
Lounibos 2002). These characteristics may help
Ae. aegypti colonize and thrive even in arid
environments such as the desert region of the
southwestern USA, where water is limited and
human settlements are widely dispersed.
It is not known when Ae. aegypti first arrived
in southern Arizona, but the species was clearly
established in the region by the 1930s (Bequaert
1946, Murphy 1953). Between 1946 and 1994,
however, the species was not detected in the
region, despite intensive surveillance efforts
(Hayes and Tinker 1958, Tinker and Hayes
1959, McDonald et al. 1973). The reasons for
the disappearance of the species are not clear, as
formal Ae. aegypti eradication programs were
not implemented in the state (PAHO 1997). The
mosquito reappeared in southern Arizona in 1994
and is now well established in towns throughout
this desert region (Engelthaler et al. 1997, Fink et
al. 1998). Genetic analyses of southern Arizona
Ae. aegypti population structure indicate a close
relationship to populations in the Pacific coast of
Mexico (Gorrochotegui-Escalante et al. 2002), as
well as some genetic exchange with populations
in the eastern USA (Merrill et al. 2005). Both

studies also suggest that human migration,

particularly along highways, may be the primary
vehicle for Ae. aegypti movement in the southwestern USA, as the natural landscape is
inhospitable to this mosquito.
While climate plays a major role in determining
the geographic range and population dynamics of
Ae. aegypti, human settlements and water use can
mitigate the effects of climate on the mosquito
populations, in some cases allowing larval development even during extended dry periods (Focks
et al. 1993a, 1993b; Rodhain and Rosen 1997).
Throughout the present global range of the
mosquito, differences in human behavior, particularly related to water storage, lead to variation
in Ae. aegypti ecology. For example, while female
mosquitoes can oviposit in a wide variety of
artificial and natural containers of water, the type
of larval habitat that produces most mosquitoes
may be large water storage tanks in one
community or region (e.g., Thavara et al. 2001)
and small plastic containers in another (Fulmali
et al. 2008).
Variability is a striking feature of this mosquitos distribution. On a small scale within a single
city, Ae. aegypti distribution may be uniform or
clumped. For example, a study in Manaus
observed fairly homogeneous Ae. aegypti distribution throughout different neighborhoods
(Rios-Velasquez et al. 2007), while a study in
another Brazilian city, Porto Alegre, found
significantly higher vector density in particular
sections of the city (Barcellos et al. 2005). Certain
human factors, particularly low socioeconomic
status, are often associated with Ae. aegypti
infestations (Von Wideguth et al. 1969, Spiegel

135

136

JOURNAL OF THE AMERICAN MOSQUITO CONTROL ASSOCIATION

et al. 2007, Caprara et al. 2009, Quintero et al.

2009), but some studies have actually found Ae.
aegypti density to be higher in wealthier neighborhoods (Barcellos et al. 2005, Maciel-de-Freitas
et al. 2007, Rios-Velasquez et al. 2007).
Efforts to understand and predict Ae. aegypti
distribution, particularly at the local level relevant for dengue control, must examine both
environmental and human factors to determine
their relationships to mosquito density. In 2003
and 2004, we conducted a systematic survey
throughout metropolitan Tucson, AZ, to identify
human and environmental factors associated with
Ae. aegypti distribution in an arid urban area. As
the goal was to tease out the relative importance
of different factors, the study involved a broad
survey of .40 geographically separated houses
rather than an in-depth examination of a few
neighborhoods. In an effort to make our findings
easy to incorporate into future modeling efforts,
we chose only those factors for which information could be obtained through public databases.
Factors examined at each mosquito sampling site
included the percentage of the site covered with
vegetation, human per capita income levels, and
human housing density, which have been associated with Ae. aegypti infestations in other studies
(e.g., Tsuda et al. 2006, Von Wideguth et al. 1969,
Cox et al. 2007, respectively), but we also assessed
the proximity of dry creek beds (washes) and
house age, as these features have been anecdotally
associated with Ae. aegypti presence in the region.
MATERIALS AND METHODS
Study area
Tucson is a large and rapidly growing city in
southern Arizona, with a metropolitan population of .800,000 (PAG 2002). Located in the
Sonoran Desert, Tucson experiences moderate
winters (average January low temperature 5
3.8uC), prolonged hot summers (average May
September high temperature 5 35.7uC), and
limited precipitation (311.1 mm annually) (NWS
2007). Some rainfall occurs during the winter, but
most rainfall occurs during the summer rainy
season between July and September.
Characterization of sampling sites
In 2003, a total of 47 sampling sites, all private
residences, were established at 3- to 4-km intervals
in a grid-like pattern across the Tucson metropolitan area (Fig. 1a). Most sites were established
by randomly selecting a house at an appropriate
distance from other sampling sites and asking
residents whether they would permit mosquito
sampling on their property. Due to residents
requests, several sampling sites were eliminated in
2004, leaving a total of 42 sampling sites.

VOL. 27, NO. 2

Human and environmental factors potentially

associated with Ae. aegypti distribution were
determined for each site using publicly available,
geographically referenced data provided by the
Pima County Department of Transportation
(Pima DOT 2005). Residents remained anonymous, and all data used to characterize sites were
derived from public sources, not interviews. The
factors and their ranges are summarized in
Table 1. The number of dry creek beds within
200 m of each site was determined using
Geographical Information System (GIS) analysis
of Pima DOT (2005). Estimates of vegetation
cover were based on visual examination of GIS
images developed from aerial photos taken
between August and September 2002 (Pima
DOT 2005). Values reflect the percentage of green
area within 50-m radius of traps. Sites were
classified into 1 of 4 vegetation cover levels: 1 5
,10% of the area covered in green vegetation, 2 5
10 to 25% coverage, 3 5 25 to 50% coverage, and
4 5 .50% coverage. Information on human
population density and per capita income was
derived from 2000 census results. Rather than
measure human density on a household level,
census blocks were used to represent the density of
the neighborhood around the sampling site. In
urban areas, a census block is typically an actual
city block. Per capita income figures were derived
from census tract data for the tract containing
each sampling site. Census tracts have between
2,500 and 8,000 persons and, when first created,
are designed to be homogeneous with respect to
population characteristics, economic status, and
living conditions (US Census Bureau 2000).
House age for each site was determined using
land-parcel information available to the public
(Pima DOT 2005).
Mosquito sampling
Aedes aegypti presence and abundance were
measured outdoors at each of the sampling sites
using 2 sets of the Centers for Disease Control
and Prevention enhanced ovitraps as designed by
Reiter et al. (1991). Mosquitoes were only
sampled outdoors because indoor sampling was
considered too invasive by many residents, and
because indoor house environments are typically
shut off from the outdoors by well-fitting doors
and windows. Sampling was conducted using
ovitraps, as these are efficient and highly sensitive
indictors of Ae. aegypti presence (Fay and
Eliason 1966) and are much less labor intensive
than traditional house or Breteau indices (Morato et al. 2005). Each enhanced trap set consists
of a pair of 1-qt jars painted black, containing
hay-infusion water and lined with seed germination paper (Anchor Paper Co., St. Paul, MN).
One jar in the trap contains pure hay infusion as
an attractant, while the other contains a 10%

JUNE 2011

Fig. 1.

AEDES AEGYPTI DISTRIBUTION

137

Cumulative Aedes aegypti egg counts at each site in (a) 2003 and (b) 2004.

mixture of hay infusion and water to create an

ideal oviposition site. In our study, the 2 jars were
placed side by side in a shaded location in or near
likely mosquito resting areas. The 2 sets of traps

were set out at locations between 5 and 10 m

apart. Traps were left at sites for approximately
72 h and then removed. All eggs collected on the
germination papers were counted, hatched, and

138

JOURNAL OF THE AMERICAN MOSQUITO CONTROL ASSOCIATION

Table 1.

VOL. 27, NO. 2

Characterization of sampling sites.

Site characteristic

Range of values

Elevation
Number of dry creek beds within 200 m of traps
Vegetation cover1
Human population density (census block)2
Per capita income in 1999 (census tract)2
House age (in 2003)

699925 m
05
1 (,10%) to 4 (.50%)
1 (0800 persons/mi2) to 8 (.9,600 persons/mi2)
$6,01853,743
484 years

1
Estimates of vegetation cover were based on visual examination of Geographical Information System images developed from
aerial photos taken between August and September 2002 (Pima County Department of Transportation 2005). Values reflect the
percentage of green area within 50-m radius of traps.
2
Information on human population density and per capita income were derived from 2000 census results and reflect the entire
census block or tract containing the sampling site, not just the site itself.

reared to the adult stage for identification.

Sampling occurred monthly during the 2003 and
2004 summer rainy seasons (July through September), the peak of mosquito activity in the
region (Walker, unpublished data).
Statistical analyses
Relationships between cumulative Ae. aegypti
egg counts and the human and environmental
factors (Table 1) were determined for each year
using multiple regression. The egg count and
human housing density data were log transformed
to improve the assumptions of normality and
homogeneity of variance. The initial analysis
involved fitting a full model using all 6 variables
and all possible 2-way interactions of those
variables. We then eliminated nonsignificant variables and interactions (P . 0.1) and created a 2nd,
reduced model. An extra sum of squares F-test
(also known as partial F-test) was used to compare
the full and reduced models to ensure that the
removed variables did not explain a significant
proportion of the variation (P , 0.05) in the
response variable (Ramsey and Shafer 2002).
For each year, logistic regression was also used
to evaluate the relationship between the odds of
Ae. aegypti presence and the same human and
environmental factors. The initial analysis involved creating a full model using all 6 variables.
Inclusion of all possible 2-way interactions was
not possible at this stage due to overfitting of the
data that destabilized the model (JMP 2001). We
then eliminated nonsignificant variables (P . 0.1)
and fit a reduced model including the significant
variables and all possible 2-way interactions.
Significance of the factors in the logistic regression
models was determined with likelihood ratio x2
tests. JMP (2001) was used for all statistical
analyses.

were collected from 16 sites in 2004. Again, all

adults reared were identified as Ae. aegypti. Culex
sp. egg rafts were observed at 20 of the sites. Egg
counts in 2003 indicated that Ae. aegypti was
widely distributed throughout the Tucson metropolitan area, but was most abundant in the
central part of the city (Fig. 1a). Aedes aegypti
was less abundant and more patchily distributed
in 2004 (Fig. 1b). Total rainfall also differed
during the 2 years. The total rainfall for 2003 was
10.05 in., primarily in August and September,
while the total rainfall for 2004 was 7.62 in., most
of which occurred in January through March.
Multiple regression models showed significant
relationships between log-transformed cumulative Ae. aegypti egg counts and the explanatory
variables in each year (2003: R2 5 0.75, df 5 16,
29, F 5 5.33, P , 0.0001; 2004: R2 5 0.30, df 5 3,
38, F 5 5.48, P 5 0.003). Table 2 shows those
explanatory variables included in the final,
reduced multiple regression models in each year.
Elevation, the number of dry creek beds within
200 m of the sampling site, percent vegetation
cover, and human population density were not
significantly associated with Ae. aegypti egg
counts in either year. In contrast, house age was
positively associated with Ae. aegypti egg counts
in both years. In 2003, per capita income was
negatively associated with egg counts. In both
years, the interaction between house age and per
capita income was also significant. In 2003 only,
the interaction between per capita income and
human population density was also significant.
Logistic regression analyses of Ae. aegypti
presence versus human and environmental factors
also showed a significant positive association
between house age and the likelihood of collecting eggs in 2003 (P 5 0.001, x2 5 10.39), but no
factors were significant in 2004.
DISCUSSION

RESULTS
A total of 4,231 eggs were collected from 31
sites in 2003. All adults reared were identified as
Ae. aegypti. Culex sp. egg rafts were also
observed at 41 of the sites. A total of 958 eggs

We found that house age was consistently

associated with Ae. aegypti density, both in a
normal rain year, 2003, and a dry year, 2004.
Mosquitoes were more abundant and, in 2003 at
least, more frequently found around older houses.

JUNE 2011
Table 2.

AEDES AEGYPTI DISTRIBUTION

139

Relationships between log-transformed Aedes aegypti egg counts and explanatory variables.1

Explanatory variable

Slope

Standard error

t-ratio

P-value

20032
Elevation
Per capita income
House age
Dry creek beds within 200 m
Vegetation class (1)
Vegetation class (2)
Human density (log)

0.179
23.97 3 1025
0.040
20.249
20.237
0.153
20.640

2.24
1.17 3 1025
0.0098
0.17
0.18
0.14
0.33

0.08
23.39
4.14
21.48
21.33
1.09
21.96

0.94
0.002*
0.0003*
0.15
0.19
0.28
0.06

Significant interactions
Income 3 house age
Income 3 human density

2.21 3 1026
26.04 3 1025

1.0 3 1026
2.6 3 1025

2.19
22.32

0.037*
0.028*

23.32 3 1026
0.043

1.03 3 1025
0.016

20.32
2.65

0.75
0.012*

3.84 3 1026

1.36 3 1026

2.83

0.007*

2004
Per capita income
House age
Significant interaction
Income 3 house age
1

The asterisk (*) indicates a statistically significant relationship between the explanatory variable and log-transformed egg counts.
2
In 2003, many 2-way variable interactions were marginally significant (P 5 0.1) in the full model, so the final model includes all 6
variables.

To our knowledge, this is the 1st study of local

distribution of Ae. aegypti to document a
relationship between mosquito abundance and
house age while accounting for the effect of other
factors potentially affecting Ae. aegypti distribution. A likely explanation for this finding is that
human habitations become more hospitable to
Ae. aegypti over time, as vegetation matures,
house structures age, and objects collect in the
yard. Further research is needed to determine
which specific characteristics of older houses
cause Ae. aegypti to be more abundant. A
preliminary examination of outdoor water containers in half of the sites used in this study did
not find significant differences in the numbers of
water-holding containers at older versus newer
houses, but did find more containers that were
positive for Ae. aegypti larvae around older
houses (Walker, unpublished data).
The inverse association between income level
and Ae. aegypti density in 2003 suggests that
houses in poorer neighborhoods provide better
habitat for mosquitoes. Houses in such neighborhoods tend to be less well maintained, often
have accumulations of objects in the yard, and
are close to vacant or abandoned properties
(Walker, personal observation). The significant
interactions of per capita income with both house
age and human housing density indicate that
income also influences the relationships of egg
counts with these other variables. Older houses in
low-income neighborhoods tended to have higher
egg counts than houses of similar age in wealthier
neighborhoods. Similarly, houses in high-density,
low-income neighborhoods tended to have higher
egg counts than houses in high-density, higherincome neighborhoods.

The lack of a significant relationship between

percent vegetation cover and mosquito presence
and abundance is interesting, as a cross-border
USA/Mexico study in the same region did find a
significant relationship between vegetation cover
and Ae. aegypti presence (Hayden et al. 2010).
While overall vegetation cover did not predict
mosquito abundance in the present study, the
positive association between house age and
mosquito density could indicate that older
vegetation (e.g., mature shade trees) could favor
Ae. aegypti by providing adult mosquito resting
sites. A study in Thailand by Tsuda et al. (2006)
that examined Ae. aegypti and Ae. albopictus
distribution along an urban to rural gradient,
however, did not find a positive relationship
between Ae. aegypti density and average tree
height. The same study found the lowest numbers
of Ae. aegypti in the intermediate or transitional
neighborhood that was also the most recently
constructed human community. In their study in
South Florida, Rey et al. (2006) found significant
negative relationships between Ae. aegypti egg
counts and several measures of vegetation cover.
The relationship between Ae. aegypti distribution
and vegetation cover thus appears complex and
not easily generalized.
Because this study focused on mosquito
distribution within a single urban area, climatic
factors such as temperature and precipitation
were not included. It is possible, however, that
microclimate differences between sites could be
sufficiently great as to explain some of the
observation variation in Ae. aegypti presence
and abundance. While microclimate analysis was
beyond the scope of this study, the study by
Hayden et al. (2010), which included both Tucson

140

JOURNAL OF THE AMERICAN MOSQUITO CONTROL ASSOCIATION

and the border cities of Nogales, did examine

microclimate variation between sites across the
entire region. That study found significant
relationships between mosquito presence and
microclimate in some years, but not others.
The dramatic difference in Ae. aegypti abundance between a relatively normal rain year and
an extremely dry year demonstrates the powerful
effect of regional climate on this species, at least
in an arid environment. These results concur with
some studies of Ae. aegypti in Southeast Asia
(Kuno 1997), but other studies of this species in
the humid tropics have not observed a strong
association between rainfall and mosquito abundance, perhaps due to this species reliance on
breeding and resting habitat created by humans
(Focks et al. 1993b, Rodhain and Rosen 1997,
Maciel-de-Freitas et al. 2007, Rios-Velasquez et
al. 2007).
A number of different models have been
developed that incorporate both entomological
and epidemiological factors to predict the spatial
distributions of both Ae. aegypti and dengue, but
house age has not been included as a possible
explanatory variable (Eisen and Lozano-Fuentes
2009). Further research is needed to determine
the mechanisms behind the observed association
between mosquitoes and older houses, and to
ascertain whether this association is limited to
this particular region and climate. If house age
proves to be predictive at other scales and in
other regions, this studys finding may be useful
in future dengue surveillance and control efforts.
ACKNOWLEDGMENTS
The authors thank Yves Carrie`re for statistical
advice and Mary Hayden for helpful comments on
the manuscript. We also appreciate the many
residents who allowed mosquito trapping on their
properties. The study was supported by the
National Institutes of Health IRACDA Fellowship
Program through the PERT Program at the Center
for Insect Science at the University of Arizona.
REFERENCES CITED
Barcellos C, Pustai AK, Weber MA, Brito MR. 2005.
Identification of places with potential transmission of
dengue fever in Porto Alegre using Geographical
Information Systems. Rev Soc Bras Med Trop
38:246250.
Bequaert J. 1946. Aedes aegypti, the yellow fever
mosquito, in Arizona. Bull Brooklyn Entomol Soc
41:157.
Caprara A, Lima JW, Marinho AC, Calvasina PG,
Landim LP, Sommerfeld J. 2009. Irregular water
supply, household usage and dengue: a bio-social
study in the Brazilian northeast. Cad Saude Publica
25(Suppl 1):S125S136.
Christophers SR. 1960. Aedes aegypti (L.), the yellow
fever mosquito: its life history, bionomics and struc-

VOL. 27, NO. 2

ture. Cambridge, United Kingdom: Cambridge Univ.

Press.
Cox J, Grillet ME, Ramos OM, Amador M, Barrera R.
2007. Habitat segregation of dengue vectors along an
urban environmental gradient. Am J Trop Med Hyg
76:820826.
Eisen L, Lozano-Fuentes S. 2009. Use of mapping and
spatial and space-time modeling approaches in operational control of Aedes aegypti and dengue. PLoS
Neglected Trop Dis [Internet] 3:e411 [accessed October
28, 2009]. Available from: www.plosntds.org/article/
info%3Adoi%2F10.1371/journal.pntd.0000411.
Engelthaler DM, Fink TM, Levy CE, Leslie MJ. 1997.
The reemergence of Aedes aegypti in Arizona. Emerg
Infect Dis 3:241242.
Fay RW, Eliason DA. 1966. Preferred oviposition sites
as a surveillance method for Aedes aegypti. Mosq
News 25:531535.
Fink TM, Hau B, Baird BL, Palmer S, Kaplan S,
Ramberg FB, Mead DG, Hagedorn HH. 1998.
Aedes aegypti in Tucson, Arizona. Emerg Infect Dis
4:12.
Focks DA, Haile DG, Daniels E, Mount GA. 1993a.
Dynamic life table model for Aedes aegypti (Diptera:
Culicidae): analysis of the literature and model
development. J Med Entomol 30:10031017.
Focks DA, Haile DG, Daniels E, Mount GA. 1993b.
Dynamic life table model for Aedes aegypti (Diptera:
Culicidae): simulation results and validation. J Med
Entomol 30:10181028.
Fulmali PV, Walimbe A, Mahadev PVM. 2008. Spread,
establishment and prevalence of dengue vector Aedes
aegypti (L.) in Konkan region, Maharashtra, India.
Indian J Med Res 127:589601.
Gorrochotegui-Escalante N, Gomez-Machorro C, Lozano-Fuentes S, Fernandez-Salas I, Munoz ML,
Farfan-Ale JA, Garcia-Rejon J, Beaty BJ, Black
WC IV. 2002. Breeding structure of Aedes aegypti
populations in Mexico varies by region. Am J Trop
Med Hyg 66:213222.
Hayden MH, Uejio C, Walker K, Ramberg F, Moreno
R, Mearns LO, Rosales C, Gameros M, ZielinskiGutierrez E, Janes CR. 2010. Microclimate and
human factors in the divergent ecology of Aedes
aegypti along the Arizona U.S./Sonora MX border.
Ecohealth 7:6477.
Hayes GR, Tinker ME. 1958. The 19561957 status of
Aedes aegypti in the United States. Mosq News
18:253257.
JMP. 2001. Statistics and graphics guide. Cary, NC:
SAS Institute.
Kuno G. 1997. Factors influencing the transmission of
dengue viruses. In: Gubler DJ, Kuno G, eds. Dengue
and dengue hemorrhagic fever. Wallingford, United
Kingdom: CABI Publishing. p 6188.
Lounibos LP. 2002. Invasions by insect vectors of
human disease. Annu Rev Entomol 47:233266.
Maciel-de-Freitas R, Marques WA, Peres RC, Cunha
SP, Lourenco de Oliveira R. 2007. Variation in Aedes
aegypti (Diptera: Culicidae) container productivity in
a slum and a suburban district of Rio de Janeiro
during dry and wet seasons. Mem Inst Oswaldo Cruz
102:489496.
McDonald JL, Sluss TP, Lang JD, Roan CC. 1973.
Mosquitoes of Arizona. Technical Bulletin 205.
Tucson, AZ: Univ. of Arizona Agricultural Experiment Station.

JUNE 2011

AEDES AEGYPTI DISTRIBUTION

Merrill SA, Ramberg FB, Hagedorn HH. 2005.

Phylogeography and population structure of Aedes
aegypti in Arizona. Am J Trop Med Hyg 72:304310.
Morato VC, Teixeira MG, Gomes AC, Bergamaschi
DP, Barreto ML. 2005. Infestation of Aedes aegypti
estimated by oviposition traps in Brazil. Rev Saude
Publica 39:553558.
Murphy D. 1953. Collection records of some Arizona
mosquitoes. Entomol News 14:233238.
National Weather Service [NWS]. 2007. National Weather
Service Forecast OfficeTucson, AZ [Internet]. Tucson,
AZ: National Weather Service Forecast Office [accessed November 11, 2009]. Available from: http://
www.wrh.noaa.gov/twc/climate/seaz_yearly_rainfall.
php.
Pan American Health Organization [PAHO]. 1997.
Dengue and dengue hemorrhagic fever in the Americas:
guidelines for prevention and control. Scientific Publication 548. Washington, DC: Pan American Health
Organization.
Pima Association of Governments [PAG]. 2002. 2002
population handbook. Tucson, AZ: Pima Association
of Governments.
Pima County Department of Transportation [Pima
DOT]. 2005. Pima County MapGuide maps [Internet].
Tucson, AZ: Technical ServiceGIS Database
Services [accessed October 26, 2005]. Available from:
http://www.dot.pima.gov/gis/maps/mapguide.
Quintero J, Carrasquilla G, Suarez R, Gonzalez C, Olano
VA. 2009. An ecosystemic approach to evaluating
ecological, socioeconomic and group dynamics affecting the prevalence of Aedes aegypti in two Colombian
towns. Cad Saude Publica 25(Suppl 1):S93S103.
Ramsey FL, Shafer DW. 2002. The statistical
sleuth: a course in methods of data analysis. Pacific
Grove, CA: Duxbury.
Reiter P, Amador MA, Colon N. 1991. Enhancement of
the CDC ovitrap with hay infusions for daily
monitoring of Aedes aegypti populations. J Am Mosq
Control Assoc 7:5255.
Rey JR, Nishimura N, Wagner B, Braks MAH,
OConnell SM, Lounibos LP. 2006. Habitat segregation of mosquito arbovirus vectors in south Florida.
J Med Entomol 43:11341141.

141

Rios-Velasquez CM, Codeco CT, Honorio NA, Sabroza PS, Moresco M, Cunha ICL, Levino A, Toledo
LM, Luz SLB. 2007. Distribution of dengue vectors
in neighborhoods with different urbanization types of
Manaus, state of Amazonas, Brazil. Mem Inst
Oswaldo Cruz 102:617623.
Rodhain F, Rosen L. 1997. Mosquito vectors and
dengue virusvector relationships. In: Gubler DJ,
Kuno G, eds. Dengue and dengue hemorrhagic fever.
Wallingford, United Kingdom: CABI Publishing.
p 4560.
Spiegel JM, Bonet M, Ibarra AM, Pagliccia N,
Ouellette V, Yassi A. 2007. Social and environmental
determinants of Aedes aegypti infestation in Central
Havana: results of a case-control study nested in an
integrated dengue surveillance programme in Cuba.
Trop Med Int Health 12:503510.
Tabachnick WJ, Powell JR. 1979. A world-wide survey
of genetic variation in the yellow fever mosquito,
Aedes aegypti. Genet Res 34:215229.
Thavara U, Tawatsin A, Chansang C, Kong-ngamsuk
W, Paosriwong S, Boon-Long J, Rongsriyan Y,
Komalmisra N. 2001. Larval occurrence, oviposition
behavior and biting activity of potential mosquito
vectors of dengue on Samui Island, Thailand. J Vector
Ecol 26:172180.
Tinker ME, Hayes GR Jr. 1959. The 1958 Aedes aegypti
distribution in the United States. Mosq News 19:
7378.
Tsuda Y, Suwonkerd W, Chawprom S, Prajakwong S,
Takagi M. 2006. Different spatial distribution of
Aedes aegypti and Aedes albopictus along an urban
rural gradient and the relating environmental factors
examines in three villages in northern Thailand. J Am
Mosq Control Assoc 22:222228.
US Census Bureau. 2000. Census tracts and block
numbering areas [Internet]. Tucson, AZ: US Census
Bureau [accessed March 9, 2005]. Available from:
http://www.census.gov/geo/www/cen_tract.html.
Von Wideguth DL, Eliason DA, Kilpatrick JW, Jakob
WL. 1969. The transitory nature of Aedes aegypti
larval habitats in an urban situation. Mosq News
29:495496.