Kissing Bugs in the United States
The Kanlal $chool Nafualilf Vol rJ No2
Department of Biological Sciences
E EMPORIA STATE UNIVERSITY Spring
2011
Kansas School Naturalist rSSN 0022-877X Published by EMPORIA STATE UNIVERSITY Editor JOHN RICHARD SCHROCK Editorial Comminee TOM EDDY BILL JENSEN MARSHALL SUNDBERG R BRENT THOMAS ERIC YANG Circulacion and Mailing ROGER FERGUSON Circulacion (chis issue) 10000 Press Run 15000 Media Designer John Decker Primed by McCormick Armscrong
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Kissing Bugs in the INTRODUCTION
Kissing bugs are bloodsuc ing =-_
m the order Hemiptera in ( e
Reduviidae Almost all knOmiddot
reside in the New World
They are best known a5
of Chagas disease a di sease 3
by transmission of the blood shy
Trypanosoma cruzi through (heir ~~~-
Vector transmission of Chaga ~
occurs with a number of e-=- _
kissing bugs Chagas is largel ~ O - --~2
to Mexico Central America an~ ~ __ -
America
Although only a few case- 0 i _ shy
Front cover Tiiatoma gerstaeckeri common in sourhern and central Texas Phoro used with permission of Mike Quinn TexasEnronet
Justin O Schmidt has a PhD in Entomology from the University of Georgia He runs a non-profit organizarion Sourhwesrern Biological Insrirure in Tucson Arizona
Lori Stevens is Professor of Biology ar The Universiry of Vermont She received her MS and PhD degrees from rhe University of Illinois Chicago Srevens is intrigued wieh ehe generic inreractions of hosrs and parasites She enjoys visiring family in Leavenworch Kansas
Patricia Dom is a Professor of Biological Sciences ar Loyola University New Orleans She received her PhD from rhe University of Maryland and began her work on Trypanosomiais during her posr-docrora l fellowship ar Sranford Medical School
Mark Mosbacher received his doctoraee of vererinary medicine from the University of Missouri He currently practices emergency veterinary medicine in Phoenix Arizona
John K10tt has a PhD in Entomology from the Universi ty of Kansas and is an Urban Entomology Specialisr Emeritus ar the University of California Riverside and lives in Sedona Arizona
Stephen A K10tt graduared from rhe Universi ty of Kansas and is Chief of Infeceious Diseases ar rhe Universiry of Arizona
transmi ned Chagas disease hamiddote
in the US rhese bugs are corn-middot
southern southeastern and sourhmiddots- --
states
We discuss the biology 0 - ~
bugs their evolu rionary hiscof- - shy
distribution in the United Srare t
they pose for transmission of T shy
pets and other domestic animals 21 _ ~ _
control of kiSSing bugs around h --
habitarions
EVOLUTION OF KISSING BUGS
The 140 or so species of rria w i cmiddot-~
or kissing bugs evolved from a re l ~
predator or assassin bug so i
because it kills its arthropod p ~ shy
injecting potent toxins and prOte shy
that liquefy the insides of the p rhey can be sucked out How shy
~~tf~ and anyone imeresred in narural ~ 2S ~= k lam Our-of-print back issues are ~ ii School Naturalist is senr free upon
~~-- 11d surf1ce mail overseas The Kansas -g opound12~e paid ar Emporia Kansas Address
oi Sciences Box 4050 Emporia Srare bull_ lt_ ~ hrne of rhe aurhors andor ediror and
= re icwed online ar wwwemporiaedu middot=-T tc-ixL lJ a ~) indexed in Wildlife Review
~ is an irregular publicarion issued
ri
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Biological
cil of Vermont She of Illinois
of hosts and
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Kissing Bugs in the United States INTRODUCTION
Kissing bugs are bloodsucking insects
in the order Hemiptera in the family
Reduviidae Almost all known species
reside in the New World
They are best known as vecrors
of Chagas disease a disease caused
by transmission of the blood parasite
Trypanosoma cruzi through their feces
Vecror transmission of Chagas disease
occurs with a number of endemic
kissing bugs Chagas is largely confined
to Mexico Central America and South
America
Although only a few cases of insectshy
transmined Chagas disease have occurred
in the US these bugs are common in
southern southeastern and southwestern
states
We discuss the biology of the
bugs their evolutionary hisrory their
distribution in the United States the risk
they pose for transmission of T cruzi ro
pets and other domestic animals and the
control of kissing bugs around human
habitations
EVOLUTION OF KISSING BUGS
The 140 or so species of rriaromines
or kissing bugs evolved from a reduviid
predaror or assassin bug so called
because it kills its arthropod prey by
injecting potent roxins and pro teases
that liquefy the insides of the prey so
they can be sucked our How many
times reduviids have forsaken their
assassin habitS ro become blood sucking
kissers has been a matter of some
controversy The fact that triaromines are
found in as diverse habitats as bird and
opossum nests (arboreal) and armadillo
burrows and rats nests as well as human
habitations (terrestrial) suggests that they
evolved from multiple different predatory
ancesrors (ie are polyphyletic) that
were already associated with arboreal or
terrestrial hosts
The twO mOSt important tribes for
human infection the Triaromini and
Rhodniini are quite divergent suggesting
distinct predatory ancestors However
a recent large study of the evolution of
the Reduviidae using multiple genes and
many morphological characters indicates
that these two tribes arose from the same
predatory ancestor (1) The remaining
three tribes still need to be examined
but so far it appears blood sucking
evolved just once ie the Triatominae are
monophyletic
Classification Hierarchy Rank Taxon
Kingdom Animalia
Phylum Arthropoda
Class Insecta
Order Hemiptera
Family Reduviidae
Subfamilies Triatominae
Rhodniini
Genera Triatoma
Paratriatoma
Figure 1 Triatoma sanguisuga the most common triatomine in Kansas and
throughout central and southeastern US Photo by Harold Baquet
Figure 2 Triatoma rubida common in foo (female note pointed projection from en L bull bull
One species Triatoma rubrofosciata
is found around the world and may have
spread in association with ships rats A few other species are found in Southeast
and East Asia however most species
(-125) are found only in the Americas
The Triatomini tribe contains two genera important for human transmission of
Chagas Triatoma and Panstrongyfus the former has the most (80) species and
includes the two most important vectors
for human transmission T inftstans
in South America and T dimidiata in
Mesoamerica (Mexico Central America and northern South America) The
Rhodniini tribe includes the genera
Rhodnius and Psammoestes with Rhodnius
profixus being an important human vector
in northern South America
KISSING BUG SPECIES IN THE UNITED STATES
Ten species of Triatominae (Table
1) are native to the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures Natural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to two genera (Paratriatoma and Triatoma)
and include four species complexes within the Triatoma (iecticularia phyllosoma
protracta rubida) The two species
reported in Kansas T sanguisuga and
T ecticularia are in an uncertain status
and lecticularia complex respectivel (Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reporred from poultry houses barns and houses
Although there are reports of serious
allergic reactions to their bite no T cruzi transmission to humans has been
reponed in Kansas Thatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida to Texas (4)
4
Figure 2 Triatoma rubida common in foothill regions of Phoenix and Tucson=-i ltc common triatomine in Kansas and
(female note pointed projection from end of the abdomen) ~~ _5 Photo by Harold Baquer
p rofixus being an importanr human vecror
in northern South America
- ~ I KISSING BUG SPECIES
IN THE UNITED STATES Ten species of Triarominae (Table
_imiddot~i4 1) are native ro the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures - - c ~atural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to twO genera (Paratriatoma and Ti-iatoma )
- 1=a and include four species complexes within
the Ti-iatoma (lecticularia phyllosoma - T- protracta rubida) The two specie
reported in Kansas T sanguisuga an ~
T fecticufaria are in an uncertain status
and lecticularia complex respectively
(Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reported
from poultry houses barns and houses
Although there are reportS of serious
allergic reactions to their bite no T cruz i transmission ro humans has been
reported in Kansas Triatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida ro Texas (4)
The other kissing bug reported from
Kansas T fecticufaria feeds on opossums
as well as wood rats (Neotoma spp) and the
rock squirrel (Spennophilus variegatus) T indictiva is reported to feed on domestic
and peridomestic animals (See Figures
2-5 for other common triatomines in the
US) All ten species in the United States
are considered sylva tic and have differenr
ecological niches Species in the leticularia
complex prefer open shrubland
woodland and wooded grassland (2)
Wooded grassland is the preferred habitat
for members of the protracta rubida and
phyllosoma complexes The one species
in the United States not in the Ti-iatoma
genus Pamtriatoma hirsuta has a broad
Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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MP
OR
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Pe
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Tri
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Kansas School Naturalist rSSN 0022-877X Published by EMPORIA STATE UNIVERSITY Editor JOHN RICHARD SCHROCK Editorial Comminee TOM EDDY BILL JENSEN MARSHALL SUNDBERG R BRENT THOMAS ERIC YANG Circulacion and Mailing ROGER FERGUSON Circulacion (chis issue) 10000 Press Run 15000 Media Designer John Decker Primed by McCormick Armscrong
The Kansas School Naturalist is sene free of charge and upon request ro reachers and anyone interesred in natueal history and nature educarion In-peine back issues are sem free as long as supply lasrs Our-of-prine back issues are sent for one dollar photocopy and poseagehandling charge per issue The Kansas School Naturalist is sent free upon requesr by media mail to all US zipcodes flrsr class to Mexico and Canada and surpound1cemail overseas The Kansas School Naturalist is published by Emporia Srare University Emporia Kansas Posrage paid ar Emporia Kansas Address all correspondence ro Ediror Kamas School Naturalist Department of Biological Sciences Box 4050 Emporia Srare University Emporia KS 66801-5087 Opinions and perspecrives expressed are rhose of rhe aurhors andor ediror and do not reflect the ofllcial posicion or endorsement of ES U Some issues can be viewed online at wwwemporiaedu ksn The Kansas School Naturalist is listed in Ulrichs International Periodicals Directory indexed in Wildlife Re-viewl Fisheries Review and appropriate issues are indexed in rhe Zoological Record The KSN is an irregulae publicaeion issued from one ro four rimes per year
Kissing Bugs in the INTRODUCTION
Kissing bugs are bloodsuc ing =-_
m the order Hemiptera in ( e
Reduviidae Almost all knOmiddot
reside in the New World
They are best known a5
of Chagas disease a di sease 3
by transmission of the blood shy
Trypanosoma cruzi through (heir ~~~-
Vector transmission of Chaga ~
occurs with a number of e-=- _
kissing bugs Chagas is largel ~ O - --~2
to Mexico Central America an~ ~ __ -
America
Although only a few case- 0 i _ shy
Front cover Tiiatoma gerstaeckeri common in sourhern and central Texas Phoro used with permission of Mike Quinn TexasEnronet
Justin O Schmidt has a PhD in Entomology from the University of Georgia He runs a non-profit organizarion Sourhwesrern Biological Insrirure in Tucson Arizona
Lori Stevens is Professor of Biology ar The Universiry of Vermont She received her MS and PhD degrees from rhe University of Illinois Chicago Srevens is intrigued wieh ehe generic inreractions of hosrs and parasites She enjoys visiring family in Leavenworch Kansas
Patricia Dom is a Professor of Biological Sciences ar Loyola University New Orleans She received her PhD from rhe University of Maryland and began her work on Trypanosomiais during her posr-docrora l fellowship ar Sranford Medical School
Mark Mosbacher received his doctoraee of vererinary medicine from the University of Missouri He currently practices emergency veterinary medicine in Phoenix Arizona
John K10tt has a PhD in Entomology from the Universi ty of Kansas and is an Urban Entomology Specialisr Emeritus ar the University of California Riverside and lives in Sedona Arizona
Stephen A K10tt graduared from rhe Universi ty of Kansas and is Chief of Infeceious Diseases ar rhe Universiry of Arizona
transmi ned Chagas disease hamiddote
in the US rhese bugs are corn-middot
southern southeastern and sourhmiddots- --
states
We discuss the biology 0 - ~
bugs their evolu rionary hiscof- - shy
distribution in the United Srare t
they pose for transmission of T shy
pets and other domestic animals 21 _ ~ _
control of kiSSing bugs around h --
habitarions
EVOLUTION OF KISSING BUGS
The 140 or so species of rria w i cmiddot-~
or kissing bugs evolved from a re l ~
predator or assassin bug so i
because it kills its arthropod p ~ shy
injecting potent toxins and prOte shy
that liquefy the insides of the p rhey can be sucked out How shy
~~tf~ and anyone imeresred in narural ~ 2S ~= k lam Our-of-print back issues are ~ ii School Naturalist is senr free upon
~~-- 11d surf1ce mail overseas The Kansas -g opound12~e paid ar Emporia Kansas Address
oi Sciences Box 4050 Emporia Srare bull_ lt_ ~ hrne of rhe aurhors andor ediror and
= re icwed online ar wwwemporiaedu middot=-T tc-ixL lJ a ~) indexed in Wildlife Review
~ is an irregular publicarion issued
ri
-middotnimiddotmiddotcrsinmiddot of
Biological
cil of Vermont She of Illinois
of hosts and
__~~ Lnimiry
--0-( [tcrions
- ~-Oh Lniersiry gt1 vrarvbnd
)()S-Jocrorai
~ t t Ka nsas
hmsas and is
Kissing Bugs in the United States INTRODUCTION
Kissing bugs are bloodsucking insects
in the order Hemiptera in the family
Reduviidae Almost all known species
reside in the New World
They are best known as vecrors
of Chagas disease a disease caused
by transmission of the blood parasite
Trypanosoma cruzi through their feces
Vecror transmission of Chagas disease
occurs with a number of endemic
kissing bugs Chagas is largely confined
to Mexico Central America and South
America
Although only a few cases of insectshy
transmined Chagas disease have occurred
in the US these bugs are common in
southern southeastern and southwestern
states
We discuss the biology of the
bugs their evolutionary hisrory their
distribution in the United States the risk
they pose for transmission of T cruzi ro
pets and other domestic animals and the
control of kissing bugs around human
habitations
EVOLUTION OF KISSING BUGS
The 140 or so species of rriaromines
or kissing bugs evolved from a reduviid
predaror or assassin bug so called
because it kills its arthropod prey by
injecting potent roxins and pro teases
that liquefy the insides of the prey so
they can be sucked our How many
times reduviids have forsaken their
assassin habitS ro become blood sucking
kissers has been a matter of some
controversy The fact that triaromines are
found in as diverse habitats as bird and
opossum nests (arboreal) and armadillo
burrows and rats nests as well as human
habitations (terrestrial) suggests that they
evolved from multiple different predatory
ancesrors (ie are polyphyletic) that
were already associated with arboreal or
terrestrial hosts
The twO mOSt important tribes for
human infection the Triaromini and
Rhodniini are quite divergent suggesting
distinct predatory ancestors However
a recent large study of the evolution of
the Reduviidae using multiple genes and
many morphological characters indicates
that these two tribes arose from the same
predatory ancestor (1) The remaining
three tribes still need to be examined
but so far it appears blood sucking
evolved just once ie the Triatominae are
monophyletic
Classification Hierarchy Rank Taxon
Kingdom Animalia
Phylum Arthropoda
Class Insecta
Order Hemiptera
Family Reduviidae
Subfamilies Triatominae
Rhodniini
Genera Triatoma
Paratriatoma
Figure 1 Triatoma sanguisuga the most common triatomine in Kansas and
throughout central and southeastern US Photo by Harold Baquet
Figure 2 Triatoma rubida common in foo (female note pointed projection from en L bull bull
One species Triatoma rubrofosciata
is found around the world and may have
spread in association with ships rats A few other species are found in Southeast
and East Asia however most species
(-125) are found only in the Americas
The Triatomini tribe contains two genera important for human transmission of
Chagas Triatoma and Panstrongyfus the former has the most (80) species and
includes the two most important vectors
for human transmission T inftstans
in South America and T dimidiata in
Mesoamerica (Mexico Central America and northern South America) The
Rhodniini tribe includes the genera
Rhodnius and Psammoestes with Rhodnius
profixus being an important human vector
in northern South America
KISSING BUG SPECIES IN THE UNITED STATES
Ten species of Triatominae (Table
1) are native to the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures Natural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to two genera (Paratriatoma and Triatoma)
and include four species complexes within the Triatoma (iecticularia phyllosoma
protracta rubida) The two species
reported in Kansas T sanguisuga and
T ecticularia are in an uncertain status
and lecticularia complex respectivel (Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reporred from poultry houses barns and houses
Although there are reports of serious
allergic reactions to their bite no T cruzi transmission to humans has been
reponed in Kansas Thatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida to Texas (4)
4
Figure 2 Triatoma rubida common in foothill regions of Phoenix and Tucson=-i ltc common triatomine in Kansas and
(female note pointed projection from end of the abdomen) ~~ _5 Photo by Harold Baquer
p rofixus being an importanr human vecror
in northern South America
- ~ I KISSING BUG SPECIES
IN THE UNITED STATES Ten species of Triarominae (Table
_imiddot~i4 1) are native ro the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures - - c ~atural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to twO genera (Paratriatoma and Ti-iatoma )
- 1=a and include four species complexes within
the Ti-iatoma (lecticularia phyllosoma - T- protracta rubida) The two specie
reported in Kansas T sanguisuga an ~
T fecticufaria are in an uncertain status
and lecticularia complex respectively
(Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reported
from poultry houses barns and houses
Although there are reportS of serious
allergic reactions to their bite no T cruz i transmission ro humans has been
reported in Kansas Triatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida ro Texas (4)
The other kissing bug reported from
Kansas T fecticufaria feeds on opossums
as well as wood rats (Neotoma spp) and the
rock squirrel (Spennophilus variegatus) T indictiva is reported to feed on domestic
and peridomestic animals (See Figures
2-5 for other common triatomines in the
US) All ten species in the United States
are considered sylva tic and have differenr
ecological niches Species in the leticularia
complex prefer open shrubland
woodland and wooded grassland (2)
Wooded grassland is the preferred habitat
for members of the protracta rubida and
phyllosoma complexes The one species
in the United States not in the Ti-iatoma
genus Pamtriatoma hirsuta has a broad
Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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~~tf~ and anyone imeresred in narural ~ 2S ~= k lam Our-of-print back issues are ~ ii School Naturalist is senr free upon
~~-- 11d surf1ce mail overseas The Kansas -g opound12~e paid ar Emporia Kansas Address
oi Sciences Box 4050 Emporia Srare bull_ lt_ ~ hrne of rhe aurhors andor ediror and
= re icwed online ar wwwemporiaedu middot=-T tc-ixL lJ a ~) indexed in Wildlife Review
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ri
-middotnimiddotmiddotcrsinmiddot of
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cil of Vermont She of Illinois
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--0-( [tcrions
- ~-Oh Lniersiry gt1 vrarvbnd
)()S-Jocrorai
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hmsas and is
Kissing Bugs in the United States INTRODUCTION
Kissing bugs are bloodsucking insects
in the order Hemiptera in the family
Reduviidae Almost all known species
reside in the New World
They are best known as vecrors
of Chagas disease a disease caused
by transmission of the blood parasite
Trypanosoma cruzi through their feces
Vecror transmission of Chagas disease
occurs with a number of endemic
kissing bugs Chagas is largely confined
to Mexico Central America and South
America
Although only a few cases of insectshy
transmined Chagas disease have occurred
in the US these bugs are common in
southern southeastern and southwestern
states
We discuss the biology of the
bugs their evolutionary hisrory their
distribution in the United States the risk
they pose for transmission of T cruzi ro
pets and other domestic animals and the
control of kissing bugs around human
habitations
EVOLUTION OF KISSING BUGS
The 140 or so species of rriaromines
or kissing bugs evolved from a reduviid
predaror or assassin bug so called
because it kills its arthropod prey by
injecting potent roxins and pro teases
that liquefy the insides of the prey so
they can be sucked our How many
times reduviids have forsaken their
assassin habitS ro become blood sucking
kissers has been a matter of some
controversy The fact that triaromines are
found in as diverse habitats as bird and
opossum nests (arboreal) and armadillo
burrows and rats nests as well as human
habitations (terrestrial) suggests that they
evolved from multiple different predatory
ancesrors (ie are polyphyletic) that
were already associated with arboreal or
terrestrial hosts
The twO mOSt important tribes for
human infection the Triaromini and
Rhodniini are quite divergent suggesting
distinct predatory ancestors However
a recent large study of the evolution of
the Reduviidae using multiple genes and
many morphological characters indicates
that these two tribes arose from the same
predatory ancestor (1) The remaining
three tribes still need to be examined
but so far it appears blood sucking
evolved just once ie the Triatominae are
monophyletic
Classification Hierarchy Rank Taxon
Kingdom Animalia
Phylum Arthropoda
Class Insecta
Order Hemiptera
Family Reduviidae
Subfamilies Triatominae
Rhodniini
Genera Triatoma
Paratriatoma
Figure 1 Triatoma sanguisuga the most common triatomine in Kansas and
throughout central and southeastern US Photo by Harold Baquet
Figure 2 Triatoma rubida common in foo (female note pointed projection from en L bull bull
One species Triatoma rubrofosciata
is found around the world and may have
spread in association with ships rats A few other species are found in Southeast
and East Asia however most species
(-125) are found only in the Americas
The Triatomini tribe contains two genera important for human transmission of
Chagas Triatoma and Panstrongyfus the former has the most (80) species and
includes the two most important vectors
for human transmission T inftstans
in South America and T dimidiata in
Mesoamerica (Mexico Central America and northern South America) The
Rhodniini tribe includes the genera
Rhodnius and Psammoestes with Rhodnius
profixus being an important human vector
in northern South America
KISSING BUG SPECIES IN THE UNITED STATES
Ten species of Triatominae (Table
1) are native to the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures Natural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to two genera (Paratriatoma and Triatoma)
and include four species complexes within the Triatoma (iecticularia phyllosoma
protracta rubida) The two species
reported in Kansas T sanguisuga and
T ecticularia are in an uncertain status
and lecticularia complex respectivel (Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reporred from poultry houses barns and houses
Although there are reports of serious
allergic reactions to their bite no T cruzi transmission to humans has been
reponed in Kansas Thatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida to Texas (4)
4
Figure 2 Triatoma rubida common in foothill regions of Phoenix and Tucson=-i ltc common triatomine in Kansas and
(female note pointed projection from end of the abdomen) ~~ _5 Photo by Harold Baquer
p rofixus being an importanr human vecror
in northern South America
- ~ I KISSING BUG SPECIES
IN THE UNITED STATES Ten species of Triarominae (Table
_imiddot~i4 1) are native ro the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures - - c ~atural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to twO genera (Paratriatoma and Ti-iatoma )
- 1=a and include four species complexes within
the Ti-iatoma (lecticularia phyllosoma - T- protracta rubida) The two specie
reported in Kansas T sanguisuga an ~
T fecticufaria are in an uncertain status
and lecticularia complex respectively
(Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reported
from poultry houses barns and houses
Although there are reportS of serious
allergic reactions to their bite no T cruz i transmission ro humans has been
reported in Kansas Triatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida ro Texas (4)
The other kissing bug reported from
Kansas T fecticufaria feeds on opossums
as well as wood rats (Neotoma spp) and the
rock squirrel (Spennophilus variegatus) T indictiva is reported to feed on domestic
and peridomestic animals (See Figures
2-5 for other common triatomines in the
US) All ten species in the United States
are considered sylva tic and have differenr
ecological niches Species in the leticularia
complex prefer open shrubland
woodland and wooded grassland (2)
Wooded grassland is the preferred habitat
for members of the protracta rubida and
phyllosoma complexes The one species
in the United States not in the Ti-iatoma
genus Pamtriatoma hirsuta has a broad
Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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Figure 1 Triatoma sanguisuga the most common triatomine in Kansas and
throughout central and southeastern US Photo by Harold Baquet
Figure 2 Triatoma rubida common in foo (female note pointed projection from en L bull bull
One species Triatoma rubrofosciata
is found around the world and may have
spread in association with ships rats A few other species are found in Southeast
and East Asia however most species
(-125) are found only in the Americas
The Triatomini tribe contains two genera important for human transmission of
Chagas Triatoma and Panstrongyfus the former has the most (80) species and
includes the two most important vectors
for human transmission T inftstans
in South America and T dimidiata in
Mesoamerica (Mexico Central America and northern South America) The
Rhodniini tribe includes the genera
Rhodnius and Psammoestes with Rhodnius
profixus being an important human vector
in northern South America
KISSING BUG SPECIES IN THE UNITED STATES
Ten species of Triatominae (Table
1) are native to the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures Natural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to two genera (Paratriatoma and Triatoma)
and include four species complexes within the Triatoma (iecticularia phyllosoma
protracta rubida) The two species
reported in Kansas T sanguisuga and
T ecticularia are in an uncertain status
and lecticularia complex respectivel (Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reporred from poultry houses barns and houses
Although there are reports of serious
allergic reactions to their bite no T cruzi transmission to humans has been
reponed in Kansas Thatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida to Texas (4)
4
Figure 2 Triatoma rubida common in foothill regions of Phoenix and Tucson=-i ltc common triatomine in Kansas and
(female note pointed projection from end of the abdomen) ~~ _5 Photo by Harold Baquer
p rofixus being an importanr human vecror
in northern South America
- ~ I KISSING BUG SPECIES
IN THE UNITED STATES Ten species of Triarominae (Table
_imiddot~i4 1) are native ro the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures - - c ~atural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to twO genera (Paratriatoma and Ti-iatoma )
- 1=a and include four species complexes within
the Ti-iatoma (lecticularia phyllosoma - T- protracta rubida) The two specie
reported in Kansas T sanguisuga an ~
T fecticufaria are in an uncertain status
and lecticularia complex respectively
(Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reported
from poultry houses barns and houses
Although there are reportS of serious
allergic reactions to their bite no T cruz i transmission ro humans has been
reported in Kansas Triatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida ro Texas (4)
The other kissing bug reported from
Kansas T fecticufaria feeds on opossums
as well as wood rats (Neotoma spp) and the
rock squirrel (Spennophilus variegatus) T indictiva is reported to feed on domestic
and peridomestic animals (See Figures
2-5 for other common triatomines in the
US) All ten species in the United States
are considered sylva tic and have differenr
ecological niches Species in the leticularia
complex prefer open shrubland
woodland and wooded grassland (2)
Wooded grassland is the preferred habitat
for members of the protracta rubida and
phyllosoma complexes The one species
in the United States not in the Ti-iatoma
genus Pamtriatoma hirsuta has a broad
Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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Figure 2 Triatoma rubida common in foothill regions of Phoenix and Tucson=-i ltc common triatomine in Kansas and
(female note pointed projection from end of the abdomen) ~~ _5 Photo by Harold Baquer
p rofixus being an importanr human vecror
in northern South America
- ~ I KISSING BUG SPECIES
IN THE UNITED STATES Ten species of Triarominae (Table
_imiddot~i4 1) are native ro the United States (2 3)
where their northern limit is determined
by intolerance of cold temperatures - - c ~atural infections of T cruzi have been
found in seven of these species (4) The
Triatominae in the United States belong
to twO genera (Paratriatoma and Ti-iatoma )
- 1=a and include four species complexes within
the Ti-iatoma (lecticularia phyllosoma - T- protracta rubida) The two specie
reported in Kansas T sanguisuga an ~
T fecticufaria are in an uncertain status
and lecticularia complex respectively
(Dorn unpublished data) They have the
widest geographical distribution of the
ten species found in the United States
The most widespread species T sanguisuga (Figure 1) and T fectufaria are
mostly sylvatic however in Manhattan
Kansas T sanguisuga was reported
from poultry houses barns and houses
Although there are reportS of serious
allergic reactions to their bite no T cruz i transmission ro humans has been
reported in Kansas Triatoma sanguisuga
is commonly associated with raccoons
and opossums and has been found in
large numbers around human dwellings
from Florida ro Texas (4)
The other kissing bug reported from
Kansas T fecticufaria feeds on opossums
as well as wood rats (Neotoma spp) and the
rock squirrel (Spennophilus variegatus) T indictiva is reported to feed on domestic
and peridomestic animals (See Figures
2-5 for other common triatomines in the
US) All ten species in the United States
are considered sylva tic and have differenr
ecological niches Species in the leticularia
complex prefer open shrubland
woodland and wooded grassland (2)
Wooded grassland is the preferred habitat
for members of the protracta rubida and
phyllosoma complexes The one species
in the United States not in the Ti-iatoma
genus Pamtriatoma hirsuta has a broad
Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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Table 1 Checklist and geographic distribution ofTriatominae of the United Stares
(Not including the cosmopolitan T rubrofoseiata)
Genera Species Species In Kansas States Distribution Complex reported from
Triatoma Uncertain T sanguisuga yes 23
Lectularia Tleeticularia yes 16
T indictiva 3
T inaassata 1
Phyllo soma T recurva 6
T gerstaeckeri 2
Protracta T protroeta 7
T neatamae 1
Rubida T rubida 6
Parotriatoma P hirsuta 5
Alabama Arizona Arkansas Florid a Georgia Illinois Indiana Kansas Kentucky Louisiana MarylandMi ssiss ippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma PennsylvaniaS Carolina Tennessee Texas Virginia
Arizona California Florida Georgia Illinois Kansas Louisiana Maryland Missouri New Mexico N Carolina Oklahoma PennsylvaniaS Carolina Tennessee Texas
Arizona New MexicoTexas
Arizona
Arizona CaliforniaColorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
Arizona California Colorado NevadaNew Mexico
potential niche much broader than its
current geographical distribution
BIOLOGYAND NATURAL HISTORY OF KISSING BUGS
Kissing bugs have a gradual
development with five immature instars
appearing similar to adults but without
wings They are large bugs with adults
ranging in length from about 12 to 36
mm Both sexes and all five instars feed on
blood ofvertebrates starting with the first
instars which take a blood meal as soon
as 2-3 days after hatching from the egg
They can develop from egg (Figure 5) to
adult in several months in warm tropical
areas when food is readily available In
temperate regions having extended cooler
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live
several months occasionally a yea r or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protraeta or T reeurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared ro other hemipterans As with
other hemipterans females readily mate
with multiple males females can srore
sperm for use months later and over their
life span can lay up to several hundred
eggs depending upon conditions (5)
6
Active dispersal is achieved by flight in
adults and by crawling in immatures (6 )
Passive dispersal occurs in some species
by transport of attached eggs to ne
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs to crawl
toward locations potentially having new
hosts Adults typically disperse by fli gh r
around dusk Nutritional s tate and
hunger are powerful drivers of dispersal
(7) Another is the classical reproducti e
strategy in which gravid females retain
eggs for deposition after flight (8 9)
Dispersal is not a primary form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of femal es
and males captured at lights and the
rapidity with which captured femal es
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
Triatomine species inhabit a variety ot
ecological environments from generalize
to highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a
limited number of host types Specialists
include Cavernieola pilosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammolestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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3 ri bution ofTriatominae of the United States - I I ) -1roiflscUlta
$tates oprtedfrom
I Distribution
Alabama Arizona Arkansas Florida Georgia Illinois Indiana Kansas Kentucky louisiana Maryland Mississippi Missouri New Jersey New Mexico N Carolina Ohio Oklahoma Pennsylvania S Carolina Tennessee Texas Virginia
ArizonaCalifornia Florida Georgia Illinois Kansas louisiana Maryland Missouri New Mexico NCarolinaOklahoma Pennsylvania SCarolina TennesseeTexas
Arizona New MexicoTexas
Arizona
ArizonaGJlifornia Colorado Nevada New Mexico Texas
New Mexico Texas
Arizona California Colorado Nevada New Mexico Texas Utah
Texas
Arizona California Colorado Nevada New Mexico Texas
ArizonaCalifornia Colorado Nevada New Mexico
periods of the year the bugs are typically
univoltine with only one generation per
year In these regions bugs seek refuge in
rodent burrows cavities caves and other
locations with moderate temperatures and
pass the unfavorable times as immatures
in various stages (Figure 4) Adults live everal months occasionally a year or
more (5) and feed throughout their
lives In captivity adults of Triatoma
rubida T protracta or T recurva live 4-5
months on average Mating lasts only
about 10 minutes a relatively short time
compared to other hemipterans As with
other hemipterans females readily mate
wi th multiple males females can stOre
sperm for use months later and over the [ ~
life span can lay up to several hundrec
eggs depending upon conditions (S
Active dispersal is achieved by flight in
adults and by crawling in immatures (6)
Passive dispersal occurs in some species
by transport of attached eggs to new
locations by birds or other animals or by
human transport of materials harboring
eggs or free-living stages Active dispersal
of immatures can occur when a shortage
of food sources stimulates bugs ro crawl
roward locations potentially having new
hosts Adults typically disperse by flight
around dusk Nutritional state and
hunger are powerful drivers of dispersal
(7) Another is the classical reproductive
strategy in which gravid females retain
eggs for deposition after flight (89)
Dispersal is not a primaly form of mate
seeking as in many moths and other
insects Evidence against mate seeking
is indicated by equal numbers of females
and males captured at lights and the
rapidity with which captured females
oviposit fertile eggs (8)
HABITAT AND HOST ASSOCIATION
TriatOmine species inhabit a variety of
ecological environmentS from generalized
ro highly specialized including human
domestic environments Within these
environments they can range from
opportunists feeding on a variety of
host species to specialists feeding on a limited number of host types Specialists
include Cavernicola piLosa which feeds
on bats Rhodnius spp living in palm
trees where they feed on inhabiting birds
and mammals and Psammofestes spp
which live in bird nests constructed of
rwigs Bird blood meals are sufficient
for the bugs to survive and reproduce
but birds are not hosts of T cruzi and
thus alter the parasite transmission cycle
Somewhat more general feeders are T rubrofasciata and members of the T
protracta complex that prefer feeding on
rodents Triatoma infestans T dimidiata
and R profixus are inhabitants of human
domestic environments where they feed
on people and their animals (6) Most
species irrespective of their preferred
habitat can become opportunistic feeders
when other species are available Species
that are generally opportunists can also
be facultative specialists preferring to feed
mainly on one group of hosts if given the
opportunity (10) The Triatoma living
in Arizona are all non-domestic species
that often feed on packrats (Neotoma
spp) yet they readily feed on a variety of
vertebrate hosts they encounter including
humans The Arizona-Sonora Desert
Museum in Tucson Arizona is a regional
educational living zoo that inadvertently
supports large populations of T rubida
and T protracta These bugs plague many
museum animals and sometimes feed
so extensively that they have caused the
death by chronic blood loss of animals
including a rare protected mountain
rattlesnake (James L Jarchow DVM
pers commun) In Kansas Triatoma
sanguisuga like the Arizona species feeds
predominantly on Neotoma raccoons and
opossums but will readily feed on a
variety of other hosts including humans
When and how some species of
triaromines became specialists on humans
is a mystery The subfamily is believed
to have originated about 5 million years
Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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Figure 4 A fifth instar (nymphal stage) of T recurva Note the absence of figure 6 A teneral of T rubida i shy _
wings in the nymphs
8
- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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- shy --t o-ader in California especially
middotmiddotmiddotomiddotva Note the absence of Figure 6 A reneral of T rubida ienewly emerged adult
bull 9
ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
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he U
S (f
emal
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ago in the New World (11) Dates for
the arrival of human beings in the New
World are debated but their estimated
arrival sometime between 14000 to
20000 years ago is relatively recent
Thus shifts in some species of kissing
bugs to specializing on our species are
recent evolutionary events and might
well explain some of our vulnerability to
the T cruzi parasite the bugs transmit to
us
FEEDING BEHAVIOR Kissing bugs rely on multiple sensory
systems to locate hosts These include
gradients of carbon dioxide in the air
odors moisture heat and air flow (12)
They are exquisitely sensitive to carbon
dioxide exhaled by mammals when
breathing being able to detect 75 parts
per million above the background air
levels of 350 parts per million when
a trace of exhaled lactic acid another
attractant is present Cold blooded hosts
including reptiles generate very low levels
of carbon dioxide nevertheless kissing
bugs use many other host-generated
chemical cues including short chained
ali phatic amines and acids 7- to 9-carbon
aldehydes and alcohols and lactic acid to
locate hosts The bugs can also detect
infrared radiation enabling them to orient
toward warm blooded hosts from several
meters Finally they anemotactically
follow the movement of air currents
impregnated with host odors to locate
the host (12) Feeding by bugs in human
domiciles usually takes place at night
with bugs hiding during the day in
cracks and crevices away from the feeding
areas Much less is known about feeding
times for sylvatic (non-domestic) species
Likely they adjust their activity periods to
match the inactive periods of their host
thereby minimizing danger from host
defenses Triatoma rubida and T protracta
do not hesitate to approach and feed on
rodents during bright midday times (13)
a behavior that might well be adaptive for
bugs that often feed on rodents in their
burrows while the rodents are sleeping or
quiescent during the day
Once a host is located a hungry
bug extends its proboscis and inserts it
into the skin of the host Sometimes the
host detects the minor sensation caused
by a bugs probing and moves or shifts
posmon In response the bug usually
retracts somewhat before continuing
probing and feeding Considering that
a bug often imbibes more blood than
its own weight (3) feeding is relatively
rapid lasting on average 22 minutes
for T protracta 28 for T rubida and
between 11 - 28 minutes for T infestans
T dimidiata and R proixus (1 3 14)
Blood is mostly water and the bulkiness
of a large blood meal limits a bugs
movement To eliminate excess water
bugs often defecate during feeding at
completion of feeding or shortly after
leaving the feeding site (Figure 8) Some
species including those domestic species
that frequemly transmit Chagas disease
defecate rapidly and frequently often on
or near the host (14) while others defecate
less rapidly and less frequently often at a
distance from the host (3) Frequency
and proximity of defecation to the host
are considered to be critical factors in
transmitting T cruzi and Chagas
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are th - common cause of insect bites that - -_
in anaphylaxis in the United State --
medical emergency (also causeu - - ~
commonly by bee stings) results -ro ~- =
release of chemical signals in resP Qclt ~
proteins in the bugs saliva ina b _ _ usually causes the individual to r s the emergency room where epineF~~gt _
and other imerventions reverse th - _
blood pressure swollen ai[-as ~=shy
rashes that often accompany anapn- c
Victims usually are awakened at nig shy
shortness of breath difficulty b re-l ~- ~
or generalized itching Othe~ _c ominous allergies include hives se ~
of the eyes swelling at the site 0 - - -t
bite and persistent itching Patiem -_
experience anaphylaxis when re-bi ~-shy
and one adult woman has died ~C _-shythe condition The bite itself is pai
however swelling and inflammatior -shy
the site of the bite may last for middoteeL
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit CI ~
to humans in the United States lt =-__ shy
six such cases have been docunei
Infection with Trypanasoma en is common in several wildlife spe - j -
rare in domestic pets Reported Cd5eS __
10
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
~--l vluch less is known about feeding
-16 for sylva tic (non-domestic) species
~~Jd~ they adjust their activity periods to
- ch the inactive periods of their host
erltbmiddot minimizing danger from host
erenscs Triatoma rubida and T protracta
_ ~Of hesitate to approach and feed on
rienrs during bright midday times (13)
chlior that might well be adaptive for
_g [hat often feed on rodents in their
transmuting T cruzi and Chagas disease
to humans (Figure 9)
HUMAN BITE ALLERGIES FROM KISSING BUGS
Kissing bug bites are the most
common cause of insect bites that result
in anaphylaxis in the United States This
medical emergency (also caused more
commonly by bee stings) results from the
domestic species
the warmer southern stares There are no
reportS of livestock horses or pigs being
infected Only one case report exists of
infection in a domestic cat in the US
The primary wildlife species in the
United Srates that serve as host reservoirs
for T cruz i are
packrats in the genus Neotoma
and opossums ~ _ -ns while the rodents are sleeping or
-e-cnt during the day
nee a host is located a hungry
extends its proboscis and inserts it
he skin of the host Sometimes the
erects the minor sensation caused
s- Sugs probing and moves or shifts
xsiion In response the bug usually
- = ~-rS somewhat before continuing
iing and feeding Considering that
t~~ often imbibes more blood than
-1 weight (3) feeding is relatively
~i c las ting on average 22 minutes
- ~ ~- porracta 28 for T rubida and
11 - 28 minutes for T infestans
-
3w i
d diara and R prolixus (13 14)
moscly water and the bulkiness
l rge blood meal limits a bugs
bull O1em To eliminate excess water
0 o tten defecate during feeding at
_ -c~E io1 of feeding or shortly after
-T-i ~ he feeding site (Figure 8) Some
~ _-~~ i1Cluding those domestic species
-~-- E~lIently transmit Chagas disease
i-_ ~ ~ rap idly and frequently often on
bullbull -~ ile hos t (14) while others defecate
_ 2-d l and less frequently often at a
_ lt~ ~ r-om the host (3) Frequency
- E - ~igtxi ni( of defecation to the hosr
_~ odered to be critical factors in
release of chemical signals in response to
proteins in the bugs saliva Anaphylaxis
usually causes the individual to rush to
the emergency room where epinephrine
and other interventions reverse the low
blood pressure swollen airways and
rashes that often accompany anaphylaxis
Victims usually are awakened at night by
shortness of breath difficulty breathing
or generalized itching Other less
ominous allergies include hives swelling
of rhe eyes swelling at the site of the
bite and persistent itching Patients may
experience anaphylaxis when re-bitten
and one adult woman has died from
the condition The bite itself is painless
however swelling and inflammation at
the site of the bite may last for weeks
WILD AND DOMESTICATED ANIMALS INFECTED WITH Trypanosoma cruzi IN THE UNITED STATES
Kissing bugs rarely transmit Chagas
to humans in the United States only
six such cases have been documented
Infection with Trypanasoma cruz i is
common in several wildlife species but
rare in domestic pets Reporred cases in
are limited ro dogs in
rodents particularly
raccoons
and to a lesser degree
armadillos and skunks T cruzi has been
reporred in more than 20 different wildlife
species in the United States in at least 13 states (15) Wood rats and other rodent
species serve as reservoirs of infection
as rodent dens and nests may harbor
the insect vectors Infection dynamics of
T cruz i isolates appear to vary between
species wirh raccoons being infected wirh
a more virulent genetic strain in which
blood parasites levels peak sooner and
antibodies are more quickly derected rhan
occurs in opossums anorher primary
wildlife reservoir species Additionally
mice injecred with opossum derived T cruzi isolates appear to subsequently
clear the infection and survive whereas
injecting mice with the raccoon derived
strains resulted in a 75 mortality rate
in one study Despite differences in
virulence between the host species strains
pathologic cardiac changes induced by T cruzi have been observed at necropsy in
both raccoons and opossums
Prevalence of T cruz i in wildlife
species varies but infection rates as high
as 62 occurred in raccoons tested
in various states (16) Infection rates
appear to be generally highest in coastal
locations with the elevated rates assumed
11
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
T cruzi exposure in many southern
states and as far north as Minnesota
In domestic dogs as in wildlife species
infection is thought ro be primarily by
First and foremost is a thorough
inspecrion of the home both inside and
outside
During the active season (around
bull Keep window curtalins and blirci ~-
Seal potential entryways into ho es points of entry) with caulk silicone sea
Figure 7 Blood smear showing
Trypanosoma cruzi parasites among
red blood cells Phoro by Rose
Nasrazadani
ro be due ro the increased disrribution
of insect vectors secondary ro the high
humidiry and lack of a winter freeze
along the sourhern and eastern coasts
In contrast ro the Latin American
isolates that are transmitted ro humans
indirectly at the insect bite location by
fecal contamination T cruzi in wildlife
species in the US is transmitted by
oral ingestion of infected insects and ro
a lesser degree by placental transmission
in utero
In domestic pets T cruzi causing
clinical health issues has been reported
primarily 111 working dogs from
southeastern Texas and foci in Louisiana
although antibody testing has confirmed
eating infected kissing bug vecrors in
response ro skin irritation from the insect
bite and from in utero transmission from
dam ro pup
Clinical signs of infection in dogs are
primarily cardiac recognized as exercise
intolerance or generalized weakness
Diagnostic testing may reveal severe heart
arrythmias And ultimately a dilated
form of heart failure develops Treatment
rarely results in a cure and survival time
in clinically affecred dogs ranges from 0
ro GO months
Prevention of Chagas disease in
dogs depends on insect vecror control
and limiting contacr with host reservoir
species In endemic kissing bug areas
animal housing and bedding should be
treated with insecticide monthly and
kennels improved as needed ro remove
insect nesting sites
PEST MANAGEMENT OF KISSING BUGS
Insecticide spraying is the cornersrone
ofvector control programs for triatomines
in South America where domestic and
peridomestic species are prevalent and
can attain high population densities in
and around homes The sylva tic species
found in the US are only occasional
invaders and thus present a different set
of problems that rely on non-chemical
measures for control such as habitat
modification and exclusion techniques
(see table below)
bull amp
Figure 8 (above) Fecal smears
and drops from a female T rubida
Table 2 Non-chemical measure w ~c _ triaromines
Sanitation measures bull Reduce clutter (eg clothes ~ _
bedroom to reduce potentia - - ~ - - - - bull Manage vegetation around t nE - ~ ~ - =shy
firewood and debris) that n1 a -shy
Install weather stripping and tight fi insect proof dog and cat entrances a
Manage indoor and outdoor lighting a ~ - - ~
kissing bugs bull Move lights away from doors bull Replace outside white ligh 5 gt- _ _
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
_ - ~ ntected kissing bug vecmrs in
-~- ro sk in irriration from the insect
- ~ ~ from iii IItero transmission from
2~ __ bull gt ult
C gtal signs of infection in dogs are
- --- shy cardiac recognized as exercise
ltOk 1 or generalized weakness
J~~ -i testing may reveal severe heart
--~ --5 nd ultimately a dilated
t-he-arr failure develops Treatment
-L ~~- ~esul r5 in a cure and survival time
dir alh affected dogs ranges from 0
-1 onths
_~ecmion of Chagas disease in
It7 c epends on insect vector control
-c ~ -i(in g conract with host reservoir
- ifO In endemic kissing bug areas
~Ad fi ousing and bedding should be
lt-~1 2 - irh insecticide monthly and
D --u~-- improved as needed to remove
-51 - s ring sites
bull
Figure 8 (above) Fecal smears
and drops from a fem ale T rubida
PEST Mfu~AGEMENT OF KISSING BUGS
gt xri ide spraying is the cornersmne
middot C T io nrrol programs for triaromines
~ il merica where domestic and
- --l(1c-gt[ ic species are prevalel1( and
bull- 1il high population densities in
1d aol 1d homes The sylvatic species
~c n [he CS are only occasional
- middot~c dnd thus presel1( a differel1( set
- ~ -tiems that rely on non-chemical
- ~~=-gt fo r control such as habitat
~Cil[ion and exclusion techniques
- dow)
- and foremost is a thorough
-- - - ~ion of the home both inside and
ring the active season (around
Table 2 Non-chemical measures ro reduce risk of household infestation by
triaromines
Sanitation measures bull Reduce clutter (eg clothes piles of paper) inside homes particularly in the
bedroom to reduce potential hiding places for kiSSing bugs bull Manage vegetation around the home and eliminate clutter (eg piles of lumber
firewood and debris) that may prOvide small animal habitat
Install weather stripping and tight fitting insect screens on windows and doors insect proof dog and cat entrances and keep fireplace flues shut
Manage indoor and outdoor lighting at night to make homes less attractive to kissing bugs
bull Move lights away from doors or windows where they may attract insects bull Replace outside white lights with yellow lights bull Keep window curtains and blinds drawn in lighted rooms
I
Seal potential entryways into homes (eg foundation cracks and utility line points of entry) with caulk silicone seal or other appropriate materials
I
13
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
mid-spring to mid-fall) inspections
should be carried our on a regular basis to
find and destroy any stray bugs During
the day they typically hide in dark places
in cracks and crevices or under objects
For example in the bedroom they may
hide in bed sheets or blankets or under
the mattress so it is recommended that
all of these potential hiding places be
inspected especially before retmng
Outside during the day they hide in
dark sheltered places such as beneath
flowerpots or furniture emerging at night
to be seen crawling or resting on surfaces
Pay special attention to areas inside and
out where pets congregate or sleep
In addition to efforts directly aimed
at kissing bugs concrol measures may be
needed to remove their wild vertebrate
hosts sllch as packrats which are common
around homes in the Southwest Packrats
can have several dozen or more kissing
bugs inhabiting their nests Trapping the
packrats and removing their nests close
to the house can potentially reduce home
invasions However it is advisable that
only the nearest packrat nest and packrats
be removed in the hope that the kissing
bugs will move to more peripheral nests
farther from the house After the packrats
have been removed a pyrethroid dust
or spray should be applied to the old
nesting sites to eliminate any remaining
kiSSing bugs This is critical because in the
absence of their natural host the bugs will
seek out another source of blood which
might end up being the homeowner (I 7)
In the case of sensitized homeowners
who have become allergic to kissing bugs
a directed spray application of pyrethroid
insecticides in and aro und the home
may provide some added protection
Most effective indoors is a crack and
crevice application to potential harborage
sites in bedrooms and bathrooms and
outdoors a perimeter treatment along
the foundation and eaves paying close
attention to thoroughly treat entryways
such as windows and doors to prevent
their access into the home
Sensitized individuals may also elect
to sleep under a bed net (mosquito
netting) that is tucked in all around
the mattress for added protection and
using double-sided sticky tape placed
on the legs of the bed Beds should be
kept at least one foor away from walls
Sticky trap monitors placed under and
around beds may also catch wandering
bugs Finally homeowners might wish
to thoroughly vacuum the bedroom and
consider isolating their bedroom during
the active season by weather-stripping
doors and keeping doors closed
Kissing bugs are medically important
fascinating insects that likely will be
interacting with humans and their pets
wi th increasing frequency in response
to warming climates and as more homes
encroach into their habitats Learning to
recognize these bugs and to keep them
out of human habitations is an important
responsibility of homeownership
14
REFERENCES 1 Weiruch C Munro fB
phylogeny of rhe as middot3i - (Hemi prera Reduviidae shymirochondrial genes dol P~ - ~- shy
Evol 2009 53 287-299 2 Iba rra-Cerdena CN San ~le= - _~
V Townsend Pererso n A Rl middot
(2009) Ecology of -or L _
Triarominae Acra Trop 1 10 - - bull
3 Zeled6n R Beard CB Pio _ 0
Leiby DA Dorn PL R J (in press) An appraisal r ~ --=shyChagas disease in rhe L-n re-C =- ~
4 de la Rua N Srevens (201l) High generic dit _ - 1- _ ~
popularion of (Le Com e 1855) i n ter~ - ~
mirochondrial markef5 Co ~ -
and 16S ribosomal D- -co
Generics and Evolurion 5 Goda D E and C J ~ _
Popularion dynamics of Ii
under narural climaric middot0 shy
rhe Argentin e C haco -EM ~ ~ shy3 179-94
6 Schofield C J 1988 Bios [- c rhe Triarominae In Bio5middotreoi H aemarophagous Insecrs 1 - __
ed) Sys remarics Associario r _ - Volume No 37 pp 284middot31 c -~ shy
Press O xford 7 Lehane M ] P K Mch --
Whiraker and C J Schotleld ] 4 ~ _
role of remperarure and nurririofi- _ _ in flighr iniriarion by 7i-iatam 1-~ _
Acta Trop 52 27-38 8 Klorz J H P L Dorn J L _ --
L Stevens J L Pinnas J 0- ~ l-r _
and S A Klotz 2010 Ki55ill ~ - __ ~
porential disease vecrors and (~ U
anaphylaxiS Cfin In Dis ) o 6 _ - 9 McEwen P K and M ] Lehl _- shy
Relationship between Highr 2
and oviposir ion in TiaOIgtII - - - shy
(Klug) (Hem Reduviidae - shy
Entomo 117 217middot23
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
u~ Je5 in and around the home
--_2 rovide some added protecrion
-- efrecrin~ indoors is a crack and
_c lt application to potenrial harborage
~-olt r bedrooms and bathrooms and
-or a perimeter rreatmenr along
- -()ll1dation and eaves paying close
~(o n ro thoroughly treat enrryways
~~ - d windows and doors ro prevenr
~ laquo(e55 inro the home
S1si tized individuals may also elect
lteF under a bed net (mosquiro
- ~Jng that is rucked in all around
c -rress for added protection and
_ sect do uble-sided sticky tape placed
Ci - he legs of the bed Beds should be
r ~ least one foor away from walls
rap monirors placed under and
- - ~ -_ bds may also catch wandering
_~ _ Finally- homeowners mighr wish
-uroughh- vacuum rhe bedroom and
___id ~ iolaring rheir bedroom during
d~ 2Cl i season by wearher-srripping
nd keeping doors dosed - lis~ lg bugs are medically imporranr
is -_2ing insecrs rhar likely will be
- _~~~ i rh humans and rheir pers
_1reas ing frequency in response
-ing c1imares and as more homes
-~ h inro their habirars Learning ro
sect-izemiddot these bugs and (Q keep rhem
il-lnan habirarions is an imporranr
- bili r of homeownership
REFERENCES I Weiruch C Munro JB Molecular
phylogeny of the assassin bugs (Hemiptera Reduviidae) based on mirochondrial genes Mol Phylogenet Evo 2009 53 287-299
2 Ibarra-Cerdena CN Sanchez-Cordero
V Townsend Peterson A Ramsey JM (2009) Ecology of North American
Triarominae Acta Trop 110 178-186 3 Zeledon R Beard CB Pinro Dias JC
Leiby DA Dorn PL Rodrigues Coura J (in press) An appraisal of the sraws of Chagas disease in the United States
4 de la Rua N Stevens L Dorn PL (2011) High genetic diversity in a si ngle
population of Tiiatoma sanguisuga
(Le Conte 1855) inferred from rwo mitochondrial markers cytochrome b and 16S ribosomal DNA Infection Genetics and Evolution
5 Goda D E and C J Schofield 1989 Population dynam ics of Ti-iatoma inmans under nawral climatic conditions in
the Argentine Chaco Med Vet Entomol
3 179-94 6 Schofield C J 1988 Biosystematics of
the Triatominae In Biosystematics of Haematophagous Insects (M W Service ed) Systematics Association Special
Volume No 37 pp 284-312 Clarendon
Press Oxford 7 Lehane M J P K McEwen C J
Whitaker and C J Schofield 1992 The role of temperature and nutritional status in flight initiation by Triatoma infestans Acta Timiddotop 52 27-38
8 KJotz J H P L Dorn J L Logan L Stevens J L Pinnas J O Schmidt and S A KJotz 20 IO Kissing Bugs potential disease vectors and cause of anaphylaxis CLin In Dis 50 1629-34
9 McEwen P K and M J Lehane 1994 Relationship berween flight initiation and oviposition in Triatoma infestam (KJug) (Hem Reduviidae) j App
EntomoL 11 7 217-23
10 Gurtler R E L A Ceballos P OrdoZezshy
Krasnowski L A Lanati R Stariolo and
U Kitron 2009 Strong host-feeding
preferences of the vector Tiiatoma
infestans modified by vector density
implications for the epidemiology of
Chagas disease PLGS Neglect Trop Dis
3e447
11 Schofield C J and C Galvno 2009
Classification evolution and species
groups within the Triatominae Acta
Tropl1088-100
12 Guerenstein P G and C R Lazzari
2009 Hosr-seeking how rriatomines
acqui re and make use of information to
find blood Acta Ti-op 110148-583
13 KJotz S A P L Dorn J H KJOtz
] L Pinnas C Weirauch J R Kurtz
and J Schmidt 2009 Feeding behavior
of triaromines from the southwestern
United States an update on potential
risk for transmission of Chagas disease
Acta Trop III114-18
14 Zeled6n R R Alvarado and L F Jiron
1977 Observations on the feeding and
defecation patterns of three triatol11ine
species (Hemiptera Reduviidae) Acta
Timiddotop3465-77
15 Barr Sc Canine C hagas Disease
(American Trypanosomiasis) in North
America Vet CLin Small Animal 39
(2009) 1055-1064
16 Yabsley MJ and NobletGP Seroprevalence
of Trypanosoma cruz i in raccoo ns from
South Carolina and Georgia J WiLdl Dis
38 (2002) 75-83
17 Gouge DH Stoltman AJ Snyder
J L Olson C How to Bug Proof Your
Home http ca ls arizona edupubs
insectsaz l 3201
15
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)
e E
MP
OR
IA S
TATE
UN
IVE
RS
ITYr
u KA
NSA
S SC
HO
OL
NAT
UR
ALIS
T B
OX
4050
12
00 C
OM
MER
CIA
L ST
E
mpo
ria
KS 6
6801
-508
7
ADDR
ESS
SERV
ICE
REQ
UEST
ED
No
np
rofi
t O
rg
US
Pos
tage
PA
ID
Em
pori
a K
ansa
s
66
80
1-5
08
7
Pe
rmit
No
203
Tri
atom
a re
curv
a t
he l
arge
st t
riat
ome
in t
he U
S (f
emal
e)