Review
Special Issue: Zoonoses of people and pets in the USA
Hookworms of dogs and cats as agents
of cutaneous larva migrans
Dwight D. Bowman1, Susan P. Montgomery2, Anne M. Zajac3, Mark L. Eberhard2 and
Kevin R. Kazacos4
1
Department of Microbiology and Immunology, College of Veterinary Medicine, Cornell University, Campus Road, Ithaca, NY
14850, USA
2
Division of Parasitic Diseases, Centers for Disease Control and Prevention, 4770 Buford Hwy, MS F-22, Atlanta, GA 30341, USA
3
Department of Biomedical Science and Pathobiology, Virginia-Maryland Regional College of Veterinary Medicine, Virginia Tech,
Blacksburg, VA 24061, USA
4
Department of Comparative Pathobiology, School of Veterinary Medicine, Purdue University, West Lafayette, IN 47907, USA
Dogs and cats are hosts to hookworms that may cause
zoonotic disease, most notably, cutaneous larva
migrans. Ancylostoma braziliense is most often impli-
cated in dermatological lesions, and Ancylostoma cani-
num has been associated with eosinophilic enteritis and
suggested as a possible cause of diffuse unilateral sub-
acute neuroretinitis in humans. Other manifestations
include eosinophilic pneumonitis, localized myositis,
folliculitis, erythema multiforme, or ophthalmological
manifestations. Ancylostoma eggs are morphologically
indistinguishable, which complicates epidemiological
studies. Surveys of dermatologists, gastroenterologists,
and ophthalmologists would help to define the inci-
dence of these zoonotic infections. Improved diagnostic
tests are needed to identify the causative species
involved and understand the epidemiology of hook-
worm disease. This review describes the discovery of
the disease, the biology of the agents, and how that
biology may impact disease.
Dog and cat hookworms and zoonotic disease
The common hookworms of dogs, Canis familiaris, are:
Ancylostoma caninum, A. braziliense, A. ceylanicum and
Uncinaria stenocephala; and of cats, Felis catus, are: Ancy-
lostoma tubaeforme, A. braziliense, A. ceylanicum and U.
stenocephala [1]. Larval infection in humans can lead to
hookworm associated cutaneous larva migrans (CLM) [1]
and, less commonly, eosinophilic pneumonitis [2–5], loca-
lized myositis [6,7], folliculitis [8–12] erythema multiforme
[13] or ophthalmological manifestations [14]. Of these
species, only A. ceylanicum will readily develop to adults
in people, but A. caninum has, on occasion, reached adult-
hood in people, causing eosinophilic enteritis [1,15].
The geographical distribution of each species is imper-
fectly known because ranges overlap, hosts can harbor
several species simultaneously, and only the larger U.
stenocephala egg can be morphometrically distinguished
[16]. Uncinaria stenocephala occurs in dogs and cats in the
colder climates of North America, South America, Europe,
Asia, Australia and New Zealand [17,18]. Recently, mol-
Corresponding author: Bowman, D.D. (ddb3@cornell.edu)
162 1471-4922/$ – see front matter ß 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.pt.2010.01.005 Available online 25 February 2010
ecular methods were developed to distinguish Ancylostoma
eggs in feces [19,20], but most older surveys were based on
necropsy results. A. caninum and A. tubaeforme, the most
common species, are found throughout the warmer ranges
of dogs and cats [1,19,20]. Ancylostoma ceylanicum is found
in Asia (India, Taiwan, Thailand, Malaysia, Borneo,
Indonesia), Australia and South America (Surinam)
Glossary of terms
Bulla: a large blister (plural – bullae). Some bullae associated with CLM have
been several inches in diameter.
Colitis: inflammation of the large intestine
Enteritis: inflammation of the intestine, usually the small intestine.
Eosinophilia: increase in the type of white blood cells called eosinophils in the
peripheral blood. Such increases often accompany infections with helminths
and arthropods. The numbers in the peripheral blood will also often increase in
allergic reactions.
Folliculitis: inflammation of hair follicles
Gasterophilus: This is a genus of flies, called botflies, which lay eggs on the
hairs of horses (and a few other animals). The adults are free living, but the
larvae, the maggots, live and develop only in the stomach of the horse.
Hookworm: These are small nematode parasites of the intestinal tract of
vertebrates; hookworms suck blood to various degrees, depending on the
species. They are called hookworms because the anterior end is bent
backwards giving the worm a ‘hooked’ appearance.
Myiasis: Infection with maggots. The term maggot is typically reserved for the
larvae of flies, more specifically dipteran flies lacking a distinct head capsule.
Apart from the Gasterophilus mentioned in the text, other genera of maggots
such as Hypoderma of cattle have also been responsible for serpiginous track
lesions in the skin of people.
Myositis: Inflammation of the muscle.
Papule/papular: Small, solid nodule in the skin, which may be intensely
pruritic.
Paratenic host: This is a host in which a parasite can persist, without major
developmental change, but which is not required for the life cycle to be
completed. In the case of A. caninum, the larvae can infect and persist in
cockroaches. They can also persist in the muscles of small rodents following
infection by skin penetration, ingestion of larvae from soil, or by eating a
cockroach. Dogs can get infected by ingesting larvae on grass, the skin
penetration of larvae, or the ingestion of either the cockroach or mouse
paratenic hosts.
Pruritus: an unpleasant sensation that induces someone to scratch or rub the
skin
Transmammary transmission: Transmission via the milk/mammary gland.
Puppies become infected with the larvae of A. caninum while nursing. Some
percentages of larvae that enter a dog persist in the dog’s muscles. These
larvae are reactivated (probably hormonally) towards the end of gestation and
make their way to the mammary glands where they are transferred to the
suckling puppies.
Vesicle: small circumscribed fluid-filled elevations on the skin, also called
blisters. Bullae are large vesicles.
 Review Trends in Parasitology Vol.26 No.4

Figure 1. Cutaneous larval migrans (also known as ‘creeping eruption’) is a zoonotic infection with hookworm species that do not use humans as a definitive host, the most
common being A. braziliense and A. caninum. The normal definitive hosts for these species are dogs and cats. The cycle in the definitive host is very similar to the cycle for
the human species. Eggs are passed in the stool , and, under favorable conditions (moisture, warmth, shade), larvae hatch in 1–2 days. The released rhabditiform larvae
grow in the feces and/or the soil , and after 5–10 days (and two molts) they become filariform (third-stage) larvae that are infective . These infective larvae can survive 3–
4 weeks in favorable environmental conditions. On contact with the animal host , the larvae penetrate the skin and are carried through the blood vessels to the heart and
then to the lungs. They penetrate into the pulmonary alveoli, ascend the bronchial tree to the pharynx, and are swallowed. The larvae reach the small intestine, where they
reside and mature into adults. Adult worms live in the lumen of the small intestine, where they attach to the intestinal wall and suck blood. Some larvae become arrested in
the tissues, and serve as source of infection for pups via transmammary (and possibly transplacental) routes . Humans may also become infected when filariform larvae
penetrate the skin . With most species, the larvae cannot mature further in the human host, and migrate aimlessly within the epidermis, sometimes as much as several
centimeters a day. Some larvae may persist in deeper tissue after finishing their skin migration and a few species may enter the human intestine and develop further.

[19,10]. A. braziliense is found along the southeastern
Atlantic coast of North America, the Gulf of Mexico and
the Caribbean Sea, down to Uruguay in South America, in
Africa (South Africa, Somalia, Democratic Republic of
Congo, Sierra Leone), Australia and Asia (Malaysia and
Indonesia, and based on humans cases, also in Thailand)
[3,5,19,20]. In India, A. braziliense-like lesions are reported
in humans, with adults being described from animals
[21,22], but the necropsy literature is confusing because
of the morphologic similarity of A. ceylanicum and A.
braziliense [19].
Hookworm life cycle and pathogenicity
The life cycle of these nematodes begins with hookworm
larvae hatching from eggs passed in the host’s feces, devel-
oping to the infective stage (filariform larva) in soil, and
gaining access to the definitive host most commonly
through skin penetration (Figures 1 and 2) [1]. Infective
larvae can also establish infections if ingested [1]. In adult
dogs, a large proportion of A. caninum infective larvae
undergo somatic migration, and puppies are often infected
by the transmammary route [1,23]. Transmammary trans-
mission does not occur in cats or with the other species in
dogs. Ancylostoma spp. are capable of being sequestered in
the tissues of vertebrate paratenic hosts (Box 1) where they
persist without development (Figure 3); thus, infection via
hunting is a possibility [1,23]. For several species (A.
caninum, A. tubaeforme, A. ceylanicum), the adult worms
cause blood loss and anemia. Peracute and acute disease
from blood loss is particularly important and sometimes
fatal in young puppies and kittens. Other hookworms such
as U. stenocephala and A. braziliense cause negligible blood
loss, but can sometimes induce disease [23].
CLM in humans
When humans, dogs and cats are infected percutaneously
with larvae of their own host-adapted hookworms, the
lesion at the penetration site varies from nil to an erythe-
matous papular or vesicular rash (Box 1) with varying
levels of pruritus (Box 1). This rash on people in human-
hookworm endemic areas is called ‘ground itch’ [1]. In the
early 1900 s, a lesion distinct from ground itch was found to
occur within the USA (Figure 4). This lesion had a charac-
teristic serpiginous track that could grow for weeks to
months, was intensely pruritic and was ultimately self-
limiting.

163
Review
Figure 2. Hookworm larva in skin biopsy of a person with cutaneous larva migrans
(scale bar = 100 mm). Inset shows higher magnification of coiled larva in the
epidermis (scale bar = 25 mm).
Creeping eruption, ‘a linear, tortuous, and serpiginous
eruption,’ was first described by Lee in 1874, and the first
causative agent identified was a Gasterophilus maggot
(Box 1) [24]. In the USA, it was generally believed that
the creeping eruptions seen in the southern states were
myiases (Box 1), but Kirby-Smith and co-workers looked
for another cause after failing to demonstrate a geographi-
cal correlation between people with lesions and the distri-
bution of Gasterophilus in Florida. In studies that
culminated with experiments using human volunteers,
the cause of the ‘creeping eruptions’ was found to be the
larvae of A. braziliense (Figure 5) [24–26]. These larvae
produced long serpiginous tracks rather than the pruritic
papular lesions (Box 1) produced by human hookworms, A.
caninum, or A. ceylanicum [24,27]. These ‘creeping erup-
tions’ have since been identified as hookworm associated
CLM [28].
In the USA, the disease was called ‘plumber’s itch,’
because the lesions appeared on plumbers who would
crawl under the raised-floor houses of the coastal south
and work in the moist, shaded, sandy soil. The lesions
Figure 3. Larva of Ancylostoma caninum in the muscle of an experimentally
infected macaque monkey. Double lateral alae are evident on one side of the larva.
Scale bar = 50 mm.
164
Trends in Parasitology Vol.26 No.4
would begin soon thereafter and would develop into
cutaneous serpiginous lesions. In one outbreak, lesions
appeared on eight laborers working in a three-foot crawl
space under a hospital being built in Florida [29]. An
outbreak of CLM associated with a contaminated sandbox
recently occurred at a children’s camp in Florida [30].
Geographically, the distribution of hookworm associ-
ated CLM appears to mirror the geographic distribution
of A. braziliense [19,31,32]. Most reported cases have been
in tourists who frequented beaches in regions where A.
braziliense is endemic in dogs and cats [31,32]. The disease
does not occur after exposure to beaches where A. brazi-
liense is not present, e.g. the Pacific coast of the USA and
Mexico. CLM-like lesions have been reported in India [21],
but A. braziliense has not been reported there, so the
species in question is unresolved [19]. No A. braziliense
was found in a recent Australian survey of a large number
of dogs and cats, which could explain the lack of CLM
lesions in people frequenting beaches in that country [20].
Other presentations of larval hookworm infections
Other hookworm larva-associated lesions may be due, in
part, to the species involved. A. tubaeforme appears not to
cause CLM, A. ceylanicum tends not to cause CLM, but can
develop into adults in people, and A. caninum can cause
folliculitis, myositis, and, on occasion, eosinophilic enter-
itis (Box 1). The relationship between U. stenocephala and
CLM remains unclear. Percutaneous infection of human
induced serpiginous tracks can persisted for 3–4 weeks
[33], and larvae can infect dogs percutaneously causing
pedal dermatitis [34], but such larvae seldom make it to the
lungs or develop to adults in dogs [23].
After percutaneous infection, Ancylostoma larvae make
their way to the lungs and then to other body tissues of the
host [23]. This probably accounts for some cases of CLM
progressing to pneumonitis. The first case occurred in 1951
in New Orleans, LA, USA [4]. The patient developed CLM
with a papular rash immediately after fixing the pipes
under his house. Seven days later, he presented to the
hospital with pneumonitis. In the hospital, the rash per-
sisted, and third-stage hookworm larvae were found in his
sputum for the next 24 days. Larvae were also recovered
from the soil under the house. Other cases of pneumonitis
following CLM have been described in visitors to beaches
in the Gulf of Mexico, Barbados and Thailand [3,5,31,32].
In another case, a hookworm larva was found in the
muscle fiber of a man following CLM in Shreveport, LA,
USA [7]. The patient was initially treated for CLM on his
right thigh, back, left shoulder and arms. This was followed
over the next several weeks with eosinophilia and pneu-
monitis with pulmonary infiltrates. Pain and swelling in
the back and thigh developed, and, over the next year, he
received treatments with levamisole, thiabendazole,
diethylcarbamazine and mebendazole. Thirteen months
later, a biopsy revealed the intracellular larva that was
identified as A. caninum because that is the only larval
hookworm described within muscle fibers (A. braziliense
and A. tubaeforme larvae prefer the paratenic host’s muco-
sal epithelium of the nasopharynx), A. braziliense was not
reported in northern LA, and A. tubaeforme larvae were
considered minimal skin penetrators [7]. It seems that
 Review
Figure 4. Redrawn map from 1932 of the distribution of cases of cutaneous larva migrans (CLM) that was used to define the extent of the disease in the USA in the original
description of the syndrome [29]. This continues to be the primary distribution of the condition in the USA.
signs of myositis might also have occurred in the arms of
two volunteers percutaneously infected with the larvae of
A. caninum [6].
The skin lesions from A. braziliense larvae are serpigi-
nous and persistent; lesions of A. caninum follicular, pap-
ular/pustular and ephemeral; lesions of A. ceylanicum
comparatively minimal; and there may be little skin
penetration with A. tubaeforme [7,24,27]. Hookworm lar-
vae in a biopsy from a case of folliculitis in New South
Wales, Australia, were identified as A. caninum because
the only hookworms there were thought to be A. caninum,
A. tubaeforme and U. stenocephala [10]. Most cases of
hookworm associated follicular dermatitis are likely to
be caused by A. caninum [8,9,11,12].
Figure 5. Massive numbers of lesions typical of hookworm associated CLM on a
patient visiting the emergency room of Cook county Hospital in Chicago, USA,
after a vacation in Florida. Photo courtesy of Scott C Sherman, MD, FAAEM,
Department of Emergency Medicine, Cook County Hospital (Stroger), Chicago, IL.
Trends in Parasitology Vol.26 No.4
Hookworm larvae are suspected as potential causes of
diffuse unilateral subacute neuroretinitis, a form of ocular
larva migrans, which can lead to loss of vision, inflam-
mation in the posterior eye, and retinal lesions [14]. Two
different sizes of nematode larvae have been observed in
affected eyes. A. caninum, Toxocara spp, and Strongyloides
spp. have been proposed as representing the smaller lar-
vae, whereas larvae of Baylisascaris spp. have been impli-
cated as the larger larvae [14].
CLM in unexpected geographical locations
Europe: Autochthonous CLM cases have been described
from the UK, Germany, Italy and Serbia; although biopsies
were taken in several cases, hookworm larvae have not yet
been identified. In the UK, a 50-year-old presented 3 weeks
after having participated in outdoor sports in October
where he wore clothing that had been stored in a shed
where a dog was kept [35]. Eight days later, he noted about
40 insect bite-like lesions on his forearm and abdomen. At
presentation, the arm, chest and abdomen were covered
with thin serpiginous tracks. A biopsy did not reveal
larvae, but he responded to albendazole treatment. U.
stenocephala was considered the cause because of the lack
of local A. caninum. In two additional UK cases, crusted
eczematous plaques on the buttocks with serpiginous
tracks extending from their margins responded to therapy
with albendazole or ivermectin [36,37]. A 10-year-old Eng-
lish child had a 3 cm serpiginous track on his instep and
pruritus of a month’s duration [38]. Four cases have been
reported from Germany. Three had visited lakesides and
riverbanks around Berlin and had single serpiginous
tracks on the foot or hand [39]. The other, a lady from
Saxony, developed a bullous lesion (Box 1) on the back of
her hand about a month after the flooding of the Elbe forced
her to spend time in a shelter with numerous domestic and
wild animals [40]. A similar case, without bullae, occurred
in France in a man who developed multiple serpiginous
165
Review
tracks on his leg after standing for a long period in mud in
his flooded house [41]. There are three reports from Italy:
two reports of individual cases and one involving six people
who had contact with potting soil used in dried floral
arrangements [42–44]. In two cases from Serbia, one
patient had chest lesions that persisted for five months
after working construction in August and one had lesions
on his back for five months after having laid on the ground
in June repairing his car [45].
New Zealand: Three cases of autochthonous CLM have
been reported, two in children on the North Island and one
in an avid 80-year-old lady gardener on the South Island
[18,46]. A hookworm larva was identified in the biopsy
specimen from the lady. New Zealand is not known to have
any A. braziliense, has only a little A. caninum on the North
Island, and U. stenocephala has been reported from dogs,
but not cats.
Animal hookworms developing to adults in humans
A. ceylanicum is known to develop on occasion to the adult
stage in people, and can cause abdominal discomfort [1].
Immature solitary adult A. caninum worms (15 cases as of
1998) have been found in the bowel lumen, indicating
maturation within the human gut following percutaneous
infection or perhaps larval ingestion [1,15]. Patients pre-
sented with mild to severe eosinophilic enteritis, charac-
terized by abdominal pain, peripheral eosinophilia, and
eosinophilic infiltration of the bowel wall. A. ceylanicum
adults have now also been recovered from a human colon in
Taipei [47].
Hookworm treatment and prevention for companion
animals
Treatment of dogs and cats for hookworms is covered
elsewhere (see: http://www.capcvet.org/recommendations/
hookworms.html and http://www.cdc.gov/ncidod/dpd/para-
sites/ascaris/prevention.htm). Because re-infection is com-
mon, regular internal parasite control is recommended,
and, when possible, dogs and cats should be on year-round
internal parasite control to minimize the shedding of hook-
worm eggs. In Australia, concern has been raised about the
resistance of A. caninum to pyrantel [48], but this has not
been observed elsewhere. Treatment success should
always be verified through regular fecal examinations on
cats and dogs.
Hookworm treatment and prevention for humans
Although CLM is a self-limiting infection, antiparasitic
and symptomatic treatment is often sought because of
the associated discomfort and potential for secondary bac-
terial infection with this disease. Albendazole or ivermec-
tin are the treatments of choice [32]. Attempts to freeze or
surgically remove cutaneously migrating larvae are gener-
ally ineffective, and can result in further injury [32]. Elev-
ated dosages of ivermectin might also kill larvae in muscle
tissues.
Preventative measures include: wearing protective foot-
wear on beaches, covering sand boxes, cleaning up animal
waste, preventing cats and dogs from defecating in public
areas, and, when lying on beach sand, seeking sand washed
by the tide over other parts of the beach and using mat-
166
Trends in Parasitology Vol.26 No.4
tresses rather than towels [32,49]. In less developed
countries, control of this disease by mass drug adminis-
tration with ivermectin might be more feasible [50].
Next steps
Defining the burden of disease in both developed and less-
developed countries is critical to controlling this disease. In
the developed world, surveys of specialists, e.g. dermatol-
ogists, gastroenterologists and ophthalmologists, would
better define the incidence of human disease and risk
factors for infection, which would, in turn, inform preven-
tative measures. Improved diagnostics are needed to
confirm that hookworms are the source of observed lesions
and, ideally, to identify the species. A diagnostic test that
identifies the genus of worm in ocular infections is critically
needed for assessing potential risks. Recently developed
molecular methods that discriminate species of hookworm
eggs from animal feces, if widely utilized, could determine
the geographic distributions through fecal analysis rather
than necropsy. Some veterinarians would like a rapid fecal
assay for a generic hookworm infection to assist in diag-
nosis and treatment. In less-developed countries, where
CLM may cause significant health impacts (but be largely
unrecognized), community surveys quantifying the mor-
bidity due to hookworm associated CLM are needed. In
these settings, integrating treatment of CLM in existing
control programs for other infections might be the most
efficient approach [50].
Disclaimer
The findings and conclusions in this manuscript are those
of the authors and do not necessarily represent the views of
the Centers for Disease Control and Prevention.
Acknowledgements
This material developed out of a workshop held at the Centers for Disease
Control and Prevention (CDC) in Atlanta, Georgia in 2008 with invited
representatives from the Companion Animal Parasite Council (CAPC),
the CDC, the American Association of Veterinary Parasitologists (AAVP)
and others. Many people made this event possible, including the critical
help of Lonnie King and Mary Bartlett of the CDC, and Mike Paul and
Sonya Hennessey of CAPC. The authors also thank Blaine Mathison for
assistance in creating the lifecycle figure.
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