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Veterinary Research Communications
https://doi.org/10.1007/s11259-023-10171-5
REVIEW
Is leishmaniasis thenew emerging zoonosis intheworld?
EsperanzaMontaner‑Angoiti1· LolaLlobat1
Received: 20 May 2023 / Accepted: 5 July 2023
© The Author(s), under exclusive licence to Springer Nature B.V. 2023
Abstract
Leishmania is a genus of parasitic protozoa that causes a disease called leishmaniasis. Leishmaniasis is transmitted to humans
through the bites of infected female sandflies. There are several different species of Leishmania that can cause various forms
of the disease, and the symptoms can range from mild to severe, depending on species of Leishmania involved and the
immune response of the host. Leishmania parasites have a variety of reservoirs, including humans, domestic animals, horses,
rodents, wild animals, birds, and reptiles. Leishmaniasis is endemic of 90 countries, mainly in South American, East and
West Africa, Mediterranean region, Indian subcontinent, and Central Asia. In recent years, cases have been detected in other
countries, and it is already an infection present throughout the world. The increase in temperatures due to climate change
makes it possible for sandflies to appear in countries with traditionally colder regions, and the easy movement of people and
animals today, facilitate the appearance of Leishmania species in new countries. These data mean that leishmaniasis will
probably become an emerging zoonosis and a public health problem in the coming years, which we must consider control-
ling it from a One Health point of view. This review summarizes the prevalence of Leishmania spp. around the world and
the current knowledge regarding the animals that could be reservoirs of the parasite.
Keywords Infection· Leishmania· Prevalence· World· Zoonosis
Introduction
Leishmaniasis is a vector-borne disease caused by an obli-
gate parasitic protozoan, from the genus Leishmania (family
Trypanosomatidae). It is transmitted by the bite of infected
female sandflies of the genus Phlebotomus in the Old World,
and of the genus Lutzomyia in the New World at least 93
sandfly species are proven or probable vectors worldwide
(Akhoundi etal. 2017; Mann etal. 2021).
Leishmania is a wide genus, it is meaning that it has a
complex life cycle that involves two main stages, the pro-
mastigote, and the amastigote, changing according to the
host: invertebrate vector (sandfly) and vertebrate host
(human or animal), respectively. Transmission is mediated
by the sandfly and can be anthroponotic or zoonotic depend-
ing on the region (Pace 2014).
Female sandfly takes a blood meal from an infected ver-
tebrate host, ingesting macrophages or other host cells that
contain amastigotes, which are the intracellular form of
Leishmania parasites. Once inside the sandfly, the amas-
tigotes transform into promastigotes in the midgut of the
insect. These promastigotes are elongated with a hair-
like flagellum and multiply through binary fission. They
migrate to the anterior portion of the sandfly’s gut and
attach to the gut wall, where they continue to divide (Slama
etal. 2014).
After several days, the promastigotes differentiate into
infective metacyclic promastigotes. These metacyclic pro-
mastigotes are highly mobile and infectious. They migrate
toward the proboscis of the sandfly, ready to be transmitted
to a new vertebrate host. When an infected sandfly takes a
blood meal from a vertebrate host, it injects the infective
metacyclic promastigotes into the skin. Once inside the host,
the metacyclic promastigotes are engulfed by phagocytes,
such as macrophages. Within the phagocytes, the metacy-
clic promastigotes transform into amastigotes, which are
the intracellular stage of the parasite. The amastigotes mul-
tiply within the host cells, primarily within macrophages,
and form clusters called amastigote-laden parasitophorous
* Lola Llobat
maria.llobatbordes@uchceu.es
1 Molecular Mechanisms ofZoonotic Disease (MMOPS)
Group, Facultad de Veterinaria, Universidad Cardenal
Herrera-CEU, CEU Universities Valencia, Valencia, Spain
Veterinary Research Communications
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vacuoles. The presence of Leishmania parasites inside the
host cells triggers an immune response, leading to the clini-
cal manifestations of leishmaniasis. If an infected sandfly
takes a blood meal from the infected vertebrate host, it
ingests the amastigote-laden macrophages, restarting the
cycle (Solano-Gallego etal. 2012; Akhoundi etal. 2016;
Bates 2018) (Fig.1).
This heterogeneous nature of Leishmania has important
implications for disease transmission and control, as inter-
ventions must target both the vertebrate and invertebrate
hosts to effectively reduce disease transmission (Akhoundi
etal. 2016). The primary hosts are vertebrates commonly
infecting canids, rodents, marsupials, mongooses, bats,
hyraxes, and humans, being the disease is an important
zoonosis around the world (Millán etal. 2014; Akhoundi
etal. 2017). In fact, according to the World Health Organi-
zation (WHO), more than 1 billion people live at risk of
infection in endemic areas for leishmaniasis, with approxi-
mately 30,000 new cases of visceral leishmaniasis (VL)
and more than one million new cases of cutaneous leish-
maniasis (CL) occurring annually (www. who. int/ health-
topics/ leish mania sis).
The purpose of this review is to summarize the current
knowledge of the species that are affected by this parasite
and to raise awareness, that Leishmania is infecting a wide
and growing range of species, and its importance as a pos-
sible reservoir as it is an emergent zoonotic disease.
Classication ofLeishmania genus
There are many different species of Leishmania, which
are classified based on their geographic distribution, host
range, and clinical manifestations of the disease. Currently,
54 Leishmania species are identified (Akhoundi etal. 2016).
Thirty have been proven to affect mammals, of which at
least 21 are pathogenic to humans. Overall, the classification
of Leishmania is complex and continues to evolve as new
research is conducted on the parasites and their interactions
with hosts and vectors. Figure2 shows the complete clas-
sification of Leishmania species.
For this review, we summarize the most relevant Leish-
mania species in terms of human and animal health in four
complexes: L. donovani; L. tropica; L.mexicana; L. brazil-
iensis. L. donovani includes L. donovani (syn, L. archibaldi)
and L. infantum (syn., L chagasi) can cause cutaneous leish-
maniasis (CL) and visceral leishmaniasis (VL) in humans
and animals. These parasites are found in parts of Europe,
Asia, Africa, America (especially South America) the Mid-
dle East. L. tropica complex includes L. tropica (syn. L kil-
licki) and L. aethiopica. These species can cause CL, and
we can encounter them in parts of Asia, Africa, and the
Middle East, L. mexicana complex includes several species
L. mexicana (syn. L. pifanoi), L. amazonensis (syn. L. gar-
hami), L. aristidesi, L. forattinii, L. venezuelensis, L.waltoni
found in Central and South America, causing CL in humans,
Fig. 1 Life cycle of Leishmania parasite. Created with Biorender.com
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L. brazilensis complex theseparasites can cause CL and
mucocutaneous leishmaniasis (MCL) in the Americas, and
L. major complex includes L. major, L. gerbili, L. turanica
and L. arabica. These species are primarily found in North
Africa and the Middle East, where it is a significant cause
of cutaneous leishmaniasis in these regions.
Leishmania spp. intheworld
Leishmaniasis is endemic in many regions of the world,
including parts of Africa, Asia, Europe, and the Americas
(Figs.3 and 4). The prevalence of the disease is influenced
by a variety of factors, including climate, environment, and
socioeconomic conditions (Torres-Guerrero etal. 2017).
In Europe, the disease is endemic in several countries,
particularly in the Mediterranean basin. The prevalence of
Leishmania and the species causing the disease can vary
between regions and even within countries (Palma etal.
2021; Van der Auwera etal. 2022).
The main species found in Europe are L. infantum and
L. tropica. L. infantum is the most common species found
and is endemic in the Mediterranean area, affecting mainly
mammals and humans, and responsible for causing human
visceral and cutaneous leishmaniasis, and canine leishmani-
osis. L. tropica has been reported in Greece, and neighbor-
ing countries causing sporadically anthroponotic cutaneous
leishmaniasis (Alvar etal. 2012; Antoniou etal. 2013; Ber-
riatua etal. 2021; El Idrissi etal. 2022).
Although infection of Leishmania is common in Spain,
Italy, Portugal, Greece, and France, the epidemiological
trend of Leishmania spp. is varying in the last few years
as L. infantum is spreading northward, mostly due to the
traveling of infected dogs, becoming a real issue in northern
latitudes where sandfly vectors are either absent or present
in very low densities as is occurring in Germany and the
United Kingdom, where Leishmania parasites are regularly
imported by dogs and humans from endemic regions leading
to a continual risk of autochthonous transmission (Naucke
etal. 2008; Ready 2010; Oerther etal. 2020).
Furthermore, the introduction of exotic Leishmania spe-
cies or strains into Europe via the traveling of humans and
domestic animals. A retrospective analysis of leishmania-
sis cases was conducted in 15 European centers where they
Fig. 2 Classification of Leishmania species. Adapted from Akhoundi etal. (2016) and Klatt etal. (2019)
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found a total of 9 different species of Leishmania, all of them
except L. infantum considered as imported or travel-related
infections (Millán etal. 2014; Van der Auwera etal. 2022).
Lastly, there is evidence based on molecular markers, that
European vectors of Leishmania have extended their ranges
northward (Aransay etal. 2003; Perrotey etal. 2005; Ready
2010). Different studies are conducted to understand if cli-
mate change is influencing the distribution of the presence
of Leishmania species in Europe, but so far has been only
theorized (Maroli etal. 2008; Ready 2010).
In the Americas, leishmaniasis is endemic in many coun-
tries, particularly in the Andean and Amazonian regions.
Africa 9,165 cases
Americas 37,502 cases
Eastern Mediterranean 174,920 cases
Europe 201 cases
South-East Asia 2 cases
GLOBAL 221,790 cases
0
< 100
100 - 999
1000 - 4999
>= 5000
Not applicable
No autochthonous cases reported
No data
Fig. 3 Distribution of cutaneous leishmaniasis (CL) around the world
0
< 100
100 - 499
500 - 999
>= 1000
Not applicable
No autochthonous cases reported
No data
Africa 3,825 cases
Americas 1,604 cases
Eastern Mediterranean 4,660 cases
Europe 214 cases
South-East Asia 1,464 cases
GLOBAL 11,767 cases
Fig. 4 Distribution of visceral leishmaniasis (VL) around the world
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The prevalence and species of Leishmania causing the dis-
ease can vary between regions and even within countries,
making leishmaniasis a complex disease, encountering up
to ten species within the same territory (Herrera etal. 2020).
Leishmaniasis is a complex and endemic disease in the
American continent, due to the high concentration of dif-
ferent species in the same country, encountering up to ten
species within the same territory (Ramírez etal. 2016; Mon-
talvo etal. 2017; Herrera etal. 2020).
An interactive database of the distribution of the differ-
ent Leishmania species in the Americas has been created by
Herrera etal. (2020), reporting a total of 20 species with a
wide range of hosts. L. amazonensis, L. brazilensis, L. guay-
anensis, L. infantum, L. mexicana, L. panamensis, and L.
peruviana are the most reported in the bibliography affecting
humans. Table1 summarizes the different species that have
been described in hosted vertebrates by country in Americas.
In the United States (U.S.), leishmaniasis is the most
common imported infection, mostly acquired during travel
abroad. Except L. mexicana which is considered endemic
in the U.S., most reported in Texas and southeast Okla-
homa and L. infantum is present in dogs in the U.S. with
a prevalence of up to 20% (range 2–20%). In humans is
considered an imported disease, as they consider the posi-
tive cases mostly related to the American soldiers that
return from Iraq, as 19.5% of the return with asymptomatic
visceral leishmaniasis, and general immigration. Sandfly
transmission to humans is possible but not confirmed in
U.S. (McHugh etal. 1996; Clarke etal. 2013; McIlwee
etal. 2018; Curtin and Aronson 2021).
The most common species found in Asia are L. infantum
and L. donovani, being responsible for most of the VL and
CL in humans (Ready 2014). India has the highest burden
of VL in the world, with 33,000 new cases reported in 2019
according to the National Vector Borne Disease Control Pro-
gram (Lukes etal. 2007; Ghatee etal. 2020; Kumar etal.
2020). Another country with a high burden of VL is Afghan-
istan with over 5,000 new cases reported in 2020, according
to the Ministry of Public Health (Knight etal. 2022). In Iran,
a study published in 2020 found that the prevalence of CL
was highest in the central and eastern regions of the country,
with over 9,000 cases reported in 2018 (Ghatee etal. 2020).
In Pakistan, there were over 12,000 cases of CL reported in
2019, according to the Ministry of National Health Services,
Regulations, and Coordination (Kayani etal. 2021). MCL
presentation, typical of L. brazilensis complex infection, is
rare in Asia (Strazzulla etal. 2013).
The disease is endemic in many countries in Africa,
particularly in North Africa, East Africa, and parts of
West Africa. The prevalence of Leishmania spp. can
vary between regions and even within countries being L.
donovani and L. infantum as the most common species
causing leishmaniasis in Africa (Alvar etal. 2021). The
prevalence of leishmaniasis in Africa ranges from 14%-
50%, the highest range found in Algeria with the second
highest incidence in CL, commonly caused for L. major
in this area (Aoun and Bouratbine 2014; Beniklef etal.
2021). Overall, the prevalence of leishmaniasis in Africa
is difficult to estimate due to the lack of adequate reporting
and surveillance systems in many regions. However, it is
believed that the disease is underdiagnosed and underre-
ported in many parts of the continent (Jones and Welburn
2021; Viana de Almeida etal. 2021).
Methods fordetection ofparasite
Serological methods
Involves the detection of antibodies produced by the immune
system against Leishmania spp. parasites in the blood or
other body fluids of infected individuals. IFAT (Indirect Flu-
orescent Antibody Test), ELISA (Enzyme-Linked Immuno-
sorbent Assay) and Western Blot (WB) are three commonly
used serological tests for the diagnosis of leishmaniosis. The
principle of these three techniques is based on the detection
of antibodies against the Leishmania parasite in a patient's
blood serum (Persichetti etal. 2017).
In the IFAT test, a small amount of Leishmania antigen
is placed on a slide and mixed with the patient's blood sam-
ple. If the patient has previously been infected, their blood
will contain antibodies that will stick to the antigen. Next,
a fluorescently labeled secondary antibody is added. If the
patient's antibodies are present, the secondary antibody
will bind to them. If the sample fluoresces indicates that
the patient has antibodies against Leishmania (Ryan etal.
2002). ELISA method works on a similar principle but in
this test the secondary antibody is enzyme-labeled instead
of fluorescent-labeled. For ELISA tests it is common to use
microtiter plates, where the Leishmania antigen is attached,
and the patient's serum is added. If the patient has antibodies
against Leishmania, these antibodies will bind to the antigen
on the plate, and the secondary antibody will also bind to the
patient’s antibodies if present. A substrate is added helping
to produce an enzymatic reaction that will produce a color
change that can be detected with a spectrophotometer (Alhajj
and Farhana 2023).
Both IFAT and ELISA are sensitive and specific tests
for the diagnosis of leishmaniasis, but they differ in
their sensitivity and specificity depending on the antigen
used, the stage of the disease, and the population being
tested. IFAT tests show acceptable sensitivity (87–100%)
and specificity (77–100%), but it is more expensive and
requires sophisticated laboratory. ELISA is the preferred
laboratory test for the serodiagnosis of leishmaniasis as
it is easier and faster to perform and can be done using
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Table 1 Leishmania spp. found in the Americas classified by country and vertebrate host. O: order
Subgenus Species Countries Vertebrate Host Reference
Leishmania L. donovani Brazil, Colombia, Mexico, Nicaragua, Panama,
Venezuela
O. Primates: Human (Homo sapiens) Harris etal. (1998); Monroy-Ostria etal. (2000)
O. Carnivora: Dog (Canis lupus familiaris), Zulueta etal. (1999)
O. Didelphimorphia: Didelphis albiventris, Brazil-
ian common opossum (Didelphis aurita), Com-
mon opossum (Didelphis marsupialis)
Sherlock etal. (1984)
O. Rodentia: Black rat (Rattus rattus) Zulueta etal. (1999)
L. infantum Argentina, Bolivia, Brazil, Colombia, Costa Rica,
Honduras, Mexico, Nicaragua, Peru, Uruguay,
Venezuela
O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O. Carnivora: Molina's hog-nosed skunk (Cone-
patus chinga), Cat (Felis silvestris catus), Dog
(Canis lupus familiaris), Patagonian fox (Pseu-
dalopex griseus), Crab eating fox (Cerdocyon
thous), Lion (Panthera leo), Jaguar (Panthera
onca), Bush dog (Spheotos venaticus)
Dahroug etal. (2010, 2011); Silva etal. (2014); Mil-
lán etal. (2016); Almeida etal. (2018); Lopes etal.
(2020)
O. Chiroptera: Dog-faced bat (Molossus molossus),
Pallas's long-tongued bat (Glossophaga soricina),
Black myotis bat (Myotis nigricans), Ghost bat
(Diclidurus scutatus), Seba's short-tailed bat
(Carollia perspicillata)
Savani etal. (2010); Gómez-Hernández etal. (2017)
L. major Argentina, Ecuador, Peru, Mexico O. Primates: Human (Homo sapiens), Red-handed
howler monkey (Alouatta belzebul) Calvopina etal. (2006); Lau etal. (2014); Pasquali
etal. (2019); Martínez etal. (2020)
O. Carnivora: Patagonian fox (Pseudalopex griseus) Millán etal. (2016)
O. Didelphimorphia: White-bellied opossum
(Didelphis albiventris), Brazilian common opos-
sum (Didelphis aurita)
Grimaldi etal. (1987); Corredor etal. (1989); Hum-
berg etal. (2012)
O. Lagomorpha: European rabbit (Oryctolagus
cuniculus) Gonçalves etal. (2012)
O. Perissodactyla: Horse (Equus ferus caballus) Soares etal. (2013)
O. Rodentia: Hamster (Cricetidae spp.), Hystri-
cognath rodents (family Hystricognathi), Mouse
(Mus musculus), Necromys laisuris, Black rat
(Rattus rattus), Proechimys canicollis
Berzunza-Cruz etal. (2000); Cássia-Pires etal.
(2014)
L. mexicana Argentina, Brazil, Belize, Bolivia, Colombia,
Ecuador, Guatemala, Honduras, Mexico, Panama,
Peru, Venezuela, Costa Rica
O. Primates: Human (Homo sapiens), Red howler
monkey (Alouatta palliata), Black howler monkey
(Alouatta pigra)
Berzunza-Cruz etal. (2000, 2009); Rovirosa-Hernán-
dez etal. (2013)
O. Carnivora: Dog (Canis lupus familiaris), Cat
(Felis silvestris catus),
López-Céspedes etal. (2012); Rivas etal. (2018);
Castillo-Ureta etal. (2019)
O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
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Table 1 (continued)
Subgenus Species Countries Vertebrate Host Reference
O. Chiroptera: Mexican fruit bat (Artibeus
jamaicensis), yellow-shouldered bat (Sturnira
lilium), Vampire bat (Desmodus rotundus),
Common bat (Carollia sowelli), Ludwig's bon-
neted bat (Sturnira ludovici), great fruit-eating
bat (Artibeus lituratus), black mustached bat
(Dermanura phaeotis), Seba’s short-tailed bat
(Carollia perspicillata), Godman’s long-tailed bat
(Choeroniscus godmani), Pallas's long-tongued
bat (Glossophaga soricina), Wagner’s mustached
bat (Pteronotus parnelli and Pteronotus persona-
tus, Black myotis bat (Myotis nigricans),
Berzunza-Cruz etal. (2015)
O. Didelphimorphia: Brazilian common opossum
(Didelphis aurita) Grimaldi etal. (1987); Guimarães-E-Silva etal.
(2017)
O. Pilosa: Mexican tamandua (Tamandua mexi-
cana) Lainson etal. (1982)
L. tropica Peru, Mexico O. Rodentia: Syrian hamster (Mesocricetus aura-
tus), Gaumer’s spiny pocket mouse (Heteromys
gaumeri and Heteromysgaumeri desmarestianus),
Mouse (Mus musculus), Hamster (Cricetidae
spp.), Nicaraguan oryzomys (Oryzomys melano-
tis), Yucatan deer mouse (Peromyscus yucat-
anicus), Hispid cotton rat (Sigmodon hispidus),
Black rat (Rattus rattus)
Chable-Santos etal. (1995)
L. amazonensis Argentina, Bolivia, Brazil, Colombia, Ecuador,
Mexico, Paraguay, Peru, Surinam, Venezuela
O. Primates: Human (Homo sapiens), Red-handed
howler monkey (Alouatta belzebul) Calvopina etal. (2006); García Bustos etal. (2016);
Guimarães-E-Silva etal. (2017); Salvioni Recalde
etal. (2019); Hoyos etal. (2020); Martínez etal.
(2020)
O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O. Carnivora: Molina's hog-nosed skunk (Cone-
patus chinga), Cat (Felis silvestris catus), Dog
(Canis lupus familiaris), Crab eating fox Cerdo-
cyon thous
Miles etal. (1980); Tolezano etal. (2007); Souza
etal. (2009); Dias etal. (2011); Buitrago etal.
(2011); Guimarães-E-Silva etal. (2017); Valdivia
etal. (2017); Paz etal. (2018); Herrera etal.
(2018); Alves Souza etal. (2019); Picón etal.
(2020)
O. Cingulata: Armadillo (Dasypodidae spp.) Lainson etal. (1982)
O. Chiroptera: Wagner's bonneted bat (Eumops
perotis), Brazilian porcupine (Coendou pre-
hensilis), Dog-faced bat (Molossus molossus),
Pallas's long-tongued bat (Glossophaga soricina),
Black myotis bat (Myotis nigricans), Ghost bat
(Diclidurus scutatus), Round-eared bats (Carollia
spp.), Bat (Sturnira spp.)
Miles etal. (1980); Savani etal. (2010); Oliveira
etal. (2015); Gómez-Hernández etal. (2017)
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Table 1 (continued)
Subgenus Species Countries Vertebrate Host Reference
O. Didelphimorphia: White-bellied opossum
(Didelphis albiventris), Common opossum
(Didelphis marsupialis), Murine mouse opossum
(Marmosa murina), Water opossum (Metachirus
nudicaudatus)
Arias and Naiff (1981); Grimaldi etal. (1987);
Guimarães-E-Silva etal. (2017)
O. Rodentia: Tropical rat (Hylaeamys spp.), Mouse
(Mus musculus), Guyanan spiny rat (Proechimys
guyannensis), Big marsh rat (Oryzomys capito),
Black rat (Rattus rattus), Hamster (Cricetinae
spp.), Hispid cotton rat (Sigmodon hispidus)
Guimarães-E-Silva etal. (2017)
L. colombiensis Colombia, Venezuela O. Primates: Human (Homo sapiens) Rodriguez-Bonfante etal. (2003)
O. Carnivora: Dog (Canis lupus familiaris), Zerpa etal. (2001)
L. iansoni Bolivia, Brazil, Colombia, Ecuador, Peru, Surinam O. Primates: Human (Homo sapiens) Kato etal. (2016); Guimarães-E-Silva etal. (2017);
Bilbao-Ramos etal. (2017)
Viannia L. brazilensis Argentina, Belize, Bolivia, Brazil, Colombia, Costa
Rica, Ecuador, Guatemala, Mexico, Nicaragua,
Panama, Paraguay, Peru, Surinam, Venezuela
O. Primates: Human (Homo sapiens), Azara's night
monkey (Aotus azarae azarae) Acardi etal. (2013); Patino etal. (2017); Araujo-
Pereira etal. (2018); Olivo Freites etal. (2018);
Copa etal. (2019)
O. Carnivora: Molina's hog-nosed skunk (Cone-
patus chinga), Cat (Felis silvestris catus), Dog
(Canis lupus familiaris), Crab eating fox (Cerdo-
cyon thous)
Buitrago etal. (2011); López-Céspedes etal. (2012);
Longoni etal. (2012); Morgado etal. (2016)
O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O. Chiroptera: Wagner's bonneted bat (Eumops
perotis), Free-tailed bats (Molossidae spp.),
Striped hairy-nosed bat (Platyrrhinus lineatus),
Jamaican fruit bat (Artibeus planirostris)
Shapiro etal. (2013); Guimarães-E-Silva etal. (2017)
O. Didelphimorphia: White-bellied opossum
(Didelphis albiventris), Brazilian common
opossum (Didelphis aurita), Common opossum
(Didelphis marsupialis), Demerara opossum
(Marmosa demerarae)
Silva etal. (2016); Guimarães-E-Silva etal. (2017)
O. Lagomorpha: European rabbit (Oryctolagus
cuniculus), Brazilian Cottontail (Sylvilagus
braziliensis)
Gonçalves etal. (2012)
O. Perissodactyla: Horse (Equus ferus caballus),
Donkey (Equus asinus) Truppel etal. (2014)
O. Pilosa: Hoffmann's two-toed sloth (Choloepus
hoffmanni) Loyola etal. (1988)
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Table 1 (continued)
Subgenus Species Countries Vertebrate Host Reference
O. Rodentia: Hamster (Cricetidae spp.), red-bellied
rice rat (Euryoryzomys russatus), Hystricognath
rodents (family Hystricognathi), Cerradomys,
Akodon cursor, Mouse (Mus musculus), Necro-
mys lasiurus, Black rat (Rattus rattus), Melano-
mys caliginosus, Micromys minutus, big marsh rat
(Oryzomys capito)
Tonelli etal. (2017); Fernández etal. (2018)
L. guayanensis Argentina, Bolivia, Brazil, Colombia, Costa Rica,
Ecuador, Mexico, Panama, Paraguay, Peru, Suri-
nam, Venezuela
O. Primates: Human (Homo sapiens) Rosales-Chilama etal. (2015); Kato etal. (2016);
Muvdi-Arenas and Ovalle-Bracho (2019)
O. Carnivora: Dog (Canis lupus familiaris), Santaella etal. (2011); Guimarães-E-Silva etal.
(2017)
O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O. Chiroptera: Round-eared bats (Carollia spp.),
Bat (Sturnira spp.)
Lourenço etal. (2018)
O. Didelphimorphia: White-bellied opossum
(Didelphis albiventris), Brazilian common
opossum (Didelphis aurita), Common opossum
(Didelphis marsupialis),
Grimaldi etal. (1991); Guimarães-E-Silva etal.
(2017)
O. Pilosa: Hoffmann's two-toed sloth (Choloepus
hoffmanni) Grimaldi etal. (1987, 1991)
O. Rodentia: Hystricognath rodents (family
Hystricognathi), Mouse (Mus musculus), Hamster
(Cricetinae spp.), Rat (Rattus spp.)
Cássia-Pires etal. (2014)
L. lidenbergi Colombia O. Primates: Human (Homo sapiens) Correa-Cárdenas etal. (2020)
L. naiffi Brazil, Colombia, Ecuador, Surinam O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O. Carnivora: Dog (Canis lupus familiaris)
O. Cingulata: Nine-banded armadillo (Dasypus
novemcinctus)
O. Didelphimorphia: opossum
O. Rodentia: Hystricognath rodents (family Hystri-
cognathi), Mouse (Mus musculus)
O. Primates: Human (Homo sapiens) Kato etal. (2013)
L. panamensis Argentina, Brazil, Colombia, Costa Rica, Ecuador,
Mexico, Nicaragua, Panama, Peru
O. Carnivora: Dog (Canis lupus familiaris) Longoni etal. (2011)
O. Primates: Human (Homo sapiens) Figueroa etal. (2009)
O. Rodentia: Hamster (Cricetinae spp.), Mouse
(Mus musculus) Berzunza-Cruz etal. (2009)
L. peruviana Mexico, Peru O. carnívora: Dog (Canis lupus familiaris), King
skunk (Conepatus rex) Llanos-Cuentas etal. (1999); Reithinger etal. (2000)
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Table 1 (continued)
Subgenus Species Countries Vertebrate Host Reference
O. Didelphimorphia: Small marsupials (Marmosa
spp.), White-bellied opossum (Didelphis albiven-
tris)
Llanos-Cuentas etal. (1999)
O. Primates: Human (Homo sapiens) Lucas etal. (1998)
O. Rodentia: Andean leaf-eared mouse (Phyllotis
andinum), small rodents (Akodon spp.), Mouse
(Mus musculus), Rice rat (Oryzomys spp.), Black
rat (Rattus rattus)
Llanos-Cuentas etal. (1999)
L. shawi Brazil, Ecuador, Peru O. Artiodctyla: Cow (Bos taurus) Guimarães-E-Silva etal. (2017)
O.Carnivora: Nasua nasua, Dog (Canis lupus
familiaris)
O. Didelphimorphia: opossum
O. Pilosa: Three-toed sloth (Bradypus tridactylu)
O. Rodentia: Syrian hamster (Mesocricetus
auratus), Mouse (Mus musculus), Hystricognath
rodents (family Hystricognathi),
O. Primates: Human (Homo sapiens), Brown
capuchin (Cebus apella), Black-bearded saki
(Chiropotes satanus)
Lainson etal. (1989); Guimarães-E-Silva etal.
(2017)
Mundinia L. enriettii Brazil O. Rodentia: Guinea pig (Cavia porcellus) Thomaz-Soccol etal. (1996)
CLADE
Paraleish-
mania
L. hertigi Brazil O. Carnivora: Crab eating fox (Cerdocyon thous) Miles etal. (1980)
O. Didelphimorphia: Murine mouse opossum
(Marmosa murina), Water opossum (Metachirus
nudicaudatus)
O. Primates: Human (Homo sapiens)
O. Rodentia: Brazilian porcupine (Coendou prehen-
silis), Guyanan spiny rat (Proechimys guyannen-
sis), Big marsh rat (Oryzomys capito)
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standard laboratory equipment. This technique is highly
sensitive, but its specificity depends upon the antigen used
(Elmahallawy etal. 2014).
If using crude soluble promastigote antigen (CSA) the
sensitivity of the test ranges form 80–100% with and speci-
ficity of 84–95%. Cross-reactions may occur in patients with
trypanosomiasis, toxoplasmosis, and tuberculosis. It is not
recommended to use selective antigenic molecules as has
very low sensitivity (37%) (Sundar and Rai 2002).
A conserved proportion of kinesin-related protein recom-
binant antigen from a cloned protein of L. chagasi (rK39)
has been reported to be highly reactive to sera from human
to canine VL, with a specificity and sensitivity of 99%.
Lastly, lipid-binding proteins have shown high levels of sen-
sitivity and specificity with the absence of cross-reactions
(Elmahallawy etal. 2014). Finally, WB method using 14-kD
and 16 kD antigens have a 100% and 91% of sensitivity for
CL and VL respectively, and no present cross-reactivity with
others, being a good method for diagnosis both two leishma-
niasis presentation (Seyyedtabaei etal. 2017).
Therefore, it is important to interpret serological results
in conjunction with other diagnostic tests, such as PCR, or
direct visualization of the parasite, to confirm the diagnosis
of leishmaniasis. In endemic areas, serology tests may have
a limited diagnostic value where many people and animals
have been previously exposed to the parasite and have devel-
oped antibodies without experiencing symptoms.
Molecular methods
Different molecular methods have been developed and evalu-
ated to ensure better specificity, sensitivity, and reproduc-
ibility in molecular diagnostics of leishmaniasis, including
multilocus enzyme electrophoresis, conventional polymerase
chain reaction (PCR) based assays, quantitative Real-Time
PCR as well as simplified PCR methods.
PCR is the most sensitive and specific technique for
the diagnosis of leishmaniasis, being advantageous over
IFAT and ELISA, as host species-specific reagents are not
required. Also, its sensitivity is greater in cases of CL, MCL,
and immunocompromised patients (Tsokana etal. 2019).
PCR uses different target sequences, which include ribo-
somal RNA genes, kinetoplast DNA (kDNA), miniexon-
derived RNA (medRNA), and the β-tubulina gene region.
The sensitivity and specificity of the test depends on the
sample used, with a wide variety of samples possible as
spleen, lymph node, bone marrow aspirates, buffy coat,
and whole blood, the latest is considered the ideal sample
as its non-invasive. With peripheral blood the PCR has a
sensitivity range from 70–100% (Elmahallawy etal. 2014).
Quantitative real-time PCR (qPCR) has replaced conven-
tional PCR for the diagnosis of leishmaniasis, as is highly
sensitive especially at the lower parasite loads, specific and
reproducible, offering the ability to monitor therapy and to
prevent relapses (Tsokana etal. 2014).
High-resolution melting PCR (HRM-PCR) is a very sen-
sitive amplification technique that enables the direct char-
acterization of amplicons in a closed-tube assay and does
not require additional precautions to prevent the crossover
of PCR products. It measures changes in the fluorescence
intensity of a DNA-intercalating dye during dis- association
from double-stranded DNA to single-stranded DNA, thus
detecting single nucleotide polymorphism. In the last few
years, this technique has become widely used mostly for
epidemiological purposes, as allows to identification of the
species of Leishmania and discriminates between Old World
and New World Leishmania spp. (Derghal etal. 2022).
From a clinical point of view, it is important to note that
PCR can demonstrate parasitemia even before the clinical
presentation starts (Galluzzi etal. 2018).
Prevalence ofLeishmania spp. according tothespecie
Worldwide at least 70 species of wild and domestic animals
have been identified as confirmed or potential reservoirs of
the leishmania parasite (Maia etal. 2018). These potential
reservoirs include human, domestic animals, wild animals,
reptiles, and other non-mammalian animals.
Humans
Leishmaniasis is a major public health problem in many
parts of the world, particularly in developing countries.
The prevalence of leishmaniasis in humans varies widely
depending on the geographic location, the type of leishma-
niasis, and other factors such as environmental conditions,
socioeconomic status, and human behavior. It is considered
that at developed countries most of the infected humans
are at a subclinical level (Singh etal. 2014). The presence
of a higher number of asymptomatic carriers makes infec-
tion control in endemic areas very challenging (Akhoundi
etal. 2017; Ibarra-Meneses etal. 2019). According to the
World Health Organization (WHO), an estimated 700,000
to 1 million new cases of leishmaniasis occur each year,
with more than 90% of cases occurring in the following
countries: Afghanistan, Algeria, Brazil, Colombia, Iran,
Iraq, Pakistan, Peru, Saudi Arabia, and Syria (www. who.
int/ health- topics/ leish mania sis).
In humans, leishmaniasis can present in three main forms,
cutaneous leishmaniasis (CL), visceral leishmaniasis (VL),
and mucocutaneous leishmaniasis (MCL), depending on
the species and the immune status of the infected human
(Alvar etal. 2012). CL is caused by different species of the
subgenus Leishmania (L. major, L. tropica, L. infantum, L.
mexicana, L. amazonensis) and of the subgenus Viannia (L.
braziliensis, L. guyanensis and L. panamensis). This is the
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most common presentation of disease in human, represent-
ing around of 90% of the cases, and it is endemic in many
parts of the world. The estimated annual incidence of CL is
0.7 to 1.2 million cases globally. The highest incidence rates
are reported in Afghanistan, Algeria, Iran, and Saudi Ara-
bia (Alvar etal. 2012; Akhoundi etal. 2017; Volpedo etal.
2021). VL is caused by the viscerotropic species L. infantum
(syn. L. chagasi in Central and South America) and L. dono-
vani. This is a potentially fatal form of leishmaniasis, and
it is endemic in several regions of the world. The estimated
annual prevalence of VL is 50,000 to 90,000 cases glob-
ally. Most cases occur in India, Bangladesh, Nepal, Sudan,
Ethiopia, and Brazil (Dixit etal. 2021). On the contrary,
MCL is a rare form of leishmaniasis caused by an infection
with the Central or South American species L. braziliensis,
L. panamensis and L. guyanensis. This presentation is often
associated with high morbidity although its prevalence is
difficult to estimate, it is thought to be much lower than CL
and VL (Boggild etal. 2019).
Companion animals (dogs andcats)
Dogs have been considered important disease reservoirs in
the domestic environment due the human proximity and inter-
action that can facilitates the transmission through the bite
of infected sandflies (Campino and Maia 2018; Pennisi etal.
2015). Cats can also be infected with the parasite, but they are
not considered to be significant reservoir hosts for the disease
until now (Pennisi 2015; Ahuir-Baraja etal. 2021).
The prevalence of Leishmania spp. infection in compan-
ion animals (dogs and cats) varies widely depending on the
geographic location, the type of leishmaniosis, and other
factors such as the presence of sandflies, the density of dog
populations, and the use of preventative measures.
Canine leishmaniosis (CanL) is endemic in many
regions of the world, particularly in the Mediterranean
basin, the Middle East, and Latin America. Domestic
dogs (Canis lupus familiaris) have been considered the
primary reservoir of L. infantum but can also be infected
by several Leishmania spp. including L. infantum, L.
donovani, L. tropica, L. braziliensis, L. peruviana and L.
panamensis but do not seem to be a significant reservoir
for the last one (Dantas-Torres 2007; Miró etal. 2017).
The prevalence of CanL varies widely depending on the
region, with estimates ranging from less than 1% to over
50%. In endemic areas, the prevalence of CanL can be as
high as 30% to 50% in some dog populations (Morales-
Yuste etal. 2022).
Clinical manifestations of CanL are dermatitis (non-
pruritic exfoliative, erosive-ulcerative, nodular, papular
and/or pustular), onychogryphosis, ocular manifestations
as blepharitis, conjunctivitis, keratoconjunctivitis, anterior
uveitis, and general manifestations as lymphadenomegaly,
loss of body weight, change in appetite, lethargy, mucous
membranes pallor, splenomegaly, polyuria, polydipsia, fever,
vomiting, and diarrhea. Other common manifestations are
mucocutaneous and mucosal ulcerative or nodular lesions,
epistaxis, lameness, atrophic masticatory myositis, vascular
disorders, and neurological disorders (Solano-Gallego etal.
2011; Lago etal. 2019).
Domestic cats have been proposed as a potential sec-
ondary reservoir in endemic areas such as the Mediter-
ranean basin and some parts of South America, although
the prevalence of feline leishmaniosis (FeL) is much lower
than CanL, and it is estimated to be less than 3% in most
endemic areas (Alcover etal. 2021). However, cats could
be a reservoir of the parasite to take into account, so the
prevalence of infection by Leishmania spp. has increased
in recent years (Ahuir-Baraja etal. 2021). Domestic cats
and other felids can also be infected by the same species
described for humans and domestic dogs, that is, L. infantum
in the Mediterranean basin, Iran, and Brazil; L. mexicana in
Texas (USA), L. braziliensis in Brazil and French Guiana,
L. amazonensis in Brazil and L. venezuelensis in Venezuela
(Soares etal. 2016).
Cats are apparently less susceptible than dogs to develop
of disease, and the clinical manifestations include skin or
mucocutaneous and cutaneous lesions, including nodules,
ulcerations or exfoliative dermatitis, and ocular manifesta-
tions (Pennisi etal. 2015).
Livestock animals (Horses andothers)
Equine leishmaniosis is a rare form of leishmaniosis and
has a lower prevalence than in other animal species like
dogs. In Europe, L. infantum has been identified as the
etiological agent of sporadic equine CL in Germany (Koe-
hler etal. 2002), Switzerland (Müller etal. 2009), Spain
(Solano-Gallego etal. 2003), and Portugal (Rolão etal.
2005). In America, autochthonous CL case produced by L.
infantum, and previously attributed to Leishmania siamen-
sis, were reported in Texas (Reuss etal. 2012). In Brazil,
different studies reported CL in horses due to L. infantum
(Soares etal. 2013) and L. braziliensis infection (Agui-
lar and Rangel 1986; Aguilar etal. 1987; Falqueto etal.
1987). In Costa Rica, five cases of CL have been reported
recently and the etiological agent was Leishmania spp.
(Ortega-García etal. 2021).
Few studies have been carried out to determine the preva-
lence of Leishmania spp. infection in horses, and this data
depends on the country. In Italy, Gazzonis etal. (2020) show
13.9% of seroprevalence of L. infantum infection in equine
population, and in Spain, the equine prevalence has been
estimated around 14.2% (Fernández-Bellon etal. 2006).
However, Sudan and Ethiopia demonstrated high seroprev-
alences (33.3%) to L. donovani in donkeys (Kenubih etal.
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2015), and in Brazil, 76.3% of the horses studied were posi-
tive for L. brazilensis (Truppel etal. 2014).
Clinical presentation of the disease is normally less
severe than in other hosts and is not a life-threatening dis-
ease. Equids normally develop skin lesions on the appear-
ance of papules or nodules in the areas where sand flies
commonly feed (eyes, muscles, neck, pinnae, scrotum, and
legs) (Escobar etal. 2019), although infection by VL and
mixed infection with CL in horses has been demonstrated
(Soares etal. 2013).
Risk factors for equine leishmaniosis correlate with
the endemicity of the area and the cohabitation with other
infected hosts, most commonly dogs (Mhadhbi and Sassi
2020). Measures such as insecticide use, and screening and
treatment of infected animals can help to reduce the risk in
these situations.
But not only horses are the livestock specie that could act
as a reservoir of Leishmania spp. Recently, other species have
been found positive for Leishmania, including cow, sheep,
goat, cattle, and pig in endemic regions as Sicily (Abbate
etal. 2020), Ethiopia (Yared etal. 2019), Iran (Rezaei etal.
2022), and Brazil (Paixão-Marques etal. 2019).
Wild animals
Leishmania parasites can be an opportunistic pathogen in
many animals, including wild animals. Depending on the
geographical area, some animals have a more active role
as a Leishmania spp. reservoir (Azami-Conesa etal. 2021).
In Europe, the main wild animal reservoir is rodents, where
the prevalence of L. infantum has been reported to be as high
as 15% in Spain (Galán-Puchades etal. 2019). However, other
wild animals could be infected by L. infantum. In fact, a fatal
case of naturally acquired leishmaniosis was described for
the first time in a Bennett's wallaby in 2011 (Macropus rufog-
riseus rufogriseus) kept in a wildlife park in Madrid, Spain
(Montoya etal. 2016). In Mediterranean basin, L. infantum
infection has been detected in small mammals as mice (Mus
spretus), red squirrel (Sciurus vulgaris) and hedgehog (Eri-
naceous europaeous), wild carnivores as marten (Martes
foina), badger (Meles meles), vison (Mustela vison), red fox
(Vulpes vulpes) (Alcover etal. 2020), American mink (Mus-
tela lutreola) (Giner etal. 2022), European rabbit (Oryctola-
gus cuniculus), European hares (Lepus europaeus) (Abbate
etal. 2019). In South America, a wide range of animals, such
as rodents, marsupials, and monkeys are the main reservoir
hosts for Leishmania spp. The prevalence of leishmania in
wild animals in South America varies widely depending on
the region and the host species (Guiraldi etal. 2022), even
reaching 60% of wild rodents in some areas of Brazil (Azami-
Conesa etal. 2021). Table2 shows different species, including
wildlife, where Leishmania spp. has been found in Europe.
In Africa, Leishmania spp. has been found in the order
Chiroptera, Order Eulipotyphla (hedgehog), Order Roden-
tia (Azami-Conesa etal. 2021), and Order Primates with a
prevalence from 13% in the Gorilla gorilla to 77% in Papio
cynocephalus Anubis (Olive baboons) (Gicheru etal. 2009;
Hamad etal. 2015).
Most of the studies conducted in Asia on wild animals are
on the Order Rodentia, but a cross-sectional study in Iran
investigated the presence of CanL in wild canines finding
L. infantum 19% of the foxes (Vulpes vulpes) and 11% of
jackals (Canis aureus) (Mohebali etal. 2016).
Birds andReptilians
Traditionally, Leishmania spp. has been a parasite that infect
exclusively species of mammals, being necessary a mam-
malian host for its replication and development. Therefore,
the prevalence of leishmania in reptiles was generally con-
sidered to be low or non-existent.
However, in the last years, some studies have reported
the presence of antibodies to Leishmania spp. in birds and
reptiles, in areas where the disease is endemic. In 2010, an
experimental study tries to demonstrate the role of birds
as hosts of L. infantum. The results of this study indicate
that chicken are not suitable host for L. infantum, whereas
domestic gooses (Anser anser) and pheasant (Phasianus col-
chicus) could have a role in the epidemiology of disease in
Brazil (Otranto etal. 2010). At the same, a research group
in Brazil described the first detection of Leishmania of the
subgenus Viannia in yellow-faced parrot (Alipiopsitta xan-
thops) in 2021 (Matheus etal. 2021). More recently, Men-
doza-Roldan etal. (2021) demonstrated the presence of L.
infantum in geckoes (Tarentola mauritanica) in Italy, also
the presence of amastigotes in the bone marrow of geckoes
suggested that the animal was not only exposed to the para-
site but also infected.
However, these findings are rare and further research is
needed to understand the significance of such findings and
their potential role in the transmission of Leishmania spp.
Although, it is possible that birds and reptiles may poten-
tially act as a source of infection for sandflies.
Treatment andprevention
Treatment of human leishmaniasis depends on different fac-
tors, such as clinical forms, efficacy, and toxicity, among
others, and the response of treatment is very heterogeneous.
The Pan American Health Organization (PAHO) has
recently published a guide to recommendations for the
treatment of human leishmaniasis (Pan American Health
Organization 2022). For cutaneous leishmaniasis, the
local treatment recommended is subcutaneous injections
of antimonial, whereas the systematic treatment is with
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Table 2 Leishmania spp. found in the Europe classified by country and vertebrate host. O: order
Subgenus Species Countries Vertebrate Host Reference
Leishmania L. donovani Spain, France, Portugal,
Italy, Greece, Turkey,
Cyprus, Netherlands
O. Primates: Human (Homo sapiens) Antoniou etal. (2009); Bart etal. (2013)
O. Carnivora: Cat (Felis catus) Pereira etal. (2019)
O. Lagomorpha: European hare (Lepus europaeus) Tsokana etal. (2016)
L. infantum Spain, France, Portugal,
Italy, Greece, Turkey,
Cyprus, Georgia,
Germany
O. Primates: Human (Homo sapiens), North-west Bornean
Orangutan (Pongo pygmaeus pigmaeus)
Moral etal. (2002); Chitimia etal. (2011); Miró etal. (2018); Iatta
etal. (2021a, b)
O. Carnivora: Dog (Canis lupus familiaris), Domestic cat (Felis
catus), Red fox (Vulpes vulpes), Wolf (Canis lupus), Golden
jackal (Canis aureus), Tiger (Panthera tigris), Egyptian mon-
goose (Herpestes ichneumon), Iberian lynx (Lynx pardons),
European wildcat (Felis silvestris silvestris), Stone marten
(Martes foina) Pine marten (Martes martes), Weasel (Mustela
nivalis), European mink (Mustela lutreola), Polecat (Mustela
putorius), European badger (Meles meles), Barbary Lion
(Panthera leo); Tiger (Panthera tigris), Egyptian Mongoose
(Herpestes ichneumon), Eurasian Otter (Lutra lutra), Beech
Marten (Martes foina), Domesticated ferret (Mustela putorius
furo), Mediterranean Monk Seal (Monachus monachus),
Brown Bear (Ursus arctos); Common Genet (Genetta genetta),
Raccoon (Procyon lotor)
Rioux etal. (1968); Bettini etal. (1980); Abranches etal. (1983);
Mancianti etal. (1994); Semião-Santos etal. (1996); Martín-
Sánchez etal. (2007); Toplu etal. (2007); Dipineto etal. (2007);
Tabar etal. (2008); Beck etal. (2008); Sobrino etal. (2008);
Verin etal. (2010); Millán etal. (2011, 2015); Chitimia etal.
(2011); Libert etal. (2012); Del Río etal. (2014); Oleaga etal.
(2018); Risueño etal. (2018); Ortuño etal. (2019); Alcover
etal. (2020); Gomes etal. (2020); Iatta etal. (2020, 2021b);
Cantos-Barreda etal. (2020); Villanueva-Saz (2022); Taddei
etal. (2022)
O. Artiodactyla: Roe deer (Capreolus capreolus), Red deer
(Cervus elaphus), Wild boar (Sus scrofa), Domestic goat
(Capra aegagrus), Domestic sheep (Ovis aries)
Rioux etal. (1968); Kantzoura etal. (2013); Taddei etal. (2022)
O. Lagomorpha: Iberian hare (Lepus granatensis), European
hare (Lepus europaeus), European rabbit (Oryctolagus cunicu-
lus)
Chitimia etal. (2011); Molina etal. (2012); Ruiz-Fons etal.
(2013); Moreno etal. (2014); Taddei etal. (2022)
O. Perissodactyla: Horse (Equus caballus), Donkey (Equus
africanus Asinus) Koehler etal. (2002); Fernández-Bellon etal. (2006); Rodrigues
etal. (2019); Nardoni etal. (2019); Escobar etal. (2019); Gaz-
zonis etal. (2020)
O. Rodentia: Black rat (Rattus rattus), Norwegian rat (Rattus
norvegicus), Rat (Rattus spp.), Edible dormouse (Glis glis),
Garden dormouse (Elyomis quercinus), House mouse (Mus
musculus), Yellow-necked mouse (Apodemus flavicollis),
Wood mouse (Apodemus sylvaticus), Algerian mouse (Mus
spretus), Porcupine (Hystricidae spp.)
Rioux etal. (1968); Bettini etal. (1980); Helhazar etal. (2013);
Zanet etal. (2014); Taddei etal. (2022)
O. Diprotodontia: Bennett’s Wallaby (Macropus rufogriseus) Ramírez etal. (2013)
O. Chiroptera: Schreibers' bats (Miniopterus schreibersii), Urban
bats (Pipistrellus pipistrellus)
Millán etal. (2011); Azami-Conesa etal. (2020)
O. Insectivora/ O. Eulipotyphla: European hedgehog (Erina-
ceous europaeus), White-toothed shrew (Crocidura russula)
Rioux etal. (1968)
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miltefosine. However, in pregnant women, patients with
electrocardiogram alterations, kidney, liver and/or heart
disease, or immunocompromised, other treatments are
most adequate, such as thermotherapy, or amphotericin
B deoxycholate, whereas for pediatric patients, the rec-
ommendation is pentavalent antimonial. This treatment is
effective for mucocutaneous and mucosal form of leishma-
niasis, and for visceral leishmaniasis, where the ampho-
tericin B deoxycholate and liposomal amphotericin B are
also adequate.
The PAHO warns that the level of safety regarding
cure with any of these treatments is moderate or low,
and all of them are toxic drug, therefore the search
for new and more effective treatment continues, and
the development of liposomal drug delivery system to
diminish the toxicity (Tuon etal. 2022). In pets, long-
term treatment with allopurinol alone or combined with
meglumine antimoniate or miltefosine is the treatment of
choice, although the clinical relapse is usual (Baneth and
Solano-Gallego 2022).
Since treatment for both humans and animals do not give
convincing or definitive results, and the emergence of resist-
ance gene in the parasite (Salari etal. 2022), prevention is
essential. In fact, the recent global report on neglected tropi-
cal diseases of the WHO includes cutaneous and visceral
leishmaniasis, and the first pillar in the road map targets for
2030 is reduced incidence and prevalence of this disease
(WHO 2023).
To control and prevention of disease, two strategies
are possible: the effective vaccine and control of disease
transmission. Currently, no vaccines are available for
human use, even though leishmaniasis could be prevent
by vaccination (Kaye etal. 2021). In dogs, four vaccines
have been marketed (Velez and Gállego 2020), although
currently only one is authorized,, with the recent sus-
pension of Leish-Tec for the Ministry of Agriculture
and Livestock (MAPA). This vaccine, LetiFend®, was
licensed in Europe in 2016, and are relative efficacy, 72%
(Reguera etal. 2016).
In this scenario, prevention methods for the proliferation
of sandfly vector and disease transmission for human and
other animals become very relevant. The barriers systems
and system control of vertebrate hosts have proven to be
the most effective measures (de Vries and Schallig 2022;
Kumosani etal. 2022). The use of deltamethrin-impregned
collars could be the one more effective prevention method
for canine leishmaniosis, reducing the sand fly feeding
around 94% (de Camargo-Neves etal. 2021; Evans etal.
2021; Paulin etal. 2018).
Certainly, a greater knowledge of the species involved in
the transmission of the parasite and its prevalence worldwide
are essential for the eradication of this zoonosis, which is
increasingly relevant.
Table 2 (continued)
Subgenus Species Countries Vertebrate Host Reference
O. Anseriformes: Greylag Goose (Anser answer), Muscovy
Duck (Cairina moschata)
Otranto etal. (2010)
O. Galliformes: Common Pheasant (Phasianus colchicus),
Chicken (Gallus gallus) Helmeted Guineafowl (Numida
meleagridis)
Otranto etal. (2010)
O. Reptilia: Geckoes (Tarentola mauritanica), lizard (Podarcis
filfolensis)
Mendoza-Roldan etal. (2021, 2022)
L. major Portugal, Turkey O. Primates: Human (Homo sapiens) Ravel etal. (2006)
O. Carnivora: Cat (Felis catus) Paşa etal. (2015); Pereira etal. (2020)
L. tropica Turkey O. Primates: Human (Homo sapiens) Toz etal. (2009)
O. Carnivora: Cat (Felis catus) Paşa etal. (2015)
L. siamensis Switzerland O. Artiodactyla: Cattle (Bos taurus) Lobsiger etal. (2010)
Sauroleishmania L. tarentolae Italy O. Primates: Human (Homo sapiens) Pombi etal. (2020)
O. Carnivora: Dog (Canis lupus familiaris) Iatta etal. (2021a)
O. Reptilia: Geckoes (Tarentola mauritanica), lizard (Podarcis
filfolensis)
Mendoza-Roldan etal. (2021, 2022)
Veterinary Research Communications
1 3
Conclusions
Leishmaniasis is a complex and wide-ranging disease that
affects a variety of hosts, including humans, companion
animals, livestock, and wildlife in many regions of the
world. Environmental changes, such as deforestation and
climatic change, are affecting the distribution of sand flies,
resulting in the expansion of the disease, and making trans-
mission to new hosts/reservoirs possible. Efforts to pre-
vent and control the spread of the disease in wild animal
populations are important for protecting both animal and
human health. Companion animals, including dogs, cats,
and livestock, including horses, bovines, and others, in
endemic areas, should be evaluated for leishmaniosis, and
guardians/owners/tutors should provide protection against
sand fly bites with the use of preventive measures.
In conclusion, leishmaniasis is a significant global health
concern that requires ongoing research and collaboration to
improve our understanding of the disease and develop effec-
tive prevention and treatment strategies. The finding of a new
wild species naturally infected by Leishmania spp. indicates
the importance of investigating infection by these parasites in
wild animals of different species to identify new hosts and
evaluate the possibility of these acting as reservoirs, contrib-
uting to a better understanding of the transmission cycles of
leishmaniasis and consequently to its control.
Acknowledgements We are grateful to University Cardenal Herrera-CEU.
Authors’ contributions E. M-A. wrote the first manuscript. L. L. con-
ceived the research, wrote, and revised the final version of manuscript.
All authors read and approved the final manuscript.
Funding This work was supported by IDOC (IDOC22-05) and
PUENTE (PUENTE22-01) projects of University CEU Cardenal Her-
rera. The funding agency was not involved in the study design, data
collection, data analysis or drafting of the manuscript.
Data availability Not applicable.
Declarations
Ethics approval and consent to participate Not applicable.
Consent for publication Not applicable.
Competing interests The authors declare that they have no competing
interests.
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