International Journal for Parasitology 44 (2014) 955967

Contents lists available at ScienceDirect

International Journal for Parasitology

journal homepage: www.elsevier.com/locate/ijpara

Invited Review

Parasitic mites of medical and veterinary importance is there

a common research agenda?
Katja Fischer a,, Shelley Walton b,
a
QIMR Berghofer Medical Research Institute, Infectious Diseases Program, Biology Department, Brisbane, Queensland, Australia
b
Inammation and Healing Research Cluster, School of Health and Sport Sciences, Faculty of Science, Health, Education and Engineering, University of the Sunshine Coast,
Sippy Downs, Queensland, Australia

a r t i c l e i n f o a b s t r a c t

Article history: There are an estimated 0.51 million mite species on earth. Among the many mites that are known to
Received 24 July 2014 affect humans and animals, only a subset are parasitic but these can cause signicant disease. We aim
Received in revised form 22 August 2014 here to provide an overview of the most recent work in this eld in order to identify common biological
Accepted 23 August 2014
features of these parasites and to inform common strategies for future research. There is a critical need
Available online 16 September 2014
for diagnostic tools to allow for better surveillance and for drugs tailored specically to the respective
parasites. Multi-omics approaches represent a logical and timely strategy to identify the appropriate
Keywords:
mite molecules. Recent advances in sequencing technology enable us to generate de novo genome
Acari
Parasitic mite
sequence data, even from limited DNA resources. Consequently, the eld of mite genomics has recently
Skin infection emerged and will now rapidly expand, which is a particular advantage for parasitic mites that cannot be
Pruritus cultured in vitro. Investigations of the microbiota associated with mites will elucidate the link between
Pyoderma parasites and pathogens, and dene the role of the mite in transmission and pathogenesis. The databases
generated will provide the crucial knowledge essential to design novel diagnostic tools, control measures,
prophylaxes, drugs and immunotherapies against the mites and associated secondary infections.
2014 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.

1. Introduction approximately 85% of them likely having an allergy to house dust

Among the Chelicerates, the Acari (mites and ticks) represent
the largest and most diverse taxon, with an estimated 0.51 mil-
lion species in total. Of over 48,000 species described to date
(Halliday et al., 2000), the vast majority are mites, with ticks
accounting for approximately 1,000 species. Mites inhabit fasci-
nating combinations of diverse ecological niches and lifestyles,
ranging from free-living, predatory or plant-feeding, to obligate
parasitic. There are major agricultural mite pests such as the free
living cosmopolitan plant feeding spider mite Tetranychus urticae,
which is known to feed on more than 1,100 plant species. Free
living allergy-causing dust mites are likely the best studied acarid
pathogens. An estimated 1015% of individuals in the Western
world suffer from asthma (Basagana et al., 2004), with
Corresponding authors at: QIMR Berghofer Medical Research Institute, P.O.
Royal Brisbane Hospital QLD 4029, Australia. Tel.: +61 7 33620417 (K. Fischer).
Inammation and Healing Research Cluster, School of Health and Sport Sciences,
Faculty of Science, Health, Education and Engineering, University of the Sunshine
Coast, Locked Bag 4, Maroochydore DC, Sippy Downs, QLD 4558, Australia. Tel.: +61
7 54302826 (S. Walton).
E-mail addresses: katja.scher@qimrberghofer.edu.au (K. Fischer),
swalton1@usc.edu.au (S. Walton).
mites (Thomas et al., 2010). Several parasitic mites, foremost
scabies mites and mite vectors of scrub typhus, are also of major
concern to human health, particularly in socioeconomically
disadvantaged communities. In addition, scabies mites and the
related sheep scab mites cause a considerable global burden of
disease in livestock and wildlife. There are many more parasitic
mites that are of medical and veterinary concern and which,
taken together, represent a signicant liability to both humans
and animals. Despite this, apart from the relatively narrow focus
on asthma-associated mite allergens, the research input into mite
borne diseases is minute. This review features solely those
parasitic mites that are clinically most relevant to humans and
domestic animals, hence it presents the tip of the iceberg. The
true range of parasitic mites is much larger. As this emerging
research eld receives increased recognition, the picture may
likely become more complete, yet more complex, in the future.
With the aim to identify common research goals in this eld,
the focus of this review is to align the most signicant parasitic
mite species of medical and veterinary importance, with regards
to their biology, the diseases they cause and the status of our
knowledge about them.

http://dx.doi.org/10.1016/j.ijpara.2014.08.003
0020-7519/ 2014 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.
956 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
2. Parasitic mites of medical and veterinary importance
2.1. Scabies and mange
Scabies is a skin infestation caused by the obligate parasitic
mite Sarcoptes scabiei. This mite burrows into the upper layers of
the skin of a wide range of mammalian hosts, including humans,
feeding on epidermal cells and serum (reviewed in Holt et al.,
2013). Scabies affects both humans and animals, with different
host-specic varieties (pathovars) of S. scabiei (Bornstein and
Samuel, 2001), such as S. scabiei var. hominis from humans, S. sca-
biei var. canis from dogs and S. scabiei var. suis from pigs. Early
studies reported on the limited capability of different pathovars
to survive on non-natural hosts, although an ongoing infestation
of dog mites on rabbits was established (Arlian et al., 1985,
1988b). More recent molecular epidemiology and phylogenetic
studies (Alasaad et al., 2013) (also reviewed in Holt and Fischer,
2013; Alasaad et al., 2014) clearly showed host-specic mite pop-
ulations. Importantly, no evidence of cross-transmission was iden-
tied between sympatric mite populations obtained from humans
and dogs living in proximity (Walton et al., 1999, 2004b). Occa-
sional infestations of humans with S. scabiei of animal origin have
been reported but appear to be short-lived and self-limiting
(Menzano et al., 2004; Bazargani et al., 2007).
2.1.1. Sarcoptes scabiei var hominis
Scabies is listed among the top 50 most prevalent diseases
worldwide, with a global prevalence of 100,625,000 in 2010
(1.5% of the world population) (Hay et al., 2014). Scabies is one
of the three most common skin disorders in children, together with
tinea and pyoderma (Andrews et al., 2009; Vos et al., 2012), and
imposes a considerable economic burden on individuals, families,
communities and health systems (Fuller, 2013; Heukelbach et al.,
2013). Owusu-Edusei et al. (2009) estimated the annual economic
burden for scabies management as US $10.4 million (Owusu-
Edusei et al., 2009). Recently the World Health Organization added
scabies to the list of Neglected Tropical Diseases, thereby recogn-
ising its impact on human health. The International Alliance for the
Control of Scabies, a newly formed organisation, proposes to
accomplish scabies control in vulnerable communities (Engelman
et al., 2013).
The life cycle of S. scabiei has four parasitic stages (egg, larva,
nymph and adult) and takes approximately 2 weeks. The adult
female mite burrows into the stratum corneum and stratum gran-
ulosum of its host (Levi et al., 2011), where she lays two to three
eggs per day. Burrowing is achieved using mouth parts, special cut-
ting surfaces on the front legs, and enzymatic secretions. The mite
releases allergenic metabolic products into the burrows, from
where they diffuse into the underlying dermis to provoke strong
hypersensitive reactions and intense itching. As a result, scratching
and disruption of the upper epidermal protective matrix ensues,
promoting infection by opportunistic bacteria. These can lead to
serious health complications. In northern Australian remote Indig-
enous communities, very high rates of scabies-associated strepto-
coccal and staphylococcal infections have been documented
(Carapetis et al., 1997; Carapetis and Currie, 1996; Lin et al.,
2009; Steer et al., 2009; Subramaniam et al., 2010). Consequently,
in populations where scabies is endemic, severe sequelae such as
acute rheumatic fever, rheumatic heart disease and acute post
streptococcal glomerulonephritis have been reported (Carapetis
et al., 1997; Carapetis and Currie, 1996; and reviewed in Hay
et al., 2013).
The healthy host immune system is able to limit mite numbers
in primary infestations to 1020 adult mites, and with subsequent
re-infestations mite numbers are reported to drop to less than ve.
Occasionally, the host becomes completely protected (Mellanby,
1944; Rodriguez-Cadenas et al., 2010). Recently, extracts derived
from scabies mites have been shown to modulate cytokine expres-
sion by keratinocytes, broblasts, dendritic cells and peripheral
blood mononuclear cells, thereby initiating inammation (Arlian
et al., 2004; Morgan and Arlian, 2010; Morgan et al., 2013). Crusted
scabies, a severe manifestation of the disease, occurs in people
with a non-protective immune response to the mite (Roberts
et al., 2005) and is a rare manifestation characterised by hyper-
infestation of the skin with thousands of mites/g of skin, often
leading to sepsis and death. Increased localised skin IL-17 secreting
T cells may play a critical role in the pathogenesis of crusted sca-
bies development (Liu et al., in press) and, although the underlying
mechanisms are still unknown, support a hypothesis that regula-
tory T cell function may be impaired. Crusted scabies mainly
occurs in immune compromised individuals or in the elderly, and
is frequently observed in nursing homes or institutions where they
often act as core transmitters (Lokuge et al., 2014). Those affected
suffer social stigma and often remain undetected and untreated for
a long time, thereby presenting a hidden reservoir of parasites
(Roberts et al., 2005; Currie and McCarthy, 2010).
Ten years ago the molecular biology of scabies was largely
unexplored. Since then signicant progress has been made on
characterising a number of S. scabiei antigens/allergens and deter-
mining their role in the mite as well as their role in immunopatho-
genesis (Mattsson et al., 2001; Pettersson et al., 2005; Mounsey
et al., 2013). This has been possible in large part due to the gener-
ation of a cDNA database of approximately 43,000 cDNA sequences
from scabies mites collected from a human patient (Fischer et al.,
2003a). This database enabled the discovery of homologs to house
dust mite allergens (Fischer et al., 2003a; Holt et al., 2003, 2004;
Dougall et al., 2005), genes implicated in drug resistance
(Mounsey et al., 2006, 2007, 2010b; Pasay et al., 2006, 2008), pro-
teins with immunodiagnostic potential (Harumal et al., 2003;
Walton et al., 2010; Jayaraj et al., 2011; Rampton et al., 2013),
and families of molecules which may have a role in pathogenesis
(Beckham et al., 2009; Bergstrom et al., 2009; Fischer et al.,
2009; Holt et al., 2003, 2004; Mika et al., 2011, 2012a, 2012c;
Mahmood et al., 2013). Sarcoptes scabiei recombinant proteins are
now being produced that are leading to advances in understanding
the biology of the mite and protective and aberrant immune
responses observed in scabies, and potentially novel therapeutic
avenues for patients (Walton, 2010; Zhang et al., 2012; Gu et al.,
2014a; Liu et al., 2014).
Upon infecting the epidermal layers, the mite would be exposed
to serum components that diffuse in from underlying dermal
microvasculature (Rapp et al., 2006). The mite ingests these com-
ponents (Rapp et al., 2006; Bergstrom et al., 2009; Walton et al.,
2010; Mika et al., 2011) and consequently must defend its intesti-
nal system against the onslaught of these host defence systems.
Complement is a crucial and rst line host defence system that is
activated upon contact of a microbe with host body uids. It is a
complex network involving approximately 40 proteins. Every suc-
cessful parasite must have defence mechanisms against this sys-
tem. Similar to many haematophagous arthropods, the scabies
mite has evolved a sophisticated arsenal of mechanisms to avoid
complement-mediated gut damage. Mite complement inhibitors,
specically serpins (Mika et al., 2012a) and Scabies Mite Inacti-
vated Serine Protease Paralogs (SMIPP-Ss) (Bergstrom et al.,
2009; Reynolds et al., 2014) have been identied as molecules
the mite may use to overcome the host defence. Members of the
extensive family of SMIPP-Ss have been shown to inhibit the acti-
vation of host complement (Bergstrom et al., 2009), particularly
the lectin pathway (Reynolds et al., 2014). Scabies mite serpins
have been documented to reduce the deposition of several
complement components, thereby blocking the human
 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
complement system (Mika et al., 2012a), and one of those (SMSB4)
has been shown to inhibit complement-mediated neutrophil func-
tion (Swe and Fischer, 2014). The mite complement inhibitors have
been shown to be present in the mite gut and released with the
mite faeces into the epidermal burrows. It is hypothesised that
they likely suppress the local host innate immune response in
the mite-infested epidermis and consequently provide favourable
conditions for the growth of co-infecting bacteria (Swe et al., in
press). In support of this increased growth, Streptococcus pyogenes
and Staphylococcus aureus were observed in whole blood bacterici-
dal assays in the presence of mite complement inhibitors (Mika
et al., 2012b; Swe and Fischer, 2014). Furthermore, the inhibitors
have been reported to decrease opsonisation of S. aureus (Swe
and Fischer, 2014) and S. pyogenes (K Fischer, personal communica-
tion), leading to reduced phagocytosis by neutrophils (Swe and
Fischer, 2014).
In a porcine model, a dramatic increase in pathogenic Staphylo-
coccus on the skin correlated with the onset of mite infestation, and
pathogenic Staphylococci persisted beyond treatment with acari-
cide and healing of the skin (Swe et al., 2014). This is in line with
epidemiological evidence of scabies-associated pyoderma
(Whitehall et al., 2013) and a decreased prevalence of impetigo fol-
lowing scabies control initiatives (Carapetis et al., 1997; Lawrence
et al., 2005). As the diversity and dynamics of the microbiota asso-
ciated with S. scabiei var. hominis have not been fully investigated,
it is unknown whether the mite requires endosymbionts for sur-
vival. Previous investigations failed to detect genes encoding Wol-
bachia surface proteins and 16S rRNA (Mounsey et al., 2005). In
ticks, endosymbionts can establish a relationship with pathogenic
bacteria (Noda et al., 1997) and manipulation of the endos-
ymbionts has been proposed as a novel means of preventing
insects from vectoring diseases (Beard et al., 2002). Thus, knowl-
edge of the microbiota associated with S. scabiei and its roles and
functions in mite biology could lead to new approaches for control.
Scabies infestations are difcult to diagnose clinically as they
are often obscured by eczema or impetigo, or are atypical
(Walton and Currie, 2007). The identication of mites by micros-
copy of skin scrapings has excellent specicity but low sensitivity
(Hengge et al., 2006). Dermatoscopy represents an interesting,
more recent, method to detect scabies mites in vivo, but is not
yet commonly used. Therefore the true global prevalence of scabies
is likely to be underestimated (Walton and Oprescu, 2013). No
commercial immunodiagnostic or molecular based tests for human
scabies is currently available. Whole-mite antigen preparations
derived from S. scabiei var. suis and the itch-mite of the red fox,
S. scabiei var. vulpes, reported a sensitivity of only 48% for human
IgG, in comparison with 80% in pig scabies and 84% in dog scabies
(Haas et al., 2005; Wells et al., 2012). A major S. scabiei var. hominis
recombinant antigen has been used to develop a highly sensitive
IgE immunodiagnostic test (Jayaraj et al., 2011; Rampton et al.,
2013). If proven efcient in clinical testing this may signal a signif-
icant breakthrough in efforts to develop diagnostic tools, to
improve surveillance and consequently control of scabies. In addi-
tion, a nested PCR method was recently reported that demon-
strated the ability to diagnose clinically suspected scabies and
the potential to determine residual mite infestation in previously
treated patients (Naz et al., 2013).
There are currently limited commercial treatment options
available for the management of scabies, and none have sufcient
ovicidal activity to allow a single treatment, necessitating the need
for repeated treatment to kill newly hatched larvae (Mounsey
et al., 2008). Present treatments include malathion (0.5%), per-
methrin (5%), and ivermectin (200 microgram/kg). Clinical assess-
ment of treatment failure in scabies is problematic, particularly
due to frequent re-infestations and residual symptoms. The inten-
sive use of pyrethroids worldwide has led to the development of
957
resistance in many arthropods, as demonstrated with head lice
(Van Leeuwen et al., 2010). Ivermectin, as the sole oral therapy,
is not approved for children aged <5 years and pregnant women.
Of concern is the reported lack of compliance with treatment
schedules for scabies, leading to increasing evidence of acaricide
tolerance/resistance and treatment failures (Andrews et al.,
2009). Resistance to topical 5% permethrin has been observed in
S. scabiei var. canis mites (Pasay et al., 2006) and increasing toler-
ance observed in S. scabiei var hominis mites (Walton et al.,
2000). Clinical and in vitro resistance of S. scabiei to ivermectin
has been documented in human crusted scabies (Currie et al.,
2004; Mounsey et al., 2009), and treatment failures in dogs
(Terada et al., 2010). Single nucleotide polymorphisms (SNPs) at
target sites in the voltage-sensitive sodium channel (vssc) gene
play a major role in conferring resistance to pyrethroids. A SNP
was identied at codon 733 in the vssc gene associated with a pop-
ulation of in vivo and in vitro permethrin-resistant S. scabiei var.
canis mites (Pasay et al., 2006). This SNP was not identied in per-
methrin-sensitive mites or those collected from treated scabies
patients in Paris, France (Andriantsoanirina et al., 2014) and thus
may be a suitable molecular marker for future use in monitoring
emerging resistance.
Tea tree oil, although not currently marketed as an acaricide,
shows promise as a topical therapy for scabies in preliminary
in vitro studies (Walton et al., 2004c) and as a combination treat-
ment in vivo for crusted scabies (Davis et al., 2013). Tea tree oil
(5%) is miticidal, ovicidal, antibacterial and anti-inammatory, and
thus could block progression to pyoderma and secondary sepsis,
as well as reducing skin hypersensitivity. A clinical trial is required
to asses this agents efcacy in the patient, especially in children.
In addition, well-designed randomised control trials of other acari-
cidal herbal or traditional medicines and the use of prophylactic
measures to prevent the transmission of scabies are greatly needed
(FitzGerald et al., 2014; Strong and Johnstone, 2011).
Another research agenda is the informed design and develop-
ment of acaricides based on molecular studies of mite essential
proteins. Building on the hypothesis that scabies mite intestinal
proteases are essential for mite survival, a number of mite prote-
ases have been a focal point of recent scabies mite molecular
research (reviewed in Holt et al., 2013) with the aim to develop
new treatment options. One serine protease (Beckham et al.,
2009), one aspartic protease (Mahmood et al., 2013) and ve cys-
teine proteases (Holt et al., 2004) were demonstrated to be present
in the mite gut and proposed to have central roles in the digestion
of host proteins, thereby dening them as possibly suitable targets
for drug development.
2.1.2. Animal scabies (mange)
Sarcoptes scabiei causes sarcoptic mange in companion, live-
stock and wild animals. Mange has been reported from 10 orders,
27 families and 104 species of domestic, free-ranging and wild
mammals (Currier et al., 2011). There are no accurate estimates
of the prevalence of sarcoptic mange in many of the different
affected animal populations globally (Alasaad et al., 2013). New
outbreaks in wildlife are being continually reported worldwide
(Holz et al., 2011). Cross-infectivity between animal hosts was rst
postulated in emerging epizootics in sympatric wild animal host
populations, with increased morbidity and mortality observed
with apparent domestic to wildlife transmission. However recent
molecular analyses indicate outbreaks are primarily host-associ-
ated and mites collected from sympatric wild animals in Europe
clearly show a lack of gene ow between carnivore, herbivore
and omnivore host-derived Sarcoptes populations (Oleaga et al.,
2013). It now seems apparent that Sarcoptes is not a single panmic-
tic population and genetic subdivision occurs according to the host
with secondary subclustering related to geographical location
958 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
within host groups, as reported previously (Walton et al., 2004a;
further reviewed in Holt and Fischer, 2013; Alasaad et al., 2014).
Animal molecular epidemiology studies and population dynam-
ics will assist in Sarcoptes vaccine development. Transmission of
the scabies mite is primarily based on the close proximity of the
host. In mange-affected high density animal populations, a vaccine
would be highly advantageous, but conversely of limited use for
sporadic outbreaks. Vaccination is generally most efcacious in
tiny populations facing very high infection rates. The increasing
use of barn raised livestock with a high population density, in par-
allel with the increasing institutional care of the elderly and very
young in human populations, suggest a scabies vaccine may be a
necessity in the future. The development of animal models of sca-
bies will be useful for continued studies of the immune response
and development of resistance as evidenced in the dog mite/rabbit
host (Arlian et al., 1985), porcine mange (Mounsey et al., 2010a),
and rabbit mange models (Casais et al., 2014). This will lead to
the development of novel immunotherapeutic and immunodiag-
nostic measures (Tarigan and Huntley, 2005; Toet et al., in press)
as evidenced in the number of vaccination studies against scabies
already completed (reviewed in Liu et al., 2014). Mouse studies
are currently underway for the development of a scabies DNA vac-
cine (Gu et al., 2014a). However, it seems increasingly apparent
from recent molecular studies that candidate vaccine and diagnos-
tic molecules will need to be tested for efcacy against the differ-
ent host-associated varieties of mites.
2.2. Scab mites (Psoroptes ovis, Psoroptes cuniculi)
Psoroptes mites are parasitic mites that have adapted to feeding
on the surface of the skin by abrading the outer layer and feeding
on the resulting serous exudate. Sheep scab is one of the most
important ectoparasitic diseases of sheep in the United Kingdom
in terms of both animal welfare and the nancial impact on pro-
ducers. Psoroptes infection is endemic in sheep in all areas of the
British Isles, with an estimated 7,000 outbreaks in 2004 (Bisdorff
et al., 2006) and is also a major welfare problem in cattle and goats,
particularly in continental Europe (Jones et al., 2008; Millar et al.,
2011). Dual infestations caused by Psoroptes and Sarcoptes mites
have been observed (Rodriguez-Cadenas et al., 2010), however dif-
ferential diagnoses are difcult due to the lack of diagnostic tools.
The highly contagious disease causes restlessness, scratching,
wool-loss, head tossing, bleeding wounds and loss of condition.
In extreme cases the disease is highly debilitating, causing signi-
cant morbidity and mortality associated with epileptiform-like sei-
zures (van den Broek and Huntley, 2003), and is a major welfare
concern (Kirkwood, 1986; van den Broek and Huntley, 2003).
Sheep scab was eradicated in Australia in the late 1800s after an
extensive quarantine and control effort, having been introduced
with the rst sheep importations in 1788. It remains a notiable
disease in all jurisdictions in Australia due to the serious implica-
tions of reintroduction.
Recent transcriptomic studies on the host response to sheep
scab (Burgess et al., 2010, 2011a, 2012b) have provided key
insights into the inammatory response to infestation. A P. ovis
cDNA microarray (Burgess et al., 2012a), built on a large collection
of P. ovis expressed sequence tags (ESTs) (Burgess et al., 2011b),
will enable future studies to analyse gene expression in this impor-
tant ectoparasite. In terms of development of diagnostic tools, two
major ruminant acute phase proteins (haptoglobin and serum
amyloid A) have been shown to indicate sheep scab infestation
(Wells et al., 2013) and may supplement the recently developed
Pso o 2 serological assay (Nunn et al., 2011; Burgess et al.,
2012c). A series of vaccine candidates has been tested previously
(Smith et al., 2001; Smith and Pettit, 2004; Nisbet and Huntley,
2006; Zheng et al., 2013) and novel approaches to mite control
have been explored, including the use of pathogenic fungi against
P. cuniculi (Perrucci et al., 2008). With the transcriptomic data
now available and the P. ovis mite genome size being identied
as relatively small (Mounsey et al., 2012), a Psoroptes genome pro-
ject seems achievable. In the advent of this, the complete mito-
chondrial genome sequence of P. cuniculi was recently published
(Gu et al., 2014b). A genome database would allow the systematic
selection of potential vaccine candidates or chemotherapeutic tar-
get proteins, which could potentially be reinforced by the use of
RNA interference (RNAi) technology, as previously implemented
for other mites (Khila and Grbic, 2007; Campbell et al., 2010).
2.3. Demodicosis
Demodex mites are burrowing follicle mites feeding on sebum
which produced by the sebaceous glands in hair follicles. Approx-
imately 60 Demodex spp. have been reported in many domestic
mammals as well as in humans.
2.3.1. Human demodicosis (Demodex folliculorum, Demodex brevis)
Humans are colonised by two Demodex spp., D. folliculorum and
D. brevis. These mites are found primarily in the facial epidermis
with the two species differing marginally in sizes and habitats.
Demodex folliculorum is found mainly in the upper segment of
the sebaceous duct of the hair follicle, while D. brevis is mainly
found in the sebaceous and Meibomian glands (Lacey et al.,
2011). They are described as negatively phototaxic but details of
their life cycle remain uncertain (Lacey et al., 2011). Currently
Demodex mites are primarily identied microscopically by skin
scrapings, however non-invasive dermoscopy will likely improve
diagnosis in the future (Segal et al., 2010). Demodex mites can
cause substantial cutaneous barrier disruption when feeding and
have been shown to penetrate the dermis (reviewed in Forton,
2012). Demodex mites accumulate bacteria in their intestinal sys-
tems. As they lack an anus (Desch and Nutting, 1972, 1977), gut
contents including bacteria, digestive enzymes and faeces are
released all at once when a mite dies and decomposes. This process
is thought to trigger an immune reaction, inammation and tissue
damage. When present in low density the mites cause no symp-
toms. However, if they reach high densities Demodex mites may
play a role in promoting rosacea, a common facial skin disease,
known to affect between 5% and 20% of the worlds population
(reviewed in Jarmuda et al., 2012). The characteristic symptoms
of rosacea are erythema (permanent redness) and telangiectasia
(small widened blood vessels visible near the surface of the skin).
In advanced rosacea red papules and pustules develop. While the
pathogenesis of rosacea remains unclear and is likely multifacto-
rial, a recent meta-analysis has shown a statistically signicant
association between Demodex infection and rosacea (Zhao et al.,
2010). Roseceais correlates with a generalised increase in inam-
mation markers and has been described as a vascular system disor-
der (Smith et al., 2007). It can be related to an immune disorder
(Yamasaki et al., 2011) or to immunosuppression due to topical
steroids, cancer chemotherapy, HIV/AIDS infection or chronic dial-
ysis treatment (summarised in Lacey et al., 2011). It is thought that
an increased expression of pattern recognition receptors elicits an
exaggerated immune response against foreign protein, likely the
intruding mites and bacteria. Several molecular features of the
inammatory processes in rosacea have been identied to be an
overproduction of Toll-like receptor 2 (Yamasaki et al., 2011), of
a kallikrein (KLK5) (Yamasaki et al., 2007), and of abnormal forms
of cathelicidin (Yamasaki et al., 2007). The disorder can coincide
with, and is difcult to differentiate from, acne vulgaris and/or
seborrhoeic dermatitis.
Increased levels of Demodex mites often correlate with rosacea-
like symptoms (reviewed in Forton, 2012). In a recent study using
 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
confocal laser scanning microscopy, Demodex mites were detected
in 11/12 rosacea patients (91%), compared with only in 3/10
healthy individuals (30%) (Harmelin et al., 2013). The mean per-
centage of follicles containing Demodex was 22% in the patient
group versus 1% in controls. Conventional parasitological examina-
tion of the same cohorts was positive in only 5/12 patients (45%)
and in none of the healthy individuals, demonstrating the unreli-
ability of current diagnostic methods. Even less data are available
to identify the roles of the mites in disease development. Demodex
mites were positively correlated with an increased activation of
the innate immune system (Lacey et al., 2007; Casas et al., 2012).
Bacillus oleronius was previously isolated by culture from Demodex
mites obtained from a rosacea patient (Lacey et al., 2007). Interest-
ingly, a total of 80% of a cohort of 26 patients with erythematote-
langiectatic rosacea showed immunological reactivity to proteins
from B. oleronius (OReilly et al., 2012a) and further work demon-
strated that several proteins produced by this microbe activated
neutrophils to migrate and to produce inammatory cytokines
(OReilly et al., 2012b). A recent culture-independent 16S rRNA
library sequencing approach compared the microbial communities
of individual mites collected from patients with the two subtypes
of rosacea and healthy volunteers (Murillo et al., 2014). This
groundwork data set revealed a Demodex-specic microbiota
including possible symbiotic bacteria and human pathogens. Inter-
estingly, distinct differences in the Demodex mite-associated mic-
robiota were observed, depending on the hosts disease status.
This pilot study indicated that the microbial balance in the mites
is tightly linked with the mite-triggered inammation in rosacea,
pending conrmation through follow-up work providing increased
sample numbers and sequencing depth.
To date almost no molecular data exist on the Demodex mites
themselves, which limits available diagnostic capabilities as well
as therapeutic measures against the parasite. The current consen-
sus seems to be that Demodex mites normally have a commensal
relationship with humans; they may even full mutualistic roles
in clearing the follicular canal of bacteria and other microbes
(Lacey et al., 2011). They seem to be tolerated by the host innate
immune system or possibly down-regulate it by the release of
yet unknown mite molecules. However, if mite numbers increase
to a critical level, due to an immune defect or external immuno-
suppression in the host, the proliferation of Demodex may promote
the development of roseacea (Forton, 2012). It is possible that
increased mite numbers overwhelm the local innate immune sys-
tem through mechanisms specically targeting complement acti-
vation, as seen in other parasitic mites (reviewed in Fischer et al.,
2012). Identication of the intricate mechanisms of interaction
between the parasites, the bacteria and the host will likely lead
to a better understanding of the disease and to the development
of novel control measures.
2.3.2. Demodectic mange (Demodex canis, Demodex cati)
In animals the pathogenic potential of Demodex mites is well
documented, as demodectic mange is a potentially lethal condition
(Plant et al., 2011). There are multiple host specic Demodex spp. in
domestic and companion animals, among them D. canis being the
most commonly seen in dogs (Ravera et al., 2013), D. cati and Dem-
odex gatoi in cats (Bernstein et al., in press; Frank et al., 2013), and
Demodex caprae, Demodex ovis and Demodex bovis causing skin
nodules in domestic goats, sheep and cattle, respectively (Scott
et al., 2001). Currently diagnosis requires microscopic analysis of
a deep skin scrape, a hair pluck sample or an acetate tape impres-
sion (Pereira et al., 2012), combined with the observation of con-
current clinical signs. Treatment with macrocyclic lactones is the
current therapy (reviewed in Ferrer et al., in press), but non-com-
pliance with long-term treatment regimes is a challenge, as it is
with many other parasitic mite infestations. The fact that Demodex
959
cannot be cultured and no molecular databases exist for this para-
site has been a major limitation to biological investigation. As in
humans, the animal host immune system seems to detect and tol-
erate the mites to a certain extent, and the exact mechanisms of an
activated immune response remain poorly understood (Ferrer
et al., in press). An impact of mite burden on the healthy skin mic-
robiota is equally possible in animals as observed in humans. The
skin microbiome of dogs has been recently investigated
(Rodrigues Hoffmann et al., 2014), hence this aspect of democido-
sis in animals is likely to be assessed in the near future.
2.4. Chiggers (Trombiculidae; Leptotrombidium pallidum and others)
Globally, nearly 3,000 species of Trombiculidae, or harvest mites,
have been reported. They are ubiquitous in hot and humid regions,
and in temperate regions they are more prevalent during summer.
The most common hosts of their parasitic larvae (chiggers) are
small rodents and humans can be accidental hosts (reviewed in
Zhan et al., 2013). Six species of the genus Leptotrombidium trans-
mit chigger-borne rickettsiosis or scrub typhus, caused by the obli-
gate intracellular bacterium Orientia tsutsugamushi. The most
dominant disease-carrying Leptotrombidium spp. are Leptotrombid-
ium pallidum and Leptotrombidium scuttellare, which are endemic in
Asia, the Pacic region and Australia. Accordingly, scrub typhus is
highly prevalent in the AsiaPacic region including Japan, Korea,
the eastern part of Russia, India, Pakistan, the southwestern Pacic
islands (Kelly et al., 2009), China (Zhan et al., 2013) and Australia
(Graves and Stenos, 2009). Mortality rates of up to 40% have been
reported (Tamura et al., 1995), more than one million cases are
recorded annually and more than one billion people are estimated
to be at risk (Kim et al., 2014). Leptotrombidium mites may serve as
vectors for other pathogens affecting humans such as Hantaan
virus (Houck et al., 2001) causing epidemic haemorrhagic fever
and Bartonella tamiae, the causative agent of human bartonellosis
(Kabeya et al., 2010). Infestation with trombiculid mite larvae is
a common cause of dermatitis in domestic animals with Neo-
trombicula autumnalis, Euschoengastia latchmani, Straelensia cynotis
and Walchia americana being the most common mite species
known to cause trombiculidosis in cats, dogs and horses (reviewed
in Le Net et al., 2002; Ramirez et al., 2009).
In contrast to other parasitic mites, chiggers appear to have a
wide range of hosts and a low host specicity. An extremely high
species diversity of chigger mites was reported in Yunnan Province,
China. This was found in an extensive epidemiological study exam-
ining the chigger populations of over 10,000 small mammal indi-
vidual hosts harbouring almost 100,000 chiggers, which were
individually identied microscopically (Zhan et al., 2013). The
study investigated only larval chigger mites which were identied,
based on morphological features, to represent 224 species, 22 gen-
era and three subfamilies of the family Trombiculidae. Leptotrom-
bidium scuttellare and Leptotrombidium deliense, both vectors for
scrub typhus, were identied as the dominant mite species. Never-
theless a single species of rodent could harbour more than 100
morphologically different species of chigger mites. Vice versa, most
chigger mite species identied infested a wide range of hosts. No
genetic analysis or cross-infestation experiments were undertaken
in this study. The low host specicity of these parasites, in combi-
nation with the possible vector role of the mites for multiple infec-
tious diseases, makes these mites potentially signicant zoonotic
agents.
For the major part of their life cycle, Trombiculidae are free-liv-
ing predatory soil dwellers, feeding on small arthropods and their
eggs (Zhan et al., 2013). Only their obligate parasitic larval stage,
the chigger, takes on the vector role for the above-mentioned
bacterial diseases. Compared with the 12 year phase of the free-
living nymphs and adult mites, the parasitic chigger stage is
960 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
relatively short-lived (35 days) and non-burrowing. Chiggers
exhibit an extremely efcient mode of concentrated feeding on
extra-orally digested epidermal and/or occasionally dermal tissue
(Shatrov et al., 2014). This highly nutritive liquid is acquired
through a feeding canal, the stylostome, which is formed by hyper-
keratotic host skin cells within the rst 24 h of attachment. Only
after the tube is built, presumably through mutual interaction
between larval secretions and the epidermal skin cells, does feed-
ing occur. At this stage, the epidermal layer at the site of attach-
ment is thickened, hyperkeratinised and eroded. As scabs are
formed on the skin surface, the hosts terminal dermal blood ves-
sels become increasingly dilated and leukocytes inltrate the area.
Interestingly, chiggers tend to attach closely to each other and
often aggregate on scabs left from previous larvae (Shatrov et al.,
2014). The observed crowding may be a strategy to share the effort
of modifying the host space for efcient and safe feeding. Presum-
ably chiggers, similar to other parasitic mites, must overcome the
onslaught of the hosts innate immune response, possibly through
the release of complement inhibitors as seen in many serum-con-
suming parasitic arthropods including parasitic scabies mites
(reviewed in Fischer et al., 2012).
2.5. Mite dermatitis (Cheyletiella blakei, Ornythonyssus bacoti,
Liponyssoides sanguineus, Dermanyssus gallinae, and others)
The term dermatitis, dened as an inammation of the skin,
characterised by itchy, erythematous, vesicular, weeping and
crusting patches, is broadly applied to a range of common, persis-
tent and often multifactorial skin conditions. The cause of derma-
titis is presumed to be a combination of genetic and multiple
environmental factors, amongst which the role of house dust mite
allergens has been the subject of recent research (reviewed in
Marsella and Samuelson, 2009; Fuiano and Incorvaia, 2012;
Garritsen et al., 2013). Parasitic mites release proteins homologous
to house dust mite allergens, presumably playing a similar role in
dermatitis caused by mite infestations.
2.5.1. Cheyletiella dermatitis
In humans Cheyletiella dermatitis is typically the result of con-
tact with an infected companion pet. In dogs, cats and rabbits
the causative mite species are Cheyletiella yasguri, C. blakei and
Cheyletiella parasitivorax, respectively. The most obvious clinical
signs in infested animals are white, dorsal akes (walking dan-
druff) and possibly crusts, either asymptomatic or causing alope-
cia and pruritus (McClain et al., 2009). Importantly, the severity
of dermatological problems is disproportionate to the number of
mites present. Consequently diagnosis by microscopy is difcult
and treatment is often indicated without direct microscopic evi-
dence (Saevik et al., 2004). Skin conditions in humans due to Chey-
letiella spp. are reported frequently as these mites are highly
contagious to humans. However, Cheyletiella mites do not burrow
or reproduce on human skin, and they have been described to
adopt a bite and run behaviour on the human host (Wagner and
Stallmeister, 2000). Nevertheless, Cheyletiella mite bites on human
skin result in pruritic papules, urticarial weals or excoriated ero-
sions in the body areas that have been in close contact with the
infested pet. In the absence or upon treatment of the infected
pet, symptoms in humans disappear without treatment within
3 weeks (Wagner and Stallmeister, 2000). Cheyletiella mites are
non-burrowing and free-living, but are obligate feeders, on the epi-
dermal surface, on skin debris. They may release allergens, which
would explain pruritic symptoms. Cheyletiella mite eggs are
attached to the hair shaft of the animal host in a characteristic bun-
dle of nely woven egg threads (Saevik et al., 2004). Females can
survive off the host for up to 10 days, and transmission from host
to host via bedding has been described (Wagner and Stallmeister,
2000).
2.5.2. Tropical Rat Mite Dermatitis (O. bacoti)
Tropical Rat Mite Dermatitis is caused by the non-burrowing
haematophagous mite, O. bacoti. This worldwide epizootic animal
mite naturally occurrs on rats, mice and other small rodents. If
their natural host is diminished, O. bacoti mites will feed on
humans. The rst case report of tropical rat mites causing derma-
titis in humans came from Australia, followed by cases in the USA
and in Germany (reviewed in Beck and Folster-Holst, 2009). It has
recently been estimated that approximately 80% of the wild
rodents in Germany are infested by this parasite (Beck and
Folster-Holst, 2009). Compared with other mites discussed earlier,
O. bacoti mites are long-lived and can survive without a host for up
to 6 months. They are considered to be house infesting rather
than infesting individual hosts. Public care facilities, particularly
after extermination of a house mouse or rat problem, have been
reported to be invaded by these mites. In addition, private house-
holds with well-kept rodent pets, as well as modern, well main-
tained animal research facilities (Kelaher et al., 2005), can
function as reservoirs for mite populations. It is only the females
and nymphs that nocturnally feed on the host. Eggs are laid in
nests in the environment and not on the host. Considering its rel-
atively low host specicity and its independence from the host for
most of its life cycle, this mite species occupies a large range of
possible ecological niches and is therefore difcult to target with
control strategies.
2.5.3. House mouse mite (L. sanguineus)
The primary host of L. sanguineus is the common house mouse
and likely other small rodents, serving as reservoir hosts for the
bacterium Rickettsia akari (reviewed in McClain et al., 2009). Only
nymphs and adult mites feed for short periods of 12 h on blood
and can transmit rickettsialpox to humans as accidental hosts
(Krusell et al., 2002; Ozturk et al., 2003). Transovarial transmission
of the bacterium has been reported by Nichols et al. (1952). As for
many other non-human parasitic mites, L. sanguineus can be con-
sidered house infesting and becomes a problem to humans when
a pre-existing mouse population has been eradicated. Liponyssoides
sanguineus was rst described to form clusters in crowded apart-
ment buildings in large cities on the east coast of the USA, but is
likely distributed sylvatically worldwide, with rickettsialpox being
under-reported (Diaz, 2010).
2.5.4. Poultry mite dermatitis (D. gallinae)
The poultry red mite, D. gallinae, is a ubiquitous and likely wide-
spread haematophagous ectoparasitic mite of wild, domestic and
synanthropic birds (Roy et al., 2009). It has a low host specicity
and may also feed on mammalian hosts such as dogs (Di Palma
et al., 2012), horses (Mignon and Losson, 2008), rodents and
humans (Bellanger et al., 2008), in the latter causing dermatologi-
cal problems of varying severity (Caero et al., 2008, 2011). In the
poultry industry this mite can be the cause of a decline in poultry
welfare, manifesting in the form of anaemia, dermatitis, restless-
ness and weight loss, and a decrease in egg production (Bellanger
et al., 2008). Blood spots on the eggs is another typical indicator
for red mite infestation and this can cause a reduction in the mar-
ket value of the eggs. In addition, staff or owners are often affected
by the itching due to transient poultry mite infection. Eradication
is highly challenging due to its short life cycle, its ability to live
off the host for a substantial time with high resistance to starvation
and a suspected resistance to acaricides (Beugnet et al., 1997).
Aside from its economic importance, D. gallinae is of sanitary sig-
nicance, as it may be involved in the transmission either as a res-
ervoir and/or a vector of several pathogens responsible for serious
 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
disease in both animals and humans. Viruses (equine encephalitis
viruses, the fowl pox virus), bacteria responsible for Q fever (Coxi-
ella burnetii) and Salmonella Enteritidis (Salmonella enterica), have
been found to be associated with D. gallinae (reviewed in
Valiente Moro et al., 2007, 2009a). For some of these pathogens,
multiplication within the mites and transovarial passage and
mechanical vector capacity has been demonstrated experimen-
tally, but the conrmation of D. gallinae acting as a natural biolog-
ical vector for these agents is still outstanding. The microbiota
associated with D. gallinae was recently investigated (Valiente
Moro et al., 2009b) with the aim to identify the characteristic bac-
terial composition associated with D. gallinae. Both pathogens and
symbionts were identied. Sequences from pathogenic agents
included Pasteurella multocida, Erysipelothrix rhusiopathiae and
the genus Aerococcus. Dermanyssus gallinae was found to be associ-
ated with Spiroplasma sp., a bacterium that may be symbiotic and
responsible for sexual determination in insects (Tinsley and
Majerus, 2006, 2007) and which has been discussed as a potential
vector control agent against mosquitoes (Humphery-Smith et al.,
1991a,b). Bacteria corresponding to Cardinium and Wolbachia were
identied in phytophagous Tetranychus spp. and have been consid-
ered to be symbiotic (Zhao et al., 2013a,b). This work indicates a
potential to identify targets for a control strategy within the mite
symbiont community. In addition, vaccination is considered a fea-
sible strategy for controlling the haematophagous poultry red
mite. A signicant research effort has been accomplished towards
the identication of vaccine candidates (Wright et al., 2009;
Bartley et al., 2012). A more recent study has reported for the rst
time a mites secretome and transmembranome (Schicht et al.,
2013), providing valuable insights into the related different meta-
bolic pathways of D. gallinae. Multiple molecules were identied
that putatively play a role in blood feeding and which may signif-
icantly contribute to the development of new vaccines or thera-
peutic targets against this poultry pest.
3. Common features of parasitic mites
All mites known to parasitize vertebrates, humans included,
affect the epidermal layers of the skin, causing dermatological con-
ditions of varying severity. Parasitic mites often trigger allergic
reactions and they can act as vectors for disease transmission or
allow secondary infections with opportunistic pathogens. These
clinical manifestations of mite-associated disease are important
to recognise and to emphasise, as they may easily be confused with
other dermatological conditions and may lead to severe systemic
complications.
As the vertebrate skin barrier, the epidermis is crucial for sur-
vival in providing a cool, acidic and desiccated barrier that resists
penetration by microbes while retaining moisture and nutrients
(Madison, 2003). The epidermal skin layers continuously renew
through the process of epidermal differentiation (Elias, 2005) and
this desquamation constantly eliminates potential microbial
intruders. Burrowing mites manage to evade separation from the
host through desquamation by persevering and feeding within
the moist and nutritious stratum granulosum. The particular
mechanisms allowing them to maintain themselves are still
unknown, however for P. ovis infestation, evidence of a potentially
deleterious effect of mite allergens on epidermal differentiation
has been recently provided (Stoeckli et al., 2013). Apart from the
mites, the skin surface is colonised by a huge variety of microbes
including bacteria, fungi and viruses. An amazing number of
approximately one billion bacteria have been estimated to inhabit
a typical square centimetre of human skin (Grice et al., 2008), con-
sisting of resident and transient microbes, many of those being
both commensals and potential pathogens, and taking on shifting
roles, ranging from host-benecial to pathogenic, depending on
961
the status of their skin environment (Cogen et al., 2008). Parasitic
mites have co-evolved with their vertebrate hosts and co-existing
microbes over millions of years.
Among the parasitic mites of medical and veterinary impor-
tance that affect the vertebrate host skin, there is a fascinating
range from ecto- to endo-parasitic lifestyle (Table 1). The mites
that burrow are obligate parasites and do not survive for long off
the host, which unfortunately means that they cannot be easily
grown in vitro. This has dramatically limited experimental
research. While parasitic mites take on different niches in the hab-
itat epidermis, they all rely on the same nourishment, which is
epidermal tissue containing skin cells and their debris and intersti-
tial uid, including serum. For example, the feeding strategies of
chiggers and scabies mites are completely different from one
another, but we observe a similar host response to either infesta-
tion in the form of hyperkeratosis, scab formation, neutrophil inl-
tration and inammation. All serum-ingesting mites are exposed
to their hosts antibodies, complement and coagulation systems.
Many of them must cope with the constant renewal of the epider-
mal layers through epidermal desquamation. Finally, they all
encounter other microbes present on the skin including bacteria
and fungi. While skin microbiota vary enormously depending on
body site, age, sex, climate and many more factors (Grice et al.,
2009), skin-infesting mites may share some general strategies with
regard to coexisting with the rest of the skin microora. As they
have a common environment, they may share common vulnerabil-
ities which could be utilised by research aimed at developing strat-
egies of control.
4. Common challenges and research goals
Mite infestations have a negative impact on the quality of life
for infected individuals. Despite some major advances in scabies
biomedical research during recent years, overall there appears to
have been little progress in controlling mites at a global level. A
major problem with current evaluations of the epidemiology of
disease caused by parasitic mites is that the data are often anec-
dotal, sporadic and reliant on the informed views of professionals,
dermatologists and veterinarians. As parasitic mites have common
features, they pose common challenges. The development of sensi-
tive and cheap diagnostic tests is critical to replace complicated,
time-consuming and inefcient mite collection and microscopic/
taxonomic analysis. This will impact positively on disease surveil-
lance, which is currently insufcient for all mite-related diseases,
due to the lack of easy-to-use and affordable diagnostic tools. If
linked to better reporting systems, then some of the issues sur-
rounding disease impact, epidemiology and control may be
resolved. With input from new technologies and increasing identi-
cation of disease biomarkers for ectoparasites, such as specic
signalling pathways during the host response to early infestation
(Toet et al., in press; Wells et al., 2012; Morgan et al., 2013), more
cost-effective, rapid and accurate diagnosis to detect the main par-
asitic mites will likely be developed.
In vivo culture of parasitic mites will always be difcult to
establish. While the free living T. urticae and D. pteronyssinus can
be cultured in vitro, the obligate parasitic lifestyles of S. scabiei,
Demodex spp. and P. ovis preclude successful in vitro culture. As a
result, experimental studies are extremely challenging. A trans-
species animal model (dog scabies mites on rabbits) (Arlian et al.,
1985, 1988a,b) has previously provided valuable insights into the
immunopathogenesis of scabies. More recently, natural host infes-
tation models have been developed for scabies and for sheep scab.
However they involve large animal studies, e.g. pigs in the case of S.
scabiei (Mounsey et al., 2010a; Swe et al., 2014) and sheep for P.
ovis (Burgess et al., 2010), which are logistically difcult and
expensive, thus experiments need to be targeted to a high degree.
962 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967

Table 1
Mites of medical and veterinary importance.

Mite species Ecological Feed Geographic Host Clinical disease Secondary Vector Acaricide Previous
niche distribution specicity bacterial role resistance molecular
infections reported approaches
Sarcoptes scabiei Obligate Epidermis, serum Worldwide High (H) Scabies; (A) Yes No Yes cDNA
parasite Mange database
burrowing in microbiota
epidermis genome size
Demodex spec. Obligate Epidermis, sebum Worldwide High (H) Demodicidosis Yes No No Microbiota
parasite (H) Rosacea (?)
burrowing in
epidermis
Psoroptes spec. Obligate Epidermis,blood Worldwide Low (A) Sheep scab (H) Yes No Yes Microbiota
parasite, non- Dermatitis symbionts
burrowing transcriptome
genome size
TrombiculidaeHarvest Free living, Epidermis,dermis, Europe Low (H) Scrub itch; Yes No No
mites soil dwellers; serum Western
predatory on hemisphere
insects, only
larvae are
ectoparasites
of small
rodents
Leptotrombidium spec Southeast (H) Scrub typhus Yes Yes Genome size
Asia, Japan, (tsutsugamushi
Philippines, disease)
South
Pacic, and
Australia
Cheyletiella spec. Free living Epidermal debris Low (H) Dermatitis (A) No No No
ectoparsites of on skin surface Walking dandruff
dogs, cats,
rabbits
Ornythonyssus bacoti, Free living Blood Temperate Low (H) Dermatitis Yes Noa No
ectoparasites and tropical
of large worldwide
rodents
house
infesting
Liponyssoides Free living, Blood USA, Low (H) Mite bite Yes Yes No
sanguineus nymphs and Northern asymptomatic
adults are Europe, Rickettsialpox
ectoparasites Asia, Africa
of mice
Dermanyssus gallinae Free living Blood Worldwide Low (Aa) Equine No Noa No Microbiota
ectoparasites encephalitis Symbionts
of birds viruses, fowl pox transcriptome
virus, Coxiella
burnetii,
Salmonella
Enteritidis, (H)
Dermatitis,
asthma
Dust mites Free living, Epidermal debris Worldwide na (H) Asthma, No No No Genome size
associated in dust rhinitis,
with humans anaphylaxis;
house exacerbates atopic
infesting dermatitis,

(H), human; (A), animal; na, not applicable.

a
Pathogen has been isolated but vectorial capacity of the mite has not been established.

Biomedical research into host-parasite relationships and the
host response is progressing slowly, but increased knowledge of
mite biology and molecular mechanisms is crucial in aiding the
development of specic therapeutic control measures. Under-
standing the complex network of innate and adaptive immune
responses, including the long lag period from infection to clinical
symptoms, is fundamental to developing early interventions and
decreasing transmission.
A lack of available genome sequence data is currently hamper-
ing research of the major parasitic mite species. However genome
size estimations for scabies mites, sheep scab mites, house dust
mites and chiggers exist (Mounsey et al., 2012; Kim et al., 2014),
and they are comparable to the 90 Mb complete genome of T. urti-
cae, indicating the feasibility of a full genome project of major par-
asitic mites.
5. Common strategies
The publication of the rst complete acarid genome sequence,
from the two spotted spider mite, T. urticae (Grbic et al., 2011),
marked the advent of mite genomics. This followed earlier work
undertaking a preliminary genome survey of the honey bee mite,
Varroa destructor (Cornman et al., 2010). These important agricul-
tural pests are just two of a large number of free living mite species
 K. Fischer, S. Walton / International Journal for Parasitology 44 (2014) 955967
which have a signicant impact on the health of humans, other
animals and plants. Metaseiulus occidentalis should be mentioned
as well as a highly effective predatory mite, feeding on pest mites
in agricultural crops around the world Its transcriptome (Hoy et al.,
2013) and mitochondrial genome (Dermauw et al., 2010) are
sequenced, and its nuclear genome is small at 88 Mb
(Jeyaprakash and Hoy, 2009). This mite is potentially useful as a
biological control agent and the established molecular data may
feed into improved pest mite management.
As T. urticae is easily cultured in vitro, it seems to have taken on
the role as a candidate model organism for mites, as did Caenorhab-
ditis elegans for helminths. With RNAi established in T. urticae
(Khila and Grbic, 2007) and its genome completely assembled, it
is hoped that similar progress in S. scabiei, D. pteronyssinus, P. ovis
and Leptotrombidium spp. will soon ensue. Transcriptomic studies
undertaken for P. ovis (Burgess et al., 2011b, 2012a), D. gallinae
(Schicht et al., 2014) and S. scabiei var. hominis (Fischer et al.,
2003a, 2003b) (Table 1), may soon be complemented by proteo-
mics, to decipher entire molecular frameworks of metabolic and
pathogenic mechanisms of these parasites.
The coexistance of microbes and arthropods has been empha-
sised frequently, for example by Dillon and Dillon (2004). Bacteria
may be a food source for the mite (Hogg and Lehane, 1999), they
may also supply minerals and vitamins, x nitrogen or degrade
pesticides (as discussed in Moro et al., 2011). If essential to mite
survival, they could represent potential targets for intervention
(Hogg and Lehane, 1999) or they could serve as a surrogate marker
for diagnostic tests. There are reports of potentially symbiotic bac-
teria in P. ovis (Hogg and Lehane, 1999, 2001), and poultry mites
(De Luna et al., 2009; Valiente Moro et al., 2009b). Uncharacterised
endosymbionts also found in tick species have been detected in
Dermatophagoides farinae (Valerio et al., 2005) and Cardinium has
been found in a wide distribution of arthropods, including mites
(Zchori-Fein and Perlman, 2004). Transovarial transmission of
pathogens has been observed in chiggers (Phasomkusolsil et al.,
2009; Shin et al., 2014), the house mouse mite L. sanguineus
(reviewed in McClain et al., 2009) O. bacoti (reviewed in Chung
et al., 1998) and poultry mites (Valiente Moro et al., 2009a). Finally,
it has been observed in experimental scabies infections that para-
sitic mites can inuence healthy skin microbiota, causing a shift
from commensals to pathogens at the affected site (Swe et al.,
2014b). This simple experiment exemplied that it is important
to integrate the microbiota into the picture of the parasite-host-
pathogen triangle.
6. Conclusion
Mite-associated disease seems to be caused by a large variety of
mite species, and overall this is not a well-studied research area.
There are only a few mite-transmitted dermatoses caused by mites
that specically parasitise humans (scabies, democidosis), and
many more mite species with a wider host range, some of which
can also affect humans. All parasitic mites known to date which
affect animals are associated with cutaneous disease. They are
mostly tissue-juice feeders but there are some blood feeders. All
cause dermatoses and allergies of varying severity, and their path-
ogenicity seems to depend on the immunological status of the
host. Many parasitic mites promote secondary bacterial infections,
but from what is currently known they do not seem to widely
transmit many infectious microbial diseases. Currently, we know
of only two non-human animal mites transmitting infectious dis-
eases: trombiculid larval mites may transmit scrub typhus and
house mouse mites may transmit rickettsialpox. However the roles
of the various mites, either as reservoirs or as vectors for patho-
gens, are only starting to be revealed either experimentally or
963
through microbiota projects. Most mites with environmental
phases during their life cycle exhibit close associations with their
mite islands, i.e. their ecosystems and zoonotic reservoirs. These
are frequented by their natural primary hosts. Humans and domes-
tic animals become accidental hosts when in proximity with these
mite islands or the primary hosts.
As a result of reviewing parasitic mites in humans and animals
we argue that the combined burden of the many mite infestations
recorded is grossly underestimated. The lack of diagnostic tools
may be a major contributor to this. Only a limited number of aca-
ricides are available and the emergence of drug resistance may
become a major challenge in the future, warranting a multi-
pronged approach in this eld.
Acknowledgements
We wish to dedicate this work to the late Prof. Dave J. Kemp,
who championed the work on molecular aspects of scabies, a
neglected disease particularly of the poor. Katja Fischer is sup-
ported by an Australian Research Council Future Fellowship.
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