TAXONOMY, such as Paecilimyces variotii and others, which



Purpureoicllium lilacinum (Thom) Luangsa-ard, Houbraken, Hywel-Jones
& Samson, comb. nov 2011, previously known
as Paecilomyces lilacinus,
described by Samson 1974) is one of the representative and widely
studied species of this group. The
fungus Purpureocillium spp. is
a saprophytic fungus under the family ophiocordycipitaceace of the order
Hypocreales. Different species of genus Paecilomyces have been
identified such as Paecilimyces variotii and others, which used to be
considered close relatives of P. lilacinum, but recent molecular and morphological study suggested that Paecilomyces
lilacinus and Purpureocillium lilacinum belongs to different orders Eurotiales
and Hpocreales respectively(Deng et al. 2012; Luangsa-Ard et al.
2011). P.
lilacinum is a ubiquitous, saprophytic, filamentous fungus with worldwide distribution
around and has wide range of habitats. This fungus has been isolated from
different materials and soils such as grassland, forest, cultivated and
uncultivated lands, desert, estuarine sediments and sewage and sludge, insects,
nematodes and human beings. This fungus can degrade the grain quality, food and
papers, present in the laboratory air as a contaminant, and contaminate the
cosmetic items such as skin creams, lotions and other clinical materials such
as plastic implants, catheters and others (Saberhagen
et al 1997; Saghrouni et al. 2013).

P. lilacinum is widely known as the nematophagous fungus, which has been detected mostly in the females of cyst and root knot
nematode such as Meloidogyne species and
other cyst nematodes and found to have antagonistic effect on nematodes. This
fungus has also been detected in the rhizosphere of the cereals crops such as
wheat and barley and also in decayed grape fruits (Denga et al. 2012; Rumbos et
al 2006). Purpureocillium species can survive in wide range of the
temperature and pH. The maximum temperature ranges from 8 to 38 0C
and optimum temperature range from 26 to 30 0C
(79 to 86 °F). Purpureocillium is very adaptable for its lifestyle
i.e. it may acts as entomopathogenic, mycoparasitic, saprophytic as well as
nematophagous based on the availability. P.
lilacinum produces a mycotoxin called Paecilotoxin, although its
significance is unknown and its production varies according to its strains
(Khan et al. 2003)

Taxonomy and Phylogeny of Purpureocillium

Scientific classification


Division: Ascomycota

Class: Sordariomycetes

Order: Hypocreales

Family: Ophiocordycipitaceae

Genus: Purpureocillium


The American mycologist Charles Thom, first described this
fungus in 1910, named as Penicillium. lilacinum, later in 1974 Robert A. Samson changed the name to Paecilomyces lilaciuns (Samson et al. 1974).
In the 2000s decade the genus Paecilomyces was described as a
non-monophyletic genera and Paecilomyces
nostocoides, Isaria takamizusanensis and Nomuraea atypicola were the close relatives
of Paecilomyeces lilacinus. Luangsa-ard et al. in 2011 again re-classified the genus and changed its
name to Purpureocillium lilacinum
based on the characteristic purple-colored conidia produced by this species
along with the morphological and molecular differences from Paecilomyces
lilaciuns. Previously,
the Purpureocillium lilacinum used to be considered as fungi imperfecti
and Duteromycetes because of lacking the proper sexual reproduction, but recent
whole genome sequencing and other phylogenetic analysis confirmed this fungus
belongs to ascomycetes (Fig. 1) (Prasad et al. 2015). Many species of the Paecilomyces
have been described around the world and variation observed within the species
(Fig. 2). Paecilomyces can be differentiated from Penicillium by
its phialides, which taper into a long distinct neck and produces divergent or
tangled conidial chains, while the Penicillium produces the
symmetrically branched phialides (Rumbos et al. 2006). Luangsa-Ard et al. (2011) described both
morphological and molecular characteristics of the new combination of P.
lilacinum. Based on phylogenetic analysis of the partial 18S rRNA genes,
the type species P. lilacinum belongs to Ophiocordycipitaceae,
while another type species Paecilomyces variotii of Paecilomyces belongs
to Trichocomaceae (Fig. 3) (Luangsa-Ard et al. 2011).

ITS and partial
translation elongation factor 1-? (TEF) genes were also used to identify the Penicillium adametzioides and P. lilacinum, morphologically both
species were found to be identical as the previous descriptions by Deng et al. (2012).
The phylogenetic analysis of the 18s ribosomal RNA (rRNA) genes of these two-species
revealed that these two species are actually not exactly similar because they
belonged to different families. Paecilomyces adametzioides belongs to the family Trichocomaceae, whereas P.
lilacinus moved to the new family Ophiocordycipitaceae as a new
genus Purpureocillium and renamed as a P. lilacinum (Fig. 2). Another phylogenetic analysis of
5.8S rDNA and internal transcribed spacer (ITS1 and ITS2) sequences from few
entomogenous Paecilomyces species
supported the polyphyly of the genus moreover; analysis showed the existence
these cryptic species in two orders such as Eurotiales and Hypocreales. Paecilomyces
variotii and Paecilomyces leycettanus belonged to Eurotiales same as
other species such as Talaromyces and Thermoascus. Another order
Hypocreales, three major subgroups were observed and one of which included Paecilomyces
virdis, Paecilomyces penicillatus, Paecilomyces carneus and
other two such as Paecilomyces lilacinus
and Paecilomyces marquandii (Fig.
2). However, the majority of the P. lilacinus and P.
marquandii isolates formed a distinct and distantly related subgroup, while
the other major subgroup contained Paecilomyces farinosus, Paecilomyces amoeneroseus, Paecilomyces fumosoroseus and Paecilomyces tenuipes (Inglis et al.
2006; Luangsa-Ard et al. 2011). Due to
the difficulties and complexity in differentiation of Paecilomyces spp. and P. lilacinum, MALDI–TOF MS (Matrix-assisted laser desorption
ionization–time-of-flight mass spectrometry) technique was used to
identify these two species and this method was found to be a rapid, reliable,
and logistically simplified alternative to sequence-based analysis for the
routine identification of Paecilomyces spp. and P. lilacinum (Barker et al. 2014).

Whole genome
sequencing of the P. lilacinum
provided the genetic makeup of this fungus and determined the right
phylogenetic placement (Fig. 1; Table 1.). Whole genome nucleotide sequences of
nine species belonging to Hypocreales order were used to create the
phylogenetic tree for P. lilacinum that provided the confirmation of the
P. lilacinum as a different species
from the Paecilomyces lilacinus under
the order Eurotiales (Prasad et al. 2015). Another new species named Pupureocilium
lavendulum closely related to P. lilacinum was proposed by Perdomo
et al. in 2012 by analyzing the ITS sequence of 37 clinical isolates from different origins and sources including other
references strains of P. lilacinum, Paecilomyces nostocoides and Nomuraea
atypicola, the new species was characterized by yellow diffusible pigment
with subglobose or limoniform conidia (Perdomo et al. 2012).




Table 1. Genomic features of P. lilacinum
(Adapted from Prasad et al. 2015)


P. lilacinum

Genome Size (Mb)


% GC content


Predicted Proteins


Avg. gene density (genes/mb)


Avg. gene length (bp)


Repeat content




Secreted Proteins


Secondary metabolites clusters

30 (SMURF), 46 (AntiSMASH)

PHI genes





Morphology of Purpureocillium

P. lilacinum produces moderately to fast growing mycelium colonies on MEA
media (Fig. 2). It produces the conidiophores with floccose overgrowth of
aerial mycelium in it. P. lilacinum produces purple colored conidia,
which referred to its generic name Pupureocillium. The mycelium colonies
first appears as white colony and later on develop into pink and lilac then the
sporulation starts. The reverse color usually looks as purple or yellow. The fungus produces smooth walled hyaline
vegetative hyphae of about 2.5 -4.0 µm wide. The conidiophores arise from submerged hyphae as mononematous stiff
with verticillate i.e. phialides ovate to cylindrical having distinct neck,
densely group together forming verticils of branches and cylindrical phialides
with or without very short neck. Conidia are produced as dry divergent chains
with straight to slightly curve or ellipsoidal to fusiform shaped with smooth
to slightly roughened. This fungus does not produce chlamydosporesn and spores germinate in the suitable environmental
conditions such as temperature of 25 °C (77 °F) with enough moisture

and nutrient available.

Purpureocillium lilacinum as a human pathogen

P. lilacinum is the one of
the important cause of human diseases. This pathogen is associated with several
diseases such as Chagas disease, immune system compromised for intraocular lens
implants, contaminant of antiseptic cream, tattoo- related skin infection and
cavitary pulmonary disease (Jenifer et al.
2011; Khan et al. 2012; Saghrouni et al. 2013; Trinh
and Angarone 2016). The World Health Organization (WHO) has declared the Chagas
disease as one of the greatest scourges in South America and about 16-18
millions of people are infected with this disease and more 100 millions of
people from 21 Latin American countries are at risk. For example in Argentina,
Chagas disease is considered the most dangerous endemic arhropozoonosis with 2 millions
of people infected with Trypanosoma cruzi, the
causative agent of Chagas disease. The presence of P. lilacinus was confirmed in Triatoma infestans Klug, vector of Chagas disease under phase contrast light
microscope from the all dead insect and fungus was re-isolated from insect
cadavers (Marti et al. 2006; Gerardo et al. 2006). As Paecilomyces lilacinus was described
more than a century ago as a fungus commonly found in soil, but in last decades
several reports have been published and confirming the fungus can be a causative
agent of many infectious diseases in man and other vertebrates. Several cases
are described from the patients with compromised immune system or intraocular
lens implants (Jenifer et al. 2011). Other research results also suggested that
the P. lilacinum can contaminate
cosmetics items such as antiseptic creams and (skin) lotions, and other
neutralizing agent such as sodium bicarbonate solutions for artificial lenses,
and can contaminate and colonized the medical materials such as catheters and
plastic implants (Pettit et al., 1980; Orth et al., 1996; Itin et al., 1998). A
tattoo-related skin infection was found to have association with the P.
lilacinum in kidney transplant recipient and other similar reports has been
described as cutaneous infection in
immunocompetent patient (Saghrouni et al.

lilacinum as a biocontrol agent for plant-parasitic

fungal order Hypocreales consists many fungal species, which are plant
pathogens, insect-pathogens, nematode-pathogens, plant endophytes and
mycoparasites. The fungus Purpureocillium
lilacinum is widely known as a biological control agent against different plant
parasitic nematodes, insects and pests (Fiedler et al. 2007; Parajuli et al. 2014). Purpureocillium lilacinum, previously named as Paecilomyces
lilacinus. These fungal species are known to produce a diversity of
secondary metabolites and bio-actives, which acts as biological control agent
for nematode and insect pests. P.
lilacinum produces a mycotoxin named as paecilotoxin, which is considered an
effective biological control agent against plant parasitic nematodes,
especially economically important nematode species, Meloidogyne incognita (Bonants et al. 1995; Khan et al. 2004). Different research studies
suggested that nematode suppressive soil has a substantial presence of
nematophagous fungi such as P. lilacinum and
Pochonia chlamydosporia and others. Similar to other species
in Hypocreales, P. lilacinum has a
broad host range and its various strains parasitize different species of
nematodes and insects. P. lilacinum
is also reported to exercise parasitic or endophytic lifestyles in the presence
of a host organism such as nematodes, aphids and cotton plants.

251 of P. lilacinum, is one of widely used strains in plant parasitic
nematode control that infects eggs and females of Meloidogyne spp. and causes death of the nematode embryos in 5 to 7
days. Improving suppression of Meloidogyne
spp. by Purpureocillium lilacinum
strain 251 tested in cotton, peanut, and maize. Strain 251 found to be more
effective in peanut and cotton than in maize planting cover crops could also enhance
the effectiveness of P. lilacinum. (Parajuli et al. 2014). Besides the
nematode control, P. lilacinum can as
a biological control agent of leafcutter ants in crops and plantations (Goffre
et al. 2015). HYBDPL-04 strain of P.
lilacinum found to be highly effective for management of root-knot
disease of tomato under naturally infested field conditions. This isolates
found to produce the maximum number of metabolites, which analyzed through high-pressure
liquid chromatography (Singh et al. 2013). Cotton seeds infected with
two entomopathogens B. bassiana and P.
lilacinum showed the adverse effect on cotton aphid reproduction both in
the greenhouse and field experiment (Diana et al. 2014; Lopez et al. 2014). P. lilacium also improves
mangrove ecosystem by compensating the cupper stress by increasing the
concentration of cupper in the soil and improve the carbonate bound cupper in
the soil (Gong et al. 2017).


Enzymes produced by Pupureocillium

P. lilacinum produces several enzymes, which act as a biological control
agent for nematodes, serine protease is a basic enzyme produced by this fungus
that is utilized against the biological activity of plant parasitic nematode
such as Meloidogyne hapla (Bonants et
al. 1995). Proteases and chitinases are the other enzymes, which weaken the
nematode eggs shell and prevent from penetration to plant cell and reduces the
hatching of Meloidogyne javanica
juveniles (Khan et al. 2004).



Pupureocillium is one of the commonly used nematophagous fungus under the family ophiocordycipitaceace. Previously, Purpureocillium was described as Paecilimyces lilaciuns,
but later several moophological, molecular, and phyologenetic analysis confirm
the fungus as a different group species as Pupureocillium, the name
referred as purple colored conidia. P. lilacium is a representative species
under the genus Purpureocillium, but additional species has been described for
example Pupureocilium lavendulum. This fungus is widely
distributed and detected from several material such as soils, sludge,
cultivated and unclultivated land, females of nematodes and adaptable to varied
environmental conditions. This fungus is highly adaptable to the situation and
can acts as entomopathogenic,
mycoparasitic, saprophytic as well as nematophagous upon availability.

has been used for managing plant parasitic nematode mainly cyst and root knot
nematode. Besides nematode this fungus also been used to control other insect and
pests. Besides its advantage as a control agent for nematodes and other pest,
it also considered as a human pathogen. Several clinical cases has been
detected, where this pathogen found to be associated with the human diseases. Chagas
disease, immunocompromised patent, tattoo-related skin infection, keratitis
and, as a contaminant of cosmetics and medical utilities such as plastic
implants and catheters are some of the examples where the Purpureocillium
lilacium found to associated or directly involved to cause the disease.

P. lilacinum has both the positive and negative aspects. Several research studies
suggest that this fungus can acts as effective biological control measures nematode,
insect and pest crop plants, but this pathogen also becoming dangerous human
pathogens and found to be associated with several human diseases. It is very
important to investigate more about P.
lilacinum as utilizing as a biological control agent for nematodes and
insects and its possible impacts on human beings and the environment. Further
research are required for further application of this fungus as a biological
control agent for nematodes and insects.



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