Paecilomyces lilacinus: A Nematophagous and Entomopathogenic Fungus for Sustainable Biocontrol

Overview of the Microbe#

Paecilomyces lilacinus is a filamentous ascomycete fungus widely found in soils and decaying organic matter around the world[1][4]. It produces fast-growing colonies that are typically violet-lilac in color, with erect conidiophores bearing chains of smooth to slightly rough-walled spores (conidia) – a trait that inspired its name[1]. Formerly classified in the genus Paecilomyces, it was reclassified as Purpureocillium lilacinum after molecular studies showed it is distinct from other Paecilomyces species[1][4]. This fungus is a nematophagous organism (able to parasitize and kill nematodes) as well as an entomopathogenic fungus (infecting insects and mites), which has made it an important agent in sustainable biological pest control.P. lilacinus typically lives as a saprophyte feeding on dead matter, but it can aggressively attack the eggs and other life stages of plant-parasitic nematodes and certain pest insectscelkau.in[3]. It has a cosmopolitan distribution and can be isolated not only from soil but also from decaying vegetation, compost, nematode cysts, insect cadavers, and other organic substrates[1]. Under laboratory conditions it grows optimally in the mid-20°C range and does not grow or survive at human body temperature (≈37 °C)[3][4]. This temperature limitation, along with the fact that some strains (e.g. strain 251) produce no known harmful toxins, means P. lilacinus is generally considered safe for use as a biocontrol agent[3][4]. Although it is typically a risk-group 1 organism, rare cases of opportunistic human infections (e.g. in immunocompromised patients) have been reported[1]. Overall, however, Paecilomyces lilacinus is valued in agriculture as an eco-friendly microorganism that can naturally suppress pests while also potentially benefitting plant health.

Nematode Control and Root-Knot Management#

One of the primary uses of P. lilacinus is in controlling plant-parasitic nematodes, especially root-knot nematodes (Meloidogyne spp.) and cyst nematodes (Heterodera and Globodera spp.). P. lilacinus was first reported parasitizing nematode eggs in 1979, demonstrating its potential as a biological nematicide[5]. Since then, numerous studies under greenhouse and field conditions have shown that P. lilacinus can significantly suppress nematode populations and reduce root damage[6]. The fungus primarily infects nematode eggs: its mycelia invade the protective egg masses (or cysts), penetrate eggshells, and consume the developing nematodes within[6]. P. lilacinus produces a suite of extracellular enzymes (including chitin-degrading enzymes and proteases) that help dissolve the tough eggshell and cuticle of nematodes, facilitating infection. It can even infect adult female nematodes (entering via the vulva or anus) and juveniles, thereby attacking multiple life stages of the pest[3]. In infected eggs, the fungus proliferates and forms new spores, often destroying most eggs in an egg mass within about 5 days. By killing eggs and young nematodes before they can damage roots, P. lilacinus effectively breaks the nematode life cycle.

Plants treated with P. lilacinus typically show reduced root galling and nematode counts, along with better growth and yield compared to untreated, nematode-infested plants[6]. For example, inoculating soil with P. lilacinus has been shown to cut root-knot gall numbers and egg-mass hatching rates dramatically, sometimes by 70–90% in research trials[6]. The fungus may attack nematode larvae in soil or even kill female nematodes before they finish laying eggs, thus curbing the pest’s reproductive potential[6]. Additionally, P. lilacinus can induce certain plant defense responses; characterization of infection-related enzymes in nematode-infected plants (e.g. elevated chitinases and other pathogenesis-related proteins) suggests the fungus might elicit systemic resistance that further helps control nematode damage[6]. Farmers and researchers have integrated P. lilacinus into root-knot nematode management on various crops (tomato, pepper, banana, etc.), often as a soil treatment or seed/seedling coating. This fungus is especially attractive as an alternative to chemical nematicides, which are being restricted due to environmental and health concerns[6].

Despite its successes, performance of P. lilacinus in nematode control can sometimes be inconsistent, especially under open-field conditions. The efficacy may depend on environmental factors (soil type, temperature, moisture) and the virulence of the particular fungal strain[7][6]. In some trials, P. lilacinus provided excellent nematode suppression, while in others it failed to establish well or did not significantly reduce nematode numbers[6]. Research indicates that P. lilacinus thrives in the rhizosphere when nematode hosts are present – for instance, it proliferates around nematode-damaged roots (feeding on leaked root exudates and nematode egg masses)[6]. This host-dependent boost may increase its reliability as a biocontrol agent in infested fields[6]. To improve consistency, P. lilacinus is often used as part of an integrated approach, combined with other methods like organic soil amendments or symbiotic fungi. One study showed that pairing P. lilacinus with an arbuscular mycorrhizal fungus and adding green manure yielded the best control of Meloidogyne incognita on vegetable crops[6]. Such synergistic strategies, along with careful strain selection, aim to ensure that P. lilacinus can reliably protect crops from root-knot nematodes in sustainable farming systems.

Insect and Mite Biocontrol in Greenhouses#

In addition to nematodes, Paecilomyces lilacinus can infect and kill a range of insect and mite pests, making it a broad-spectrum biocontrol agent. Notably, it has demonstrated efficacy against several major greenhouse pests. For example, Fiedler & Sosnowska (2007) tested P. lilacinus on greenhouse whiteflies, spider mites, aphids, and thrips. Their laboratory and pot trials showed the fungus was able to infect all these pests, with particularly striking results on aphids: P. lilacinus applications almost completely eliminated cotton aphid (Aphis gossypii) populations[10]. It was also effective against greenhouse whitefly (Trialeurodes vaporariorum), especially when targeting the vulnerable nymphal stages (late instar nymphs)[10]. For western flower thrips (Frankliniella occidentalis), the best control was achieved by applying P. lilacinus spore suspensions to soil, which infected the soil-dwelling prepupal and pupal stages of the thrips[10]. Even the two-spotted spider mite (Tetranychus urticae) was susceptible – the fungus could suppress mite numbers by about 60%, particularly under cooler temperature conditions (below 10 °C) that seemed to favor fungal activity[10]. These findings highlight that P. lilacinus, traditionally known as a “nematode fungus,” is also capable of controlling small arthropod pests in greenhouse crops[10].Beyond those examples, P. lilacinus has been explored as a biocontrol for certain lepidopteran pests. A Vietnamese study isolated a strain of P. lilacinus (designated PL01) that proved highly virulent against the diamondback moth (Plutella xylostella), a notorious caterpillar pest of cruciferous vegetables, as well as the Oriental leafworm moth (Spodoptera litura). In laboratory bioassays, the P. lilacinus PL01 strain caused lethal infections in P. xylostella larvae with a median lethal time (LT₅₀) of only ~2.5 days. It was similarly pathogenic to S. litura (armyworm) larvae, though taking slightly longer (LT₅₀ ~7 days) to kill them. Such rapid kill rates are comparable to those of well-known entomopathogenic fungi like Beauveria bassiana. Field and greenhouse studies suggest that P. lilacinus can contribute to managing pests like moth larvae, whiteflies, thrips, and mites when applied as a foliar spray or soil drench in cropping systems[10]. It may not yet be as widely used for insects as some specialized insect pathogens, but research is ongoing. The versatility of P. lilacinus as both a nematode egg parasite and an insect pathogen underscores its value in integrated pest management, especially in greenhouses where a single biocontrol agent that tackles multiple pests is highly advantageous. Recent trials are even targeting emergent pests (see Spotlight section), indicating expanding roles for this fungus in biocontrol.

Plant Growth Promotion and Nutrient Solubilisation#

Interestingly, Paecilomyces lilacinus has shown potential beyond direct pest suppression – it can also act as a plant growth promoter and nutrient mobilizer. This fungus is sometimes found living endophytically (within plant roots) or in close association with roots without causing harm. In such cases, it may confer benefits to the host plant. Research has revealed several mechanisms by which P. lilacinus can enhance plant growth and nutrition. For instance, a recent greenhouse study tested an endophytic strain P. lilacinus 112 on tomato seedlings and observed remarkable growth stimulation[9]. Tomato plants treated with a protein hydrolysate derived from P. lilacinus culture grew significantly larger than controls – plant biomass increased 3.5-fold, plant height 3.3-fold, and there were notable gains in stem thickness and leaf number[9]. These improvements are partly attributed to bioactive compounds produced by the fungus. P. lilacinus 112 was shown to secrete various hydrolytic enzymes and siderophores, and to solubilize important nutrients like phosphorus and zinc in the soil[9]. By breaking down complex soil nutrients into forms plants can absorb and by chelating iron with siderophores, the fungus effectively boosts the nutrient availability for its host plant.P. lilacinus may also indirectly promote plant health by suppressing other soil pathogens. In the above study, the P. lilacinus 112 isolate exhibited antifungal activity against several plant-pathogenic fungi (e.g. inhibiting Cladosporium, Rhizoctonia, and Sclerotinia species by ~50–66% in vitro)[9]. This antagonism, combined with the nematode control, means plants experience less pest pressure and disease, which translates to better growth. There is also evidence that P. lilacinus can produce phytohormone-like substances or trigger plant hormonal pathways. While research is ongoing, some isolates of P. lilacinus have been reported to produce indole-3-acetic acid (an auxin) or gibberellin-like compounds, which could directly stimulate root and shoot development (these findings mirror observations in other entomopathogenic fungi that have endophytic phases). Additionally, when P. lilacinus helps break down organic matter (such as keratin in feather waste, as tested in one biostimulant study), it recycles nutrients back into forms plants can use[9]. All these traits categorize P. lilacinus as a plant growth-promoting fungus (PGPF) in addition to being a biocontrol agent. This dual benefit is highly appealing in sustainable agriculture: farmers might use P. lilacinus formulations not only to attack pests but also to improve crop vigor and yield through natural microbial fertilization.

Spore Production, Persistence and Formulation#

To deploy Paecilomyces lilacinus in the field, effective mass-production and formulation of its spores are crucial. P. lilacinus can be grown on a variety of inexpensive substrates using fermentation techniques. In solid-state fermentation, agricultural by-products like grains and husks serve as growth media for producing large quantities of conidia. For example, researchers found that simply using steamed brown rice (alone or mixed with rice husks and bran) provided an excellent medium for sporulation – yielding high spore counts suitable for industrial-scale production. Liquid fermentation can also be used to cultivate P. lilacinus; different broth recipes (potato dextrose, molasses-based media, etc.) have been tested to maximize fungal biomass and spore yield[7]. One study reported that a semi-selective rich medium produced the highest spore load (3.28 × 10^8 spores/ml), outperforming standard potato dextrose broth in biomass production[7]. Once grown, the fungal propagules are formulated into a stable product. Common formulations include talc-based wettable powders, granules, or water-dispersible granules (WG). The spores are mixed with a carrier (like talc, clay, or diatomaceous earth), plus additives to improve shelf-life (e.g. desiccants and stabilizers).

The persistence of P. lilacinus in formulated products and in soil is an important consideration. Studies on various carrier materials have shown that spore viability can be maintained for several months under proper storage. In one comparison, P. lilacinus spores formulated in talc powder or fly-ash remained viable for at least 120 days (4 months) at room temperature[7]. In contrast, carriers like rice husk ash and vermiculite supported spore survival for a shorter period (around 75–90 days)[7]. Thus, talc-based formulations are popular for commercial products due to their longer shelf-life. P. lilacinus strain 251 – a well-known biocontrol strain – has been developed into a water-dispersible granule (WG) product (e.g. BioAct®) using solid-state fermentation and drying technologycelkau.incelkau.in. These granules contain dried spores plus formulation aids (for instance, one product contains ~6% P. lilacinus spores with skim milk powder and dispersants) and can be mixed into water and applied via drench or spray. In soil, the persistence of P. lilacinus depends on environmental conditions: it survives and remains active longer in moderate temperatures and adequate moisture, whereas very hot (>35 °C) or dry conditions can reduce its spore viability and growthcelkau.in. Fortunately, P. lilacinus can tolerate a range of soil types and can even form resilient resting structures under stress. Field studies indicate that after application, P. lilacinus can establish in the rhizosphere and sometimes persists through a growing season, especially if there are sufficient host nematodes or insect targets to sustain it[6]. Farmers often apply it at planting and then periodically (every 6–8 weeks) to ensure a continuous presence of fresh spores in the soil[3]. Overall, advances in mass-production and formulation (like improved media, and WG or oil-based formulations) have greatly enhanced the practicality of P. lilacinus as a microbial pesticide.

Challenges and Future Potential#

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While Paecilomyces lilacinus is a promising biocontrol agent, there are several challenges in its widespread use. One issue is variability in performance – field results have sometimes been inconsistent, with efficacy ranging from excellent control to negligible effect[6]. This inconsistency can stem from environmental factors (since P. lilacinus is sensitive to temperature and moisture extremes) and biological factors like strain differences or competition with native soil microbes. Selecting highly virulent, well-adapted strains is essential; for instance, strain 251 was chosen for commercialization due to its strong activity and ability to persist in various soils. Even with good strains, P. lilacinus typically works more slowly than chemical nematicides or insecticides – it may take days or weeks to reduce pest populations, whereas chemicals can act in hours. Consequently, growers must time applications in advance of pest outbreaks and manage expectations that biocontrol is often gradual and partial (seldom achieving 100% pest elimination). Another consideration is safety and regulation. Although P. lilacinus is non-pathogenic to plants and generally safe for humans, some regulatory agencies carefully evaluate any microorganism introduced to the environment. Strain 251’s registration, for example, required demonstrating it does not produce dangerous toxins or survive at body temperature[1]. The existence of rare human infections by other P. lilacinus strains means producers must ensure manufacturing is free of contaminants and advise users (especially immunocompromised individuals) to handle high concentrations of spores with care (e.g. wearing masks to avoid inhalation).

Looking ahead, there is substantial potential to improve and expand the use of P. lilacinus. Strain improvement and discovery: Ongoing research is exploring new isolates from different regions – some may have enhanced heat tolerance, faster kill rates, or broader host ranges. For instance, scientists are examining P. lilacinus strains from extreme environments to find ones that remain active at higher temperatures, which would be valuable in tropical agriculture. Genomic insights: The genome of P. lilacinus has been sequenced, and comparative genomics/transcriptomics are shedding light on the genes responsible for its parasitic abilities[2]. This could guide genetic enhancements or more targeted application methods (e.g. formulating the specific enzymes or compounds it uses to attack pests). Formulation technology: Future formulations might include encapsulated spores or improved granular products that prolong shelf-life and protect the fungus from UV and desiccation when applied to crops. Integration into cropping systems: P. lilacinus is being integrated with other sustainable practices. For example, combining it with organic amendments (compost, manures) can improve its establishment and efficacy, as nutrient-rich amendments often boost microbial biocontrol activity[6]. Similarly, using P. lilacinus alongside symbiotic fungi (like mycorrhizae) or beneficial bacteria might create a synergistic microbe consortium that tackles multiple soil health aspects simultaneously.

Another exciting area is the exploration of P. lilacinus in managing emerging pests. As discussed, it already shows activity against certain insect pests; researchers are now testing it against invasive species such as the fall armyworm and tomato leafminer with promising results (see Spotlight below). Its ability to produce bioactive metabolites (e.g. leucinostatins, which have antifungal and anti-oomycete properties) could be harnessed for controlling plant diseases like wilt fungi or downy mildews[8]. In summary, Paecilomyces lilacinus faces the typical challenges of any biocontrol agent – needing the right conditions and correct usage for consistent results – but with continued research and development, it holds great future potential. Its multifaceted nature (attacking pests and bolstering plant growth) aligns well with the goals of sustainable agriculture, making it a focal point for innovations in biological pest management.

Spotlight on Research: Paecilomyces lilacinus#

Brief Overview#

A recent notable study (Riaz et al., 2024) focused on leveraging Purpureocillium lilacinum (formerly P. lilacinus) as a biocontrol for the fall armyworm, an invasive insect pest. This research, conducted in Southern Taiwan, involved isolating a native strain of P. lilacinum from soil and evaluating its pathogenicity against fall armyworm (FAW, Spodoptera frugiperda) – a destructive caterpillar that attacks maize and many other crops[11]. The scientists first confirmed the identity of their fungal isolate (named strain “PT-02”) using morphological characteristics and molecular markers (ITS and TEF gene sequencing), which established it as Purpureocillium lilacinum[11]. They then carried out bioassays to test how effectively this fungus could infect and kill FAW at different life stages. Eggs, newly hatched larvae (neonates), and older larvae (1st and 2nd instars) of the armyworm were treated with conidial suspensions of P. lilacinum PT-02 under controlled conditions[11]. The study’s aim was not only to measure mortality rates at various spore concentrations but also to determine lethal concentration values and assess whether this fungal strain could be a viable biopesticide for managing FAW outbreaks.

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Key Insights#

The results of this study were encouraging. P. lilacinum PT-02 showed high virulence against fall armyworm eggs and very young larvae[11]. At the highest spore concentration tested (1×10^8 spores/mL), the fungus killed essentially 98–100% of FAW eggs within a few days of application[11]. Even at a tenth of that concentration (1×10^7 spores/mL), egg mortality remained 98%, indicating the fungus is extremely effective at colonizing and destroying FAW eggs before they hatch[11]. Notably, P. lilacinum was also lethal to the neonate caterpillars (the larvae that had just emerged from eggs): exposure to the fungus caused 100% mortality of neonates in the assays[11]. For slightly older larvae, the efficacy dropped but was still significant – first-instar larvae suffered between 10% and 36% mortality (depending on spore dose), and second-instar larvae had similar mortality rates within 7 days[11]. The researchers calculated median lethal concentration (LC₅₀) values to quantify potency: the LC₅₀ for eggs was on the order of ~3×10^7 spores/mL, whereas for first-instar larvae it was ~1.3×10^8 and for second-instar around 2.5×10^8 spores/mL[11]. These metrics indicate that eggs and very small larvae are far more susceptible to P. lilacinum infection than larger larvae. Another key finding was that the fungal isolate’s growth and appearance can vary depending on culture conditions (they observed differences in colony morphology at different temperatures and media, underscoring the importance of optimal conditions for mass production)[11]. Overall, the study demonstrated that P. lilacinum PT-02 is a potent pathogen of FAW in its early life stages.

Why This Matters#

This research is significant for several reasons. First, fall armyworm is a highly invasive pest that has recently spread across Africa and Asia, causing billions of dollars in crop losses annually[11. Current FAW control strategies rely heavily on chemical insecticides and genetically modified crops, which have drawbacks like resistance development and environmental harm[11]. The successful use of P. lilacinum against FAW suggests a promising eco-friendly alternative: a biological agent that can target the pest’s eggs and hatchlings, potentially curbing infestations at the very start. By proving that a P. lilacinum strain can effectively kill FAW eggs/neonates, this study opens the door to developing biopesticide products based on this fungus for FAW management. It also expands the known host range of P. lilacinum, reinforcing that this fungus is not limited to soil pests but can actively infect above-ground insect pests of major crops. This cross-domain efficacy (nematodes and insects) is particularly valuable in tropical farming systems where FAW and nematodes might co-occur – a single biocontrol agent could tackle both. Moreover, the study emphasizes the importance of targeting vulnerable pest stages; using P. lilacinum to wipe out FAW eggs can prevent the explosive outbreak of larvae that devastate crops. From a scientific perspective, the work contributes knowledge on the pathogenic mechanisms of P. lilacinum in insects, complementing what is known about its nematode parasitism. It underlines that P. lilacinum strains can be locally sourced and that classical isolation techniques combined with molecular tools are effective in procuring biocontrol strains in new regions. In summary, this spotlight study matters because it demonstrates a sustainable solution to a pressing pest problem and broadens the applicability of P. lilacinum in integrated pest management programs worldwide.

Summary Table: Spotlight Study#

Category	Details
Lead Researchers	Muhammad Riaz; Wen-Hua Chen (corresponding); Lekhnath Kafle; Min-Nan Tseng et al.
Affiliations	National Pingtung Univ. of Science & Technology; Kaohsiung District Agricultural Research & Extension Station (Taiwan)
Research Focus	Biocontrol of invasive fall armyworm (Spodoptera frugiperda) using native P. lilacinum
Key Breakthroughs	Isolated a local P. lilacinum strain (PT-02) and confirmed its identity via ITS/TEF sequencing. Demonstrated ~98–100% mortality of FAW eggs and neonate larvae under lab conditions. Calculated LC50 values, showing high potency against early pest stages.
Collaborative Efforts	Collaboration between a university research team and a regional agricultural research station in Taiwan. The study combined academic lab assays with practical insights for field application
Published Work	Egyptian Journal of Biological Pest Control (Open-access, 2024, vol. 34 art. 60) – Peer-reviewed research article.
Perspective	Offers a sustainable, non-chemical approach to FAW management. Highlights the entomopathogenic capacity of P. lilacinum beyond nematodes, with implications for global pest control efforts.
Publication Date	11 November 2024
Location	Pingtung, Taiwan (laboratory and growth-chamber experiments)
Key Findings	P. lilacinum strain PT-02 caused nearly 100% egg and neonate mortality in fall armyworm, and significant mortality (10–36%) in early instar larvae, within 7 days. The fungus shows promise as a biocontrol agent to protect crops from this invasive pest by targeting its most vulnerable life stages.

Conclusion#

Paecilomyces lilacinus (syn. Purpureocillium lilacinum) stands out as a multifaceted fungus that contributes to sustainable agriculture in several ways. As a biocontrol agent, it has proven effective against some of the most damaging soil pests – it parasitizes root-knot and cyst nematodes, reducing their populations and thus preventing the devastating galling and yield losses these nematodes can cause. At the same time, P. lilacinus has shown it can infect and suppress certain insect and mite pests, particularly in protected cultivation systems like greenhouses. This dual nematode-insect activity, combined with its ability to promote plant growth (through nutrient solubilization and possible endophytic benefits), makes P. lilacinus a valuable tool in integrated pest and crop management. The fungus can be mass-produced relatively easily and formulated into stable products that farmers can apply to soils or crops, and ongoing improvements in formulation are addressing past issues of shelf-life and field consistency.Like many biocontrol organisms, P. lilacinus does face challenges – its performance can be influenced by environmental conditions, and it generally acts more slowly than chemical pesticides. Yet, research and field experience have helped identify robust strains (such as strain 251 and others) and best-use practices (e.g. combining with organic amendments or other biocontrol agents) to enhance its efficacy. The future looks promising: genomic studies are unlocking the molecular secrets of its parasitism, and new trials (as highlighted in the spotlight) are expanding its use against major pests like the fall armyworm. There is a concerted push towards reducing chemical inputs in agriculture, and microbes like Paecilomyces lilacinus are at the forefront of this transition. In summary, P. lilacinus is an exemplary nematophagous and entomopathogenic fungus whose roles in biocontrol and plant health continue to grow. Its story – from a soil mold described over a century ago to a modern bio-nematicide and bio-insecticide – reflects the broader trend of harnessing natural enemies for sustainable crop protection. With continued research and field validation, P. lilacinus is likely to remain a “little purple” powerhouse in the biological arsenal against pests, contributing to safer and more resilient agricultural systems.

References#

1. Luangsa-ard, J.J. et al. (2011). Purpureocillium, a new genus for the medically important Paecilomyces lilacinus. FEMS Microbiology Letters, 321(2): 141–149. (Taxonomic reclassification of P. lilacinus to Purpureocillium lilacinum)adelaide.edu.aubayer.com.
   
2. Xie, J., Li, S., Mo, C., Xiao, X., Peng, D., Wang, G., & Xiao, Y. (2016). Genome and transcriptome sequences reveal the specific parasitism of the nematophagous Purpureocillium lilacinum 36-1. Frontiers in Microbiology, 7, 1084. https://doi.org/10.3389/fmicb.2016.01084.
   
3. EPA (2005). Biopesticides Registration Action Document – Paecilomyces lilacinus strain 251. U.S. Environmental Protection Agency, Office of Pesticide Programs, Biopesticides Fact Sheet 028826. (Safety profile of strain 251: origin, temperature growth limits, non-toxicity)www3.epa.govwww3.epa.gov.
   
4. Bayer CropScience (2006). Purpureocillium lilacinum 251 – Microbial Pest Control Agent Dossier (Excerpt). (Strain 251 safety summary: ubiquity in soil, lack of pathogenicity or toxin production, inability to grow at >36 °C)bayer.combayer.com.
   
5. Gaspard, J.T. et al. (1990). Paecilomyces lilacinus with Root-knot Nematode Infested Soil. Nematropica 20(1): 33–41. (Early report on using P. lilacinus against Meloidogyne; citing Jatala’s first findings in 1979)patents.google.com.
   
6. Khan, A. et al. (2022). Effect of Rhizoglomus fasciculatum and Paecilomyces lilacinus in the biocontrol of root-knot nematode, Meloidogyne incognita in Capsicum annuum. Scientific Reports 12: 3432. (Demonstrated P. lilacinus reduces galling and egg masses; synergy with mycorrhiza)pmc.ncbi.nlm.nih.govpmc.ncbi.nlm.nih.gov.
   
7. Prabhu, S. et al. (2008). Mass production and commercial formulation of Paecilomyces lilacinus. Madras Agricultural Journal 95(7-12): 415–417. (Liquid media comparison for biomass yield; spore viability in talc vs other carriers over 120 days)masujournal.orgmasujournal.org.
   
8. Wang, G., Liu, Z., Lin, R., Li, E., Mao, Z., Ling, J., Yang, Y., Yin, W.-B., & Xie, B. (2016). Biosynthesis of antibiotic leucinostatins in bio-control fungus Purpureocillium lilacinum and their inhibition on Phytophthora revealed by genome mining. PLOS Pathogens, 12(7), e1005685. https://doi.org/10.1371/journal.ppat.1005685.
   
9. Constantin, M. et al. (2022). Exploring the Potential Applications of Paecilomyces lilacinus 112. Applied Sciences 12(15): 7572. (Endophytic P. lilacinus promoting tomato growth; solubilization of Zn and P; biostimulant production from keratin waste)mdpi.commdpi.com.
   
10. Fiedler, Ż. & Sosnowska, D. (2007). Paecilomyces lilacinus as a biological agent for control of greenhouse insects and mite pests. BioControl 52: 547–558. (Lab and greenhouse tests showing P. lilacinus efficacy against whiteflies, thrips, aphids, spider mites)link.springer.comlink.springer.com.
    
11. Riaz, M. et al. (2024). Morphological and molecular characterization of Purpureocillium lilacinum and its biopesticidal effect against fall armyworm (Spodoptera frugiperda). Egyptian Journal of Biological Pest Control 34: 60. (Recent study demonstrating a native P. lilacinum strain’s high efficacy in killing FAW eggs and neonate larvae)link.springer.comlink.springer.com.