Characterization of Aspergillus species associated with dried cocoa beans in Peninsular Malaysia

Characterization of Aspergillus species associated with dried cocoa beans in Peninsular Malaysia | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Characterization of Aspergillus species associated with dried cocoa beans in Peninsular Malaysia Sahariman Nuraina-Syazana, Muhamad Zaid Zakaria, Nor Shifa Shuib, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6760467/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Cocoa ( Theobroma cacao ) is a key agricultural commodity that is highly susceptible to fungal and mycotoxin contamination, particularly during post-harvest processes such as fermentation, drying, and storage. This study aimed to characterize Aspergillus species associated with dried cocoa beans from 12 farms across six states in Peninsular Malaysia using morphological, molecular, phylogenetic, and toxigenicity analyses. Fungal isolation and enumeration were performed, followed by identification based on morphological characteristics, DNA sequencing, and phylogenetic analysis targeting the internal transcribed spacer (ITS) region and β-tubulin ( tub2 ) gene. Mycotoxin-producing potential was assessed using coconut cream agar (CCA), where fluorescence under ultraviolet (UV) light indicated the presence of secondary metabolites. A total of 43 Aspergillus isolates were identified as A. flavus (10 isolates), A. niger (nine isolates), A. tamarii (nine isolates), A. tubingensis (eight isolates), and A. aculeatus (seven isolates). One sample exceeded the hygienic threshold, with a fungal load of 2.9 × 10⁵ CFU/g, surpassing the permissible limit of 1 × 10⁴ CFU/g. All isolates exhibited fluorescence, indicating potential mycotoxin production. Moderate fluorescence intensity was observed exclusively in A. niger , while all other species showed low fluorescence levels. These findings underscore the importance of improved post-harvest handling practices to mitigate fungal contamination and mycotoxin risks. The results offer valuable insights for enhancing food safety, supporting cocoa trade, and informing agricultural policy in cocoa-producing regions. Theobroma cacao toxigenic fungi mycotoxins post-harvest handling phylogenetic analysis food safety Figures Figure 1 Figure 2 Figure 3 Introduction Cocoa ( Theobroma cocoa ) is a globally significant crop and the primary raw material in chocolate production. While West Africa remains the leading cocoa-producing region, substantial cultivation also occurs in Asia, Central America, and South America (Copetti et al. 2014 ). The T. cacao tree thrives in tropical climates characterized by high humidity, consistent rainfall (1,500–3,000 mm annually), and temperatures ranging from 20°C to 28°C. The fruit, known as a cocoa pod, contains 25–75 seeds embedded in a sweet, mucilaginous pulp. Upon drying, these seeds transition from white to violet or reddish-brown (Kim et al. 2011 ). Cocoa beans undergo several post-harvest processing steps before consumption, beginning with manual pod harvesting and opening. The extracted beans are naturally fermented in containers for several days, followed by sun drying on wooden platforms, cement surfaces, or directly on the ground. Once dried, the beans are stored, bagged, and transported to processing facilities where they are roasted, shelled, and ground into cocoa products. Each of these stages presents opportunities for microbial contamination, particularly by fungi introduced via pod surfaces, handling tools, workers’ hands, insects, and residual mucilage (Copetti et al. 2014 ). Fungal contamination is especially prevalent during the drying phase, where environmental exposure facilitates the growth of mycotoxigenic species. Several studies have reported the presence of Aspergillus species in cocoa beans, including those capable of producing aflatoxins (AFs) and ochratoxin A (OTA), mycotoxins of significant concern due to their toxicity and prevalence in food products (Mounjouenpou et al. 2008 ; Copetti et al. 2010 , 2011 ; Kedjebo et al. 2015 ). Mycotoxin contamination has been documented in cocoa-producing countries such as Côte d’Ivoire (Guehi et al. 2017 ), Cameroon (Mounjouenpou et al. 2008 ), and Brazil (de Magalhães et al. 2011 ). Aflatoxins and OTA are particularly hazardous due to their carcinogenic, nephrotoxic, and immunosuppressive effects (Copetti et al. 2012 ). Understanding the diversity and toxigenic potential of fungal species associated with cocoa beans is essential for assessing food safety risks and improving post-harvest practices. While most Malaysian studies have focused on the biochemical and physiological properties of cocoa (Farah et al. 2012 ; Agus et al. 2018 ; Ooi et al. 2020 ), limited research has addressed fungal contamination and mycotoxin risks. Therefore, this study aims to identify Aspergillus species associated with dried cocoa beans in Peninsular Malaysia using morphological, molecular, and phylogenetic approaches, and to evaluate their potential to produce mycotoxins. Materials and methods Sample collection and fungal isolation A total of 12 dried cocoa bean samples were collected from 12 farms across six states in Malaysia namely Penang, Perak, Selangor, Negeri Sembilan, Johor, and Melaka (Fig. 1 ). To eliminate surface contaminants, the beans were surface-sterilized by sequential immersion in 70% ethanol (C₂H₅OH) and 1% sodium hypochlorite (NaOCl) for 3 min each. The samples were then rinsed three times with sterile distilled water for 1 min per rinse and air-dried on sterile filter paper under aseptic conditions. The sterilized beans were plated onto potato dextrose agar (PDA) and incubated at 25 ± 2°C for 3–5 days, or until visible mycelial growth emerged. Actively growing mycelia were subcultured onto fresh PDA plates and incubated under the same conditions for 7 days to promote sporulation. To obtain pure cultures, single conidial isolation was performed by streaking a conidial suspension onto PDA using a sterile inoculating loop. Hyphal tips from 48-hour-old cultures were then excised and transferred to new PDA plates to establish monoconidial isolates for further analysis. Fungal enumeration A total of 5 g of dried cocoa beans were weighed and ground into a fine powder using a sterile mortar and pestle. One gram of the homogenized sample was transferred into a universal bottle containing 9 mL of sterile distilled water to prepare a 10⁻¹ dilution. The mixture was thoroughly shaken to ensure uniform suspension. Subsequent serial dilutions were prepared up to 10⁻⁵. From each dilution, 0.1 mL was aseptically spread onto PDA plates in triplicate. The inoculated plates were incubated at 25 ± 2°C for 5 days. After incubation, fungal colonies were counted, and the colony-forming units per gram (CFU/g) of sample were calculated following the method described by Ntuli et al. ( 2017 ). Morphological characterization Pure fungal isolates obtained were cultured on Czapek yeast extract agar (CYA) and malt extract agar (MEA) to assess their morphological characteristics. To prevent conidial dispersion, a drop of sterilized paraffin oil was placed at the center of each plate. The cultures were incubated at 25 ± 2°C for 7 days. Macroscopic features, including mycelial growth rate, were measured after 3 days of incubation. Colony morphology such as texture, color, and pigmentation on the reverse side was evaluated after 7 days on both CYA and MEA. For microscopic examination, fungal structures including conidia, phialides, and conidiophores were observed using the slide mount method, with preparations made on glass slides using sterile distilled water as the mounting medium. Observations were conducted using a compound light microscope (CX41, Olympus, Japan) equipped with a digital camera (KY-F55BE, JVC, Japan) to document morphological features. Molecular identification and phylogenetic analysis Molecular identification was carried out to confirm the identity of the fungal isolates obtained. Each isolate was cultured in potato dextrose broth (PDB) and incubated at 25 ± 2°C for 2 days. The resulting mycelia were harvested, air-dried on sterile filter paper, and ground into a fine powder using liquid nitrogen. Approximately 50 mg of the powdered mycelia was transferred into a sterile 1.5 mL microcentrifuge tube for genomic DNA extraction. DNA was extracted using the InnuPREP Plant DNA Kit (Analytik Jena, Germany), following the manufacturer’s protocol. Two genetic markers were targeted for amplification namely the internal transcribed spacer (ITS) region and the β-tubulin ( tub2 ) gene. The ITS region was amplified using the primer pair ITS1 (5′-TCCGTAGGTGAACCTGCGG-3′) and ITS4 (5′-TCCTCCGCTTATTGATATGC-3′) (White et al. 1990 ), while the tub2 gene was amplified using primers Bt2a (5′-GGTAACCAAATCGGTGCTGCTTTC-3′) and Bt2b (5′-ACCTCAGTGTAGTGACCCTTGGC-3′) (Glass and Donaldson 1995 ). PCR reactions were prepared in 50 µL volumes containing 8 µL of 5× Green GoTaq buffer, 8 µL of 25 mM MgCl₂, 1 µL of 10 mM dNTP mix, 8 µL of each primer (5 µM for ITS and 1 µM for tub2 ), 0.3 µL of Taq polymerase (5 U/µL), 1 µL of genomic DNA, and 15.7 µL of sterile distilled water. Amplification was performed using a MyCycler Thermal Cycler (Bio-Rad, USA). For the ITS region, the thermal cycling conditions included an initial denaturation at 94°C for 1 min 35 s, followed by 35 cycles of denaturation at 95°C for 35 s, annealing at 59°C for 55 s, and extension at 72°C for 1 min 30 s, with a final extension at 72°C for 10 min. For the tub2 gene, the protocol included an initial denaturation at 94°C for 5 min, followed by 35 cycles of denaturation at 94°C for 1 min, annealing at 57°C for 1 min, and extension at 72°C for 1 min, with a final extension at 72°C for 7 min. PCR products were separated by electrophoresis on 1.0% agarose gels stained with FloroSafe DNA stain (First BASE) in 1× TBE buffer. Electrophoresis was conducted at 80 V and 400 mA for 90 min. DNA bands were visualized using a Bio-Rad Gel Doc™ XR System and documented with Quantity One® software. Fragment sizes were estimated using a 100 bp GeneRuler™ DNA ladder (Thermo Scientific, USA). The amplified products were then sent to First BASE Laboratories Sdn Bhd (Seri Kembangan, Malaysia) for purification and sequencing. The resulting DNA sequences were aligned using MEGA12 software (Kumar et al. 2024 ). Sequence identities were confirmed through BLAST searches against the NCBI GenBank database. All sequences generated were submitted to GenBank. Phylogenetic relationships were inferred using the maximum likelihood (ML) method in MEGA12. Prior to tree construction, the best-fit nucleotide substitution model was determined. Individual gene trees were constructed using the Kimura 2-parameter model (Kimura 1980 ), while the combined ITS and tub2 dataset was analyzed using the Tamura 3-parameter model (Tamura 1992 ), with 1,000 bootstrap replications to assess branch support. Reference sequences and accession numbers used in the phylogenetic analysis are listed in Table 1 . Table 1 Fungal isolates and their corresponding GenBank accession numbers used in the phylogenetic analysis Aspergillus species Isolate Host Locality GenBank accession number Reference ITS tub2 A. aculeatinus CBS 121060 ET Coffea arabica Thailand EU159211 EU159220 Noonim et al. 2008 A. aculeatinus CBS 121871 Coffea robusta Thailand EU159206 EU159227 Noonim et al. 2008 A. aculeatinus CBS 121877 Coffee arabica Thailand EU159213 EU159224 Noonim et al. 2008 A. aculeatus CBS 172.66 ET Soil India FJ629320 FJ629271 Varga et al. 2011 A. aculeatus CBS 11480 Soil India AY585557 AY585539 Noonim et al. 2008 A. aculeatus CBS 62078 Unknown Not stated AY585556 AY585538 Noonim et al. 2008 A. aculeatus DT57 Theobroma cacao Malaysia: Melaka PQ835672 PV654129 Present study A. aculeatus PT51 Theobroma cacao Malaysia: Penang PQ835666 PV654123 Present study A. aculeatus PT52 Theobroma cacao Malaysia: Penang PQ835667 PV654124 Present study A. aculeatus TI53 Theobroma cacao Malaysia: Negeri Sembilan PQ835668 PV654125 Present study A. aculeatus TI54 Theobroma cacao Malaysia: Negeri Sembilan PQ835669 PV654126 Present study A. aculeatus TI55 Theobroma cacao Malaysia: Negeri Sembilan PQ835670 PV654127 Present study A. aculeatus RU56 Theobroma cacao Malaysia: Negeri Sembilan PQ835671 PV654128 Present study A. arachidicola CBS 117610 ET Arachis glabrata Argentina EF409241 EF203158 Pildain et al. 2008 A. awamori CBS 557.65 ET Unknown Not stated AM087614 AY820001 Samson et al. 2004 A. awamori ATHUM 5181 Unknown Greece EU982010 EU982069 Krimitzas et al. 2013 A. awamori NRRL 4951 Unknown Brazil KC796388 KC796358 Krimitzas et al. 2013 A. caelatus CBS 763.97 ET Soil USA MH862672 EF203127 Varga et al. 2011 A. fasciculatus CBS 110.55 ET Air contaminant Brazil FJ491463 EF203135 Varga et al. 2011 A. fijiensis CBS 313.89 ET Soil Fiji FJ491680 FJ491688 Varga et al. 2011 A. flavus CBS 100927 EN Cellophane South Pacific Islands MH864264 KJ175479 Frisvad et al. 2019 A. flavus CBS 100558 Peanut Indonesia MG517645 KJ175478 Frisvad et al. 2019 A. flavus CBS 816.96 Arachis hypogea Brazil MG517628 MG518011 Frisvad et al. 2019 A. flavus BD21 Theobroma cacao Malaysia: Perak PQ835646 PQ839564 Present study A. flavus BD22 Theobroma cacao Malaysia: Perak PQ835647 PQ839565 Present study A. flavus DT26 Theobroma cacao Malaysia: Melaka PQ835651 PQ839569 Present study A. flavus DT27 Theobroma cacao Malaysia: Melaka PQ835652 PQ839570 Present study A. flavus DT28 Theobroma cacao Malaysia: Melaka PQ835653 PQ839571 Present study A. flavus JN29 Theobroma cacao Malaysia: Melaka PQ835654 PQ839572 Present study A. flavus JN210 Theobroma cacao Malaysia: Melaka PQ835655 PQ839573 Present study A. flavus PJ23 Theobroma cacao Malaysia: Penang PQ835648 PQ839566 Present study A. flavus SB24 Theobroma cacao Malaysia: Penang PQ835649 PQ839567 Present study A. flavus SB25 Theobroma cacao Malaysia: Penang PQ835650 PQ839568 Present study A. foetidus CBS 564.65 Unknown Japan FJ629329 GU296697 Samson et al. 2004 A. foetidus NRRL 4750 Alcoholic beverage Japan FJ491683 FJ491690 Varga et al. 2011 A. kambarensis CBS 542.69 ET Unknown Japan FJ491468 EF203136 Samson et al. 2004 A. neoniger CBS 115656 ET Mangrove water Venezuela FJ491682 FJ491691 Varga et al. 2011 A. neoniger NRRL 62634 Mangrove water Venezuela KC796401 KC796361 Varga et al. 2011 A. niger CBS 42064 ET Unknown Not stated AY585553 AY820002 Samson et al. 2004 A. niger CBS 55765 Unknown USA EF661186 AY820001 Samson et al. 2004 A. niger CBS 101705 Carpet dust Canada FJ629343 AY820005 Samson et al. 2004 A. niger BP39 Theobroma cacao Malaysia: Johor PQ835665 PQ839599 Present study A. niger CR31 Theobroma cacao Malaysia: Perak PQ835657 PQ839591 Present study A. niger PJ32 Theobroma cacao Malaysia: Penang PQ835658 PQ839592 Present study A. niger PJ33 Theobroma cacao Malaysia: Penang PQ835659 PQ839593 Present study A. niger TK34 Theobroma cacao Malaysia: Selangor PQ835660 PQ839594 Present study A. niger TK35 Theobroma cacao Malaysia: Selangor PQ835661 PQ839595 Present study A. niger TI36 Theobroma cacao Malaysia: Negeri Sembilan PQ835662 PQ839596 Present study A. niger RU37 Theobroma cacao Malaysia: Negeri Sembilan PQ835663 PQ839597 Present study A. niger RU38 Theobroma cacao Malaysia: Negeri Sembilan PQ835664 PQ839598 Present study A. oryzae CBS 100925 ET Arachis hypogaea Argentina KJ175432 EF203138 Pildain et al. 2008 A. oryzae NRRL 447 ET Eurotium oryzae USA NR_135395 EF661483 Peterson 2008 A. oryzae ATHUM 4958 Indoor air Greece EU982022 EU982081 Krimitzas et al. 2013 A. parasiticus CBS 100926 EN Arachis hypogaea Argentina KJ175437 EF203155 Pildain et al. 2008 A. parasiticus ATHUM 5037 Indoor air Greece EU982020 EU982079 Krimitzas et al. 2013 A. parasiticus ATHUM 5038 Indoor air Greece EU982021 EU982080 Krimitzas et al. 2013 A. pseudotamarii CBS 766.97 ET Soil USA MH862673 EF203125 Varga et al. 2011 A. tamarii CBS 104.13 ET Activated carbon Unknown origin MH854614 EF203123 Frisvad et al. 2019 A. tamarii CBS 117626 Arachis hypogaea Argentina NR_135325 EF203122 Pildain et al. 2008 A. tamarii CBS 133375 Indoor air Germany MH279382 MH279885 Frisvad et al. 2019 A. tamarii BD11 Theobroma cacao Malaysia: Perak PQ835674 PQ839574 Present study A. tamarii DT16 Theobroma cacao Malaysia: Melaka PQ835679 PQ839579 Present study A. tamarii DT17 Theobroma cacao Malaysia: Melaka PQ835680 PQ839580 Present study A. tamarii PJ12 Theobroma cacao Malaysia: Penang PQ835675 PQ839575 Present study A. tamarii PJ13 Theobroma cacao Malaysia: Penang PQ835676 PQ839576 Present study A. tamarii PJ14 Theobroma cacao Malaysia: Penang PQ835677 PQ839577 Present study A. tamarii PJ15 Theobroma cacao Malaysia: Penang PQ835678 PQ839578 Present study A. tamarii TA18 Theobroma cacao Malaysia: Johor PQ835681 PQ839581 Present study A. tamarii TA19 Theobroma cacao Malaysia: Johor PQ835682 PQ839582 Present study A. terricola CBS 620.95 Soil USA FJ531244 FJ491476 Varga et al. 2011 A. thomii CBS 120.51 ET Culture contaminant Not stated MH856773 EF203133 Pildain et al. 2008 A. tubingensis CBS 11732 ET Kuro-koji Japan GQ379912 AY820012 Samson et al. 2004 A. tubingensis CBS 42565 Unknown Japan NR_131293 AY820009 Samson et al. 2004 A. tubingensis CBS 134.48 Unknown Not stated FJ629354 FJ629305 Noonim et al. 2008 A. tubingensis BP47 Theobroma cacao Malaysia: Johor PQ835690 PQ839589 Present study A. tubingensis BP48 Theobroma cacao Malaysia: Johor PQ835691 PQ839590 Present study A. tubingensis CR41 Theobroma cacao Malaysia: Perak PQ835684 PQ839583 Present study A. tubingensis JN44 Theobroma cacao Malaysia: Melaka PQ835687 PQ839586 Present study A. tubingensis JN45 Theobroma cacao Malaysia: Melaka PQ835688 PQ839587 Present study A. tubingensis JN46 Theobroma cacao Malaysia: Melaka PQ835689 PQ839588 Present study A. tubingensis PJ42 Theobroma cacao Malaysia: Penang PQ835685 PQ839584 Present study A. tubingensis TK43 Theobroma cacao Malaysia: Selangor PQ835686 PQ839585 Present study A. sydowii CBS 593.65 ET Dairy product Greece EU982025 EF652274 Krimitzas et al. 2013 EN Ex-neotype culture, ET Ex-type culture Mycotoxin screening using coconut cream agar (CCA) All fungal isolates obtained were screened for their potential to produce AF and OTA using coconut cream agar (CCA) (Maman et al. 2021 ). Seven-day-old pure cultures grown on PDA were subcultured onto CCA plates. To prevent conidial dispersion, a drop of sterile paraffin oil was applied to the center of each plate. The inoculated plates were incubated at 25 ± 2°C for 10 days. A CCA plate without fungal inoculation served as a negative control, while a plate inoculated with a non-mycotoxigenic fungal species ( Colletotrichum sp.) was used as a positive control to validate the assay. After the incubation period, the reverse side of each CCA plate was examined under ultraviolet (UV) light at a wavelength of 365 nm. The presence fluorescence was used as an indicator of mycotoxin production. Results Fungal isolation and enumeration A total of 43 Aspergillus isolates were recovered from dried cocoa bean samples collected from 12 farms across Peninsular Malaysia. To quantify fungal contamination, serial dilutions were performed on each sample, followed by plating on PDA to determine colony-forming units per gram (CFU/g). The fungal counts are summarized in Table 2 . Most samples exhibited fungal loads below the maximum permissible limit of 1 × 10⁴ CFU/g, as stipulated by food hygiene standards (Good Manufacturing Practice 2008 ). However, the sample from Pokok Jenerih, Penang, recorded a higher fungal count of 2.9 × 10⁵ CFU/g, exceeding the recommended threshold and indicating potential post-harvest contamination issues. Table 2 Aspergillus species isolated from dried cocoa beans collected across various locations in Peninsular Malaysia. The table includes fungal enumeration (CFU/g), sequence similarity to reference isolates in GenBank, and fluorescence intensity on coconut cream agar (CCA) Sampling location a Fungal enumeration (CFU/g) Fungal species Fungal code Sequence similarity with GenBank b Fluorescence on CCA ITS tub2 Bagan Datuk, Perak 1.3 x 10³ A. flavus BD21 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + BD22 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + A. tamarii BD11 MH854614 (99%) EF203122 (100%) EF203123 (99%) + Chemor, Perak 1.1 x 10³ A. niger CR31 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ A. tubingensis CR41 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + Pokok Jenerih, Penang 2.9 x 10⁵ A. flavus PJ23 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + A. niger PJ32 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ PJ33 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ A. tamarii PJ12 MH854614 (99%) EF203122 (100%) EF203123 (99%) + PJ13 MH854614 (99%) EF203122 (100%) EF203123 (99%) + PJ14 MH854614 (99%) EF203122 (100%) EF203123 (99%) + PJ15 MH854614 (99%) EF203122 (100%) EF203123 (99%) + A. tubingensis PJ42 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + Sungai Bakap, Penang 1.2 x 10³ A. flavus SB24 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + SB25 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + Tanjung Karang, Selangor 1.8 x 10³ A. niger TK34 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ TK35 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ A. tubingensis TK43 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + Parit Tujuh, Selangor 2.2 x 10³ A. aculeatus PT51 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + PT52 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + Tanjung Ipoh, Negeri Sembilan 1.4 x 10³ A. aculeatus TI53 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + TI54 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + TI55 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + A. niger TI36 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ Rembau, Negeri Sembilan 2.4 x 10³ A. aculeatus RU56 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + A. niger RU37 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ RU38 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ Durian Tunggal, Melaka 1.8 x 10³ A. aculeatus DT57 FJ629320 (100%) FJ629271(99%) AY585538 (99%) + A. flavus DT26 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + DT27 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + DT28 MH854614 (99%) KJ175479 (99%) KJ175478 (98%) + A. tamarii DT16 MH854614 (99%) EF203122 (100%) EF203123 (99%) + DT17 MH854614 (99%) EF203122 (100%) EF203123 (99%) + Jasin, Melaka 2.0 x 10³ A. flavus JN29 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + JN210 MH864264 (100%) KJ175479 (99%) KJ175478 (98%) + A. tubingensis JN44 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + JN45 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + JN46 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + Batu Pahat, Johor 1.8 x 10 3 A. niger BP39 AY585553 (99%) AY820002 (99%) AY820001 (98%) ++ A. tubingensis BP47 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + BP48 FJ629354 (100%) FJ629305 (98%) AY820009 (99%) + Tangkak, Johor 1.4 x 10³ A. tamarii TA18 MH854614 (99%) EF203122 (100%) EF203123 (99%) + TA19 MH854614 (99%) EF203122 (100%) EF203123 (99%) + Morphological and molecular identification Initial identification based on colony morphology grouped the 43 Aspergillus isolates into five distinct species. Molecular confirmation was performed through DNA sequencing of the ITS region and the tub2 gene. BLAST analysis against the GenBank database confirmed all isolates with 98–100% sequence similarity to known Aspergillus species. The identified species included A. aculeatus , A. flavus , A. niger , A. tamarii , and A. tubingensis (Table 2 ). The morphological characteristics of each species were consistent with descriptions by Pitt and Hocking ( 2009 ) and Samson et al. ( 2019 ). Among the Aspergillus species isolated, A. aculeatus was represented by seven isolates, accounting for 16% of the total. These isolates were recovered from four different samples, resulting in a prevalence rate of 33.3% (Table 2 ). On both MEA and CYA, the colonies appeared dark brown and granular, with the reverse sides ranging from white to pale yellow. After three days of incubation, colony diameters measured between 3.0 and 3.2 cm on MEA and 3.1 to 3.3 cm on CYA. Microscopic examination revealed subspheroidal to ellipsoidal conidia measuring 1.4–2.0 µm in diameter. The conidiophores were smooth, hyaline, and measured 85–95 µm in length (Table 3). Aspergillus flavus exhibited the highest isolation frequency at 23%, with five isolates recovered from five different samples, corresponding to a prevalence rate of 41.7% (Table 2 ). On CYA, colonies were green-yellow, while on MEA, they appeared yellowish-white with a floccose, tufted texture. Colony diameters ranged from 1.6 to 1.8 cm on CYA and 2.2 to 2.4 cm on MEA. Over time, sclerotia developed on the colonies, initially white and gradually turning black. The reverse side of the colonies was cream-colored. Microscopically, the conidia were globose with rough walls, and the conidial heads were typically radiate, measuring 300–400 µm in diameter. Vesicles were spherical to sub-spherical, and the conidiophores were hyaline with roughened walls (Table 3). Aspergillus niger was the most prevalent species, represented by nine isolates (21%) and detected in six samples, resulting in a prevalence rate of 50% (Table 2 ). Colonies on CYA were dark brown, while those on MEA appeared black. The reverse sides of the colonies were cream to dull yellow on both media. The isolates exhibited rapid growth, with colony diameters ranging from 1.9 to 2.1 cm on CYA and 1.9 to 2.3 cm on MEA. Microscopically, the conidia were globose, brownish to black, and verruculose. The conidiophores were long, smooth, aseptate, and terminated in swollen apices forming vesicles. The conidial heads were dark brown to black, spherical to radiate, and measured 300–500 µm in diameter. Vesicles were spherical and covered with short terminal branches (metulae) (Table 3). Similarly, A. tamarii was represented by nine isolates (21%) and was recovered from four samples, indicating a prevalence rate of 33.3% (Table 2 ). On CYA, colonies were powdery olive-green, while on MEA, they appeared yellowish-green. A distinct white margin surrounded the colonies on both media. The reverse sides were green on CYA and brown on MEA. The isolates grew rapidly at 25 ± 2°C, with colony diameters ranging from 1.4 to 1.7 cm on CYA and 1.7 to 1.9 cm on MEA. The conidia were globose to subglobose with rough surfaces. Conidial heads were globose to radiate, measuring 300–500 µm in diameter. The conidiophores were long, unbranched, hyaline, and rough-walled, terminating in broadly clavate vesicles (Table 3). Aspergillus tubingensis was identified in eight isolates (19%) recovered from five samples, corresponding to a prevalence rate of 41.7% (Table 2 ). Colonies on CYA were dark brown, while those on MEA appeared black. The reverse sides of the colonies were creamish dull yellow to yellow on both media. The isolates exhibited rapid growth, with colony diameters ranging from 2.4 to 2.6 cm on CYA and 2.7 to 3.5 cm on MEA. Microscopically, the conidia were globose and brownish to black, and the conidiophores were smooth and elongated. The conidial heads were globose to radiate, measuring 300–500 µm in diameter (Table 3). Phylogenetic analysis Phylogenetic trees constructed individually from ITS and tub2 sequences were insufficient to clearly resolve the species-level relationships among the Aspergillus isolates (Supplementary Figs. 1 and 2). In both trees, several isolates clustered ambiguously with multiple reference species, indicating limited discriminatory power when using single-locus data. To overcome this limitation, a concatenated dataset combining ITS and tub2 sequences was used to generate a ML phylogenetic tree (Fig. 2 ). The resulting tree provided significantly improved resolution, with all isolates forming well-supported clades (95–99%) corresponding to their respective Aspergillus species. Bootstrap values at key nodes were high, indicating strong statistical support for the monophyly of each species group. Distinct clades were clearly delineated and visually highlighted using colored boxes, each representing a different Aspergillus species. This combined-locus approach successfully resolved the taxonomic placement of all isolates, confirming their identities as A. aculeatus , A. flavus , A. niger , A. tamarii , and A. tubingensis . These results were fully consistent with BLAST analyses, reinforcing the reliability of the molecular identifications. Mycotoxigenic potential of Aspergillus isolates The mycotoxigenic potential of Aspergillus isolates was assessed using CCA, a qualitative medium that detect the presence of fluorescent secondary metabolites. All 43 isolates tested exhibited fluorescence, with varying levels of intensity. Low fluorescence intensity (‘+’) was observed in all isolates of A. flavus , A. tamarii , A. tubingensis , and A. aculeatus . In contrast, moderate fluorescence intensity (‘++’) was observed exclusively in A. niger isolates. A total of nine A. niger isolates from Chemor, Pokok Jenerih, Tanjung Karang, Tanjung Ipoh, Rembau, and Batu Pahat displayed this level of fluorescence. No isolates showed an absence of fluorescence or high fluorescence intensity (‘+++’) (Table 2 ) (Fig. 3 ). Discussion The recovery of 43 Aspergillus isolates from dried cocoa beans across 12 farms in Peninsular Malaysia underscores the ubiquitous presence of this fungal genus in post-harvest cocoa environments. Fungal load variation among samples can be attributed to differences in farm-level practices, including harvesting, fermentation, drying, and climatic factors. Notably, while most cocoa samples showed fungal counts below the generally accepted threshold of 1 × 10⁴ CFU/g, a significantly elevated count of 2.9 × 10⁵ CFU/g was recorded at Pokok Jenerih. This localized anomaly likely results from suboptimal drying conditions, a factor known to promote fungal proliferation (Mounjouenpou et al. 2008 ; Sánchez-Hervás et al. 2008 ). Additionally, biological vectors such as insects, birds, and rodents may introduce spores during the sun-drying phase, further exacerbating contamination risks (Delgado-Ospina et al. 2021 ). To mitigate such risks, strict adherence to the Codex Alimentarius Commission’s Code of Practice (2013) is essential. The guidelines recommend spreading cocoa beans in layers not exceeding 6 cm, ensuring uniform drying to a final moisture content of 6–8%, and avoiding re-wetting by covering beans during inclement weather or nighttime (Copetti et al. 2014 ). These post-harvest handling protocols are vital for minimizing fungal growth and reducing the risk of subsequent mycotoxin contamination. Morphological and molecular analyses identified five Aspergillus species in dried cocoa beans namely A. aculeatus , A. flavus , A. niger , A. tamarii , and A. tubingensis . The dual use of ITS and tub2 region/gene sequences provided robust resolution at the species level, with BLAST and phylogenetic analysis showing high concordance. The predominance of A. flavus (23%), A. niger (21%), and A. tamarii (21%) is particularly significant, as these species are frequently implicated in food spoilage and mycotoxin contamination. These findings align with previous studies reporting the presence of Aspergillus sections Flavi and Nigri in cocoa (Guehi et al. 2007 ; Sánchez-Hervás et al. 2008 ; Nugroho et al. 2013 ). The widespread presence of these species suggests they are well-adapted to the cocoa bean microenvironment, likely due to their thermotolerance and ability to survive in low-moisture substrates. Morphological characteristics were consistent with established taxonomic keys (Pitt and Hocking 2009 ; Samson et al. 2019 ), further validating the identifications. Among the isolates, A. flavus was the most frequently recovered species, with 10 isolates identified. This species is a known producer of AFs, particularly aflatoxin B₁ (AFB 1 ), one of the most potent naturally occurring carcinogens (Klich et al. 2000 ). However, accurate identification can be challenging due to its morphological and genetic similarity to A. oryzae . Despite these challenges, the concatenated ITS– tub2 phylogenetic tree clearly resolved A. flavus and A. oryzae into distinct clades, confirming the accuracy of species-level discrimination in this study. This distinction is critical, given that A. flavus is toxigenic, whereas A. oryzae is generally non-aflatoxigenic (Wei and Jong 1986 ; Nakamura et al. 2011 ). Aspergillus tamarii , another member of section Flavi , was also frequently isolated. Though generally considered weakly or non-aflatoxigenic, some isolates have been shown to produce aflatoxins under conducive conditions. Klich et al. ( 2000 ) reported that A. tamarii possesses genes homologous to the AF biosynthesis cluster and can produce both AFs and sterigmatocystin under specific environmental conditions. Thus, while not typically a major aflatoxin contributor, its toxigenic potential should not be entirely dismissed in tropical post-harvest environments. Species in section Nigri , including A. niger , A. tubingensis , and A. aculeatus , share overlapping morphological features such as dark conidia and similar colony textures, complicating species identification using phenotypic traits alone. These black Aspergilli are commonly found in food and agricultural products and are recognized both for their industrial applications and for their potential to produce ochratoxins (Nielsen et al. 2009 ). Due to the limited resolution of ITS sequences alone, this study adopted a multi-locus approach combining ITS and tub2 markers. This method significantly improved species-level resolution, as previously demonstrated by Zulkifli and Zakaria ( 2017 ) and Yodsing et al. ( 2018 ). Phylogenetic analysis revealed distinct clades for each species with strong bootstrap support (96–99%), confirming the efficacy of this approach in resolving cryptic species within section Nigri . The use of CCA medium provided preliminary insights into the mycotoxin-producing potential of the recovered isolates. Fluorescent metabolites were detected in all Aspergillus isolates, suggesting widespread biosynthetic capacity for mycotoxin production. The fluorescence observed under UV light, indicative of metabolic activity, varied among species. Notably, moderate fluorescence intensity was recorded only in A. niger isolates, suggesting potential production of OTA or fumonisins (FBs), known secondary metabolites of this species. Interestingly, A. flavus isolates exhibited only low fluorescence. This unexpected result may reflect inter-isolate variability, environmental influences, or limitations of the CCA assay’s sensitivity to AFs. CCA’s high lipid content, particularly its medium-chain fatty acids like lauric acid, provides a conducive substrate for toxin biosynthesis (Dyer and McCammon 1994 ). Nevertheless, as the assay is qualitative, confirmatory analysis using advanced techniques such as high-performance liquid chromatography (HPLC) or liquid chromatography–mass spectrometry (LC-MS/MS) is strongly recommended to validate and quantify mycotoxin production (Mohamed et al. 2013 ). This study confirms the diversity and prevalence of Aspergillus species in dried cocoa beans in Peninsular Malaysia, with species such as A. flavus , A. niger , and A. tamarii dominating the post-harvest fungal community. The combination of morphological and molecular tools enabled accurate species-level identification, while preliminary assessments of toxigenic potential highlighted the need for further mycotoxin quantification. The findings reinforce the importance of good post-harvest practices and molecular surveillance to safeguard cocoa quality and food safety. Declarations Conflict of interest: The authors declare no competing interests. Funding This research was supported by Universiti Sains Malaysia through the Bridging Grant (Project No: R501-LR-RND003-0000000743-0000) and the Fundamental Research Grant Scheme (FRGS/1/2019/WAB01/USM/02/1) provided by the Ministry of Higher Education, Malaysia. The first author, Nuraina Syazana, also received financial support through the Postgraduate Research Attachment (PGRA) from the Institute of Postgraduate Studies (IPS), Universiti Sains Malaysia. Author Contribution Conceptualization, SN-S and MHM; Experimentation, data collection and formal analysis, SN-S, MZZ, and NSS; Writing—original draft preparation, SN-S; Writing—review and editing, SN-S, MZZ, NSS, Y-HL, and MHM; Supervision and resources, MHM and Y-HL. All authors have reviewed the manuscript. Acknowledgement The authors extend their sincere gratitude to the Malaysian Cocoa Board (MCB) for granting permission to collect cocoa bean samples. Special appreciation is also given to the MCB staff for their invaluable assistance and support during fieldwork activities. Data Availability Data is provided within the manuscript. References Agus BAP, Mohamad NN, Hussain N (2018) Composition of unfermented, unroasted, roasted cocoa beans and cocoa shells from Peninsular Malaysia. 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J Food Sci Eng 3(9):472 Ooi TS, Ting ASY, Siow LF (2020) Influence of selected native yeast starter cultures on the antioxidant activities, fermentation index and total soluble solids of Malaysia cocoa beans: A simulation study. LWT 122:108977 Peterson SW (2008) Phylogenetic analysis of Aspergillus species using DNA sequences from four loci. Mycologia 100(2):205–226 Pildain MB, Frisvad JC, Vaamonde G, Cabral D, Varga J, Samson RA (2008) Two novel aflatoxin-producing Aspergillus species from Argentinean peanuts. Int J Syst Evol Microbiol 58(3):725–735 Pitt JI, Hocking AD (2009) Fungi and Food Spoilage. In Springer eBooks Samson RA, Houbraken JAMP, Kuijpers AF, Frank JM, Frisvad JC (2004) New ochratoxin A or sclerotium producing species in Aspergillus section Nigri . Stud Mycol 50(1):45–56 Samson RA, Houbraken J, Thrane U, Frisvad JC, Andersen B (2019) Food and Indoor Fungi (2nd ed.) Westerdijk Fungal Biodiversity Institute Westerdijk Laboratory Manual Series Vol. 2 Sánchez-Hervás M, Gil JV, Bisbal F, Ramón D, Martínez-Culebras PV (2008) Mycobiota and mycotoxin producing fungi from cocoa beans. Int J Food Microbiol 125(3):336–340 Tamura K (1992) Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G + C-content biases. Mol Biol Evol 9(4):678–687 Varga J, Frisvad JC, Samson R (2011) Two new aflatoxin producing species, and an overview of Aspergillus section Flavi . Stud Mycol 69(1):57–80 Wei D, Jong S (1986) Production of aflatoxins by strains of the Aspergillus flavus group maintained in ATCC. Mycopathologia 93(1):19–24 White TJ, Bruns T, Lee SJWT, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR protocols: a guide to methods and applications. 18(1):315–322 Yodsing N, Lekphrom R, Sangsopha W, Aimi T, Boonlue S (2018) Secondary metabolites and their biological activity from Aspergillus aculeatus KKU-CT2. Curr Microbiol 75:513–518 Zulkifli NA, Zakaria L (2017) Morphological and molecular diversity of Aspergillus from corn grain used as livestock feed. HAYATI J Biosci 24(1):26–34 Tables Tables 1 to 3 are available in the Supplementary Files section. Additional Declarations No competing interests reported. Supplementary Files Table1Referencesequencesforphylogeneticanalysis.docx Table 1 Fungal isolates and their corresponding GenBank accession numbers used in the phylogenetic analysis. Table2.docx Table 2 Aspergillus species isolated from dried cocoa beans collected across various locations in Peninsular Malaysia. The table includes fungal enumeration (CFU/g), sequence similarity to reference isolates in GenBank, and fluorescence intensity on coconut cream agar (CCA). Table3Morphologicalcharacteristics.jpg Table 3 Morphological characteristics of Aspergillus species on CYA and MEA after 7 days of incubation. SupplementaryFigure1ITSMLtree.docx Supplementary Fig. 1 Maximum likelihood (ML) tree based on ITS sequences. Isolates from this study are shown in bold, and Aspergillus sydowii is used as the outgroup. Bootstrap values ≥70% are shown at the nodes. SupplementaryFigure2BetatubulinMLtree.docx Supplementary Fig. 2 Maximum likelihood (ML) tree based on tub2 sequences. Isolates from this study are shown in bold, and Aspergillus sydowii is used as the outgroup. Bootstrap values ≥70% are shown at the nodes. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6760467","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":465989903,"identity":"49000644-2e32-43bb-b60b-1cf70bc6ec36","order_by":0,"name":"Sahariman Nuraina-Syazana","email":"","orcid":"","institution":"School of Biological Sciences, Universiti Sains Malaysia","correspondingAuthor":false,"prefix":"","firstName":"Sahariman","middleName":"","lastName":"Nuraina-Syazana","suffix":""},{"id":465989904,"identity":"bad3a5ae-9935-4def-8996-f478928b1407","order_by":1,"name":"Muhamad Zaid Zakaria","email":"","orcid":"","institution":"Food and Industrial Chemistry Section, Environmental and Industrial Analysis Division, Department of Chemistry Malaysia, Penang State","correspondingAuthor":false,"prefix":"","firstName":"Muhamad","middleName":"Zaid","lastName":"Zakaria","suffix":""},{"id":465989905,"identity":"23f98dda-3e14-4725-b506-c47ae3ae7770","order_by":2,"name":"Nor Shifa Shuib","email":"","orcid":"","institution":"Food and Industrial Chemistry Section, Environmental and Industrial Analysis Division, Department of Chemistry Malaysia, Penang State","correspondingAuthor":false,"prefix":"","firstName":"Nor","middleName":"Shifa","lastName":"Shuib","suffix":""},{"id":465989906,"identity":"7659f301-0dbe-488e-8b9e-f40fcb0fe64d","order_by":3,"name":"Yin-Hui Leong","email":"","orcid":"","institution":"National Poison Centre, Universiti Sains Malaysia","correspondingAuthor":false,"prefix":"","firstName":"Yin-Hui","middleName":"","lastName":"Leong","suffix":""},{"id":465989907,"identity":"5b3fec4e-8b3b-4371-8e99-8f2a975df111","order_by":4,"name":"Masratul Hawa Mohd","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAy0lEQVRIiWNgGAWjYBADGQMQ+YF4DQkMPCAtjDNI1sLMQ4xi3fbjzz78/HGYx5z9jOFn27a6aAaxMwZ4tZidyTGe2ZNwmMeyJ8dYOrftcG6DdA4BLQdymBl4gFoMDuRuAGo5QISW888fM/4BaTn/dvNvy7Y6IrTcSDBmBttyI3ebNGMbMzFa3hgzy6SlA7W8/2bZc+5wbpt0WgEBh6U/ZnxjYy1ncD4t+caPsrrcfunkDXi1YAI2Bg78DsMG2B+QrGUUjIJRMAqGNQAAghpHuFLVaLwAAAAASUVORK5CYII=","orcid":"","institution":"School of Biological Sciences, Universiti Sains Malaysia","correspondingAuthor":true,"prefix":"","firstName":"Masratul","middleName":"Hawa","lastName":"Mohd","suffix":""}],"badges":[],"createdAt":"2025-05-27 14:38:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6760467/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6760467/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":84080456,"identity":"85922723-e753-4863-ac5f-f58718ce6233","added_by":"auto","created_at":"2025-06-06 14:02:19","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":219232,"visible":true,"origin":"","legend":"\u003cp\u003eSampling locations of dried cocoa beans collected from 12 farms across Peninsular Malaysia.\u003c/p\u003e","description":"","filename":"Figure1Samplinglocations.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/d7ebb1e0722ab2cb44bad63c.jpg"},{"id":84080013,"identity":"618b2689-8405-4a13-a89d-9640513b04cd","added_by":"auto","created_at":"2025-06-06 13:54:19","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":650443,"visible":true,"origin":"","legend":"\u003cp\u003eConsensus maximum likelihood (ML) phylogenetic tree based on concatenated ITS and\u003cem\u003e tub2\u003c/em\u003e sequences. Isolates from this study are shown in bold, and \u003cem\u003eAspergillus sydowii\u003c/em\u003e is used as the outgroup. Bootstrap support values ≥70% are indicated at the nodes. The scale bar represents the expected number of nucleotide substitutions per site.\u003c/p\u003e","description":"","filename":"Figure2Phylogenetictree.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/3e5ada531dffd20b0c656faa.jpg"},{"id":84080019,"identity":"351c416e-27a0-47a9-b211-fa75ec4cb4b5","added_by":"auto","created_at":"2025-06-06 13:54:20","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":2235533,"visible":true,"origin":"","legend":"\u003cp\u003eMycotoxigenic potential of \u003cem\u003eAspergillus\u003c/em\u003eisolates grown on coconut cream agar (CCA) after 10 days of incubation. Low fluorescence intensity was observed in \u003cem\u003eA. aculeatus\u003c/em\u003e (A), \u003cem\u003eA. flavus\u003c/em\u003e(B), \u003cem\u003eA. tamarii\u003c/em\u003e (C), and \u003cem\u003eA. tubingensis\u003c/em\u003e (D). Moderate fluorescence intensity was observed in \u003cem\u003eA. niger\u003c/em\u003e (E). \u003cem\u003eColletotrichum\u003c/em\u003esp. was included as a non-mycotoxigenic positive control (F), and a negative control (G).\u003c/p\u003e","description":"","filename":"Figure3Mycotoxigenicpotential.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/4361100e3167c9d0503fba92.jpg"},{"id":85577853,"identity":"b79a81f6-8221-4df6-b8a2-3ab54db5daf2","added_by":"auto","created_at":"2025-06-27 20:01:29","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4578400,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/7e80018d-23cd-4ae9-955d-dba8dc41897a.pdf"},{"id":84080012,"identity":"566756a5-5e69-4c28-a78e-9838a2bae175","added_by":"auto","created_at":"2025-06-06 13:54:19","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":40588,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 1 \u003c/strong\u003eFungal isolates and their corresponding GenBank accession numbers used in the phylogenetic analysis.\u003c/p\u003e","description":"","filename":"Table1Referencesequencesforphylogeneticanalysis.docx","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/7ff2f691621df2ba7c70fe7e.docx"},{"id":84080015,"identity":"7171d81c-5f57-4e53-a993-fa8463ff99b9","added_by":"auto","created_at":"2025-06-06 13:54:19","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":29951,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e \u003cem\u003eAspergillus\u003c/em\u003e species isolated from dried cocoa beans collected across various locations in Peninsular Malaysia. The table includes fungal enumeration (CFU/g), sequence similarity to reference isolates in GenBank, and fluorescence intensity on coconut cream agar (CCA).\u003c/p\u003e","description":"","filename":"Table2.docx","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/eef2a8bfccc09732b14abe83.docx"},{"id":84080020,"identity":"0eea354d-de42-4ac9-9958-cd4013c69da8","added_by":"auto","created_at":"2025-06-06 13:54:20","extension":"jpg","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":8313055,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 3\u003c/strong\u003e Morphological characteristics of \u003cem\u003eAspergillus\u003c/em\u003e species on CYA and MEA after 7 days of incubation.\u003c/p\u003e","description":"","filename":"Table3Morphologicalcharacteristics.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/6ba1689d288d8e2f264efc44.jpg"},{"id":84080017,"identity":"edf6515d-1968-4aef-9502-32e759c63b32","added_by":"auto","created_at":"2025-06-06 13:54:20","extension":"docx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":105896,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSupplementary Fig. 1 \u003c/strong\u003eMaximum likelihood (ML) tree based on ITS sequences. Isolates from this study are shown in bold, and \u003cem\u003eAspergillus sydowii\u003c/em\u003e is used as the outgroup. Bootstrap values ≥70% are shown at the nodes.\u003c/p\u003e","description":"","filename":"SupplementaryFigure1ITSMLtree.docx","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/4c199d720fe6c5ca4395c96e.docx"},{"id":84080016,"identity":"be8dbc07-21f7-46c2-b859-eb66cadacfb1","added_by":"auto","created_at":"2025-06-06 13:54:20","extension":"docx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":23318,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSupplementary Fig. 2 \u003c/strong\u003eMaximum likelihood (ML) tree based on \u003cem\u003etub2 \u003c/em\u003esequences. Isolates from this study are shown in bold, and \u003cem\u003eAspergillus sydowii\u003c/em\u003e is used as the outgroup. Bootstrap values ≥70% are shown at the nodes.\u003c/p\u003e","description":"","filename":"SupplementaryFigure2BetatubulinMLtree.docx","url":"https://assets-eu.researchsquare.com/files/rs-6760467/v1/e49f97366c4c5f3fd8fc27bd.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Characterization of Aspergillus species associated with dried cocoa beans in Peninsular Malaysia","fulltext":[{"header":"Introduction","content":"\u003cp\u003eCocoa (\u003cem\u003eTheobroma cocoa\u003c/em\u003e) is a globally significant crop and the primary raw material in chocolate production. While West Africa remains the leading cocoa-producing region, substantial cultivation also occurs in Asia, Central America, and South America (Copetti et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The \u003cem\u003eT. cacao\u003c/em\u003e tree thrives in tropical climates characterized by high humidity, consistent rainfall (1,500\u0026ndash;3,000 mm annually), and temperatures ranging from 20\u0026deg;C to 28\u0026deg;C. The fruit, known as a cocoa pod, contains 25\u0026ndash;75 seeds embedded in a sweet, mucilaginous pulp. Upon drying, these seeds transition from white to violet or reddish-brown (Kim et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCocoa beans undergo several post-harvest processing steps before consumption, beginning with manual pod harvesting and opening. The extracted beans are naturally fermented in containers for several days, followed by sun drying on wooden platforms, cement surfaces, or directly on the ground. Once dried, the beans are stored, bagged, and transported to processing facilities where they are roasted, shelled, and ground into cocoa products. Each of these stages presents opportunities for microbial contamination, particularly by fungi introduced via pod surfaces, handling tools, workers\u0026rsquo; hands, insects, and residual mucilage (Copetti et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFungal contamination is especially prevalent during the drying phase, where environmental exposure facilitates the growth of mycotoxigenic species. Several studies have reported the presence of \u003cem\u003eAspergillus\u003c/em\u003e species in cocoa beans, including those capable of producing aflatoxins (AFs) and ochratoxin A (OTA), mycotoxins of significant concern due to their toxicity and prevalence in food products (Mounjouenpou et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Copetti et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Kedjebo et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Mycotoxin contamination has been documented in cocoa-producing countries such as C\u0026ocirc;te d\u0026rsquo;Ivoire (Guehi et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), Cameroon (Mounjouenpou et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), and Brazil (de Magalh\u0026atilde;es et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Aflatoxins and OTA are particularly hazardous due to their carcinogenic, nephrotoxic, and immunosuppressive effects (Copetti et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUnderstanding the diversity and toxigenic potential of fungal species associated with cocoa beans is essential for assessing food safety risks and improving post-harvest practices. While most Malaysian studies have focused on the biochemical and physiological properties of cocoa (Farah et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Agus et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Ooi et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), limited research has addressed fungal contamination and mycotoxin risks. Therefore, this study aims to identify \u003cem\u003eAspergillus\u003c/em\u003e species associated with dried cocoa beans in Peninsular Malaysia using morphological, molecular, and phylogenetic approaches, and to evaluate their potential to produce mycotoxins.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eSample collection and fungal isolation\u003c/h2\u003e \u003cp\u003eA total of 12 dried cocoa bean samples were collected from 12 farms across six states in Malaysia namely Penang, Perak, Selangor, Negeri Sembilan, Johor, and Melaka (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). To eliminate surface contaminants, the beans were surface-sterilized by sequential immersion in 70% ethanol (C₂H₅OH) and 1% sodium hypochlorite (NaOCl) for 3 min each. The samples were then rinsed three times with sterile distilled water for 1 min per rinse and air-dried on sterile filter paper under aseptic conditions. The sterilized beans were plated onto potato dextrose agar (PDA) and incubated at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C for 3\u0026ndash;5 days, or until visible mycelial growth emerged. Actively growing mycelia were subcultured onto fresh PDA plates and incubated under the same conditions for 7 days to promote sporulation. To obtain pure cultures, single conidial isolation was performed by streaking a conidial suspension onto PDA using a sterile inoculating loop. Hyphal tips from 48-hour-old cultures were then excised and transferred to new PDA plates to establish monoconidial isolates for further analysis.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eFungal enumeration\u003c/h3\u003e\n\u003cp\u003eA total of 5 g of dried cocoa beans were weighed and ground into a fine powder using a sterile mortar and pestle. One gram of the homogenized sample was transferred into a universal bottle containing 9 mL of sterile distilled water to prepare a 10⁻\u0026sup1; dilution. The mixture was thoroughly shaken to ensure uniform suspension. Subsequent serial dilutions were prepared up to 10⁻⁵. From each dilution, 0.1 mL was aseptically spread onto PDA plates in triplicate. The inoculated plates were incubated at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C for 5 days. After incubation, fungal colonies were counted, and the colony-forming units per gram (CFU/g) of sample were calculated following the method described by Ntuli et al. (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eMorphological characterization\u003c/h3\u003e\n\u003cp\u003ePure fungal isolates obtained were cultured on Czapek yeast extract agar (CYA) and malt extract agar (MEA) to assess their morphological characteristics. To prevent conidial dispersion, a drop of sterilized paraffin oil was placed at the center of each plate. The cultures were incubated at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C for 7 days. Macroscopic features, including mycelial growth rate, were measured after 3 days of incubation. Colony morphology such as texture, color, and pigmentation on the reverse side was evaluated after 7 days on both CYA and MEA. For microscopic examination, fungal structures including conidia, phialides, and conidiophores were observed using the slide mount method, with preparations made on glass slides using sterile distilled water as the mounting medium. Observations were conducted using a compound light microscope (CX41, Olympus, Japan) equipped with a digital camera (KY-F55BE, JVC, Japan) to document morphological features.\u003c/p\u003e\n\u003ch3\u003eMolecular identification and phylogenetic analysis\u003c/h3\u003e\n\u003cp\u003eMolecular identification was carried out to confirm the identity of the fungal isolates obtained. Each isolate was cultured in potato dextrose broth (PDB) and incubated at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C for 2 days. The resulting mycelia were harvested, air-dried on sterile filter paper, and ground into a fine powder using liquid nitrogen. Approximately 50 mg of the powdered mycelia was transferred into a sterile 1.5 mL microcentrifuge tube for genomic DNA extraction. DNA was extracted using the InnuPREP Plant DNA Kit (Analytik Jena, Germany), following the manufacturer\u0026rsquo;s protocol.\u003c/p\u003e \u003cp\u003eTwo genetic markers were targeted for amplification namely the internal transcribed spacer (ITS) region and the β-tubulin (\u003cem\u003etub2\u003c/em\u003e) gene. The ITS region was amplified using the primer pair ITS1 (5\u0026prime;-TCCGTAGGTGAACCTGCGG-3\u0026prime;) and ITS4 (5\u0026prime;-TCCTCCGCTTATTGATATGC-3\u0026prime;) (White et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), while the \u003cem\u003etub2\u003c/em\u003e gene was amplified using primers Bt2a (5\u0026prime;-GGTAACCAAATCGGTGCTGCTTTC-3\u0026prime;) and Bt2b (5\u0026prime;-ACCTCAGTGTAGTGACCCTTGGC-3\u0026prime;) (Glass and Donaldson \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). PCR reactions were prepared in 50 \u0026micro;L volumes containing 8 \u0026micro;L of 5\u0026times; Green GoTaq buffer, 8 \u0026micro;L of 25 mM MgCl₂, 1 \u0026micro;L of 10 mM dNTP mix, 8 \u0026micro;L of each primer (5 \u0026micro;M for ITS and 1 \u0026micro;M for \u003cem\u003etub2\u003c/em\u003e), 0.3 \u0026micro;L of Taq polymerase (5 U/\u0026micro;L), 1 \u0026micro;L of genomic DNA, and 15.7 \u0026micro;L of sterile distilled water. Amplification was performed using a MyCycler Thermal Cycler (Bio-Rad, USA). For the ITS region, the thermal cycling conditions included an initial denaturation at 94\u0026deg;C for 1 min 35 s, followed by 35 cycles of denaturation at 95\u0026deg;C for 35 s, annealing at 59\u0026deg;C for 55 s, and extension at 72\u0026deg;C for 1 min 30 s, with a final extension at 72\u0026deg;C for 10 min. For the \u003cem\u003etub2\u003c/em\u003e gene, the protocol included an initial denaturation at 94\u0026deg;C for 5 min, followed by 35 cycles of denaturation at 94\u0026deg;C for 1 min, annealing at 57\u0026deg;C for 1 min, and extension at 72\u0026deg;C for 1 min, with a final extension at 72\u0026deg;C for 7 min.\u003c/p\u003e \u003cp\u003ePCR products were separated by electrophoresis on 1.0% agarose gels stained with FloroSafe DNA stain (First BASE) in 1\u0026times; TBE buffer. Electrophoresis was conducted at 80 V and 400 mA for 90 min. DNA bands were visualized using a Bio-Rad Gel Doc\u0026trade; XR System and documented with Quantity One\u0026reg; software. Fragment sizes were estimated using a 100 bp GeneRuler\u0026trade; DNA ladder (Thermo Scientific, USA). The amplified products were then sent to First BASE Laboratories Sdn Bhd (Seri Kembangan, Malaysia) for purification and sequencing.\u003c/p\u003e \u003cp\u003eThe resulting DNA sequences were aligned using MEGA12 software (Kumar et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Sequence identities were confirmed through BLAST searches against the NCBI GenBank database. All sequences generated were submitted to GenBank. Phylogenetic relationships were inferred using the maximum likelihood (ML) method in MEGA12. Prior to tree construction, the best-fit nucleotide substitution model was determined. Individual gene trees were constructed using the Kimura 2-parameter model (Kimura \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1980\u003c/span\u003e), while the combined ITS and \u003cem\u003etub2\u003c/em\u003e dataset was analyzed using the Tamura 3-parameter model (Tamura \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1992\u003c/span\u003e), with 1,000 bootstrap replications to assess branch support. Reference sequences and accession numbers used in the phylogenetic analysis are listed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFungal isolates and their corresponding GenBank accession numbers used in the phylogenetic analysis\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eAspergillus\u003c/em\u003e species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eIsolate\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eHost\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eGenBank accession number\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eReference\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eITS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003etub2\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 121060\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eCoffea arabica\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eThailand\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU159211\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU159220\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 121871\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eCoffea robusta\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eThailand\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU159206\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU159227\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 121877\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eCoffee arabica\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eThailand\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU159213\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU159224\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 172.66\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIndia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 11480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIndia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585557\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY585539\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 62078\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNot stated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585556\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY585538\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT57\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835672\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654129\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePT51\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835666\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654123\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePT52\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835667\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654124\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTI53\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835668\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654125\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTI54\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835669\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654126\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTI55\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835670\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654127\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eRU56\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835671\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePV654128\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. arachidicola\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 117610\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eArachis glabrata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eArgentina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEF409241\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203158\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePildain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. awamori\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 557.65\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNot stated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAM087614\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. awamori\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATHUM 5181\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGreece\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU982010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU982069\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. awamori\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNRRL 4951\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBrazil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eKC796388\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKC796358\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. caelatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 763.97\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH862672\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203127\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. fasciculatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 110.55\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAir contaminant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBrazil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ491463\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203135\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. fijiensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 313.89\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFiji\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ491680\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ491688\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 100927\u003csup\u003eEN\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCellophane\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSouth Pacific Islands\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFrisvad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 100558\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePeanut\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIndonesia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMG517645\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175478\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFrisvad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 816.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eArachis hypogea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBrazil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMG517628\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMG518011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFrisvad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBD21\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835646\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839564\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBD22\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835647\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839565\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT26\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835651\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839569\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT27\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835652\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839570\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT28\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835653\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839571\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eJN29\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835654\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839572\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eJN210\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835655\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839573\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ23\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835648\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839566\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eSB24\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835649\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839567\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eSB25\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835650\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839568\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. foetidus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 564.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629329\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eGU296697\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. foetidus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNRRL 4750\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAlcoholic beverage\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ491683\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ491690\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. kambarensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 542.69\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ491468\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203136\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. neoniger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 115656\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMangrove water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVenezuela\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ491682\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ491691\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. neoniger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNRRL 62634\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMangrove water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVenezuela\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eKC796401\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKC796361\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 42064\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNot stated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 55765\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEF661186\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 101705\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCarpet dust\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCanada\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629343\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820005\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBP39\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835665\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839599\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eCR31\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835657\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839591\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ32\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835658\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839592\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ33\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835659\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839593\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTK34\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Selangor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835660\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839594\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTK35\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Selangor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835661\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839595\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTI36\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835662\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839596\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eRU37\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835663\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839597\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eRU38\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835664\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839598\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. oryzae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 100925\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eArachis hypogaea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eArgentina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eKJ175432\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203138\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePildain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. oryzae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNRRL 447\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eEurotium oryzae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNR_135395\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF661483\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePeterson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. oryzae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATHUM 4958\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIndoor air\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGreece\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU982022\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU982081\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. parasiticus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 100926\u003csup\u003eEN\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eArachis hypogaea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eArgentina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eKJ175437\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePildain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. parasiticus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATHUM 5037\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIndoor air\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGreece\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU982020\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU982079\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. parasiticus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eATHUM 5038\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIndoor air\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGreece\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU982021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEU982080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. pseudotamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 766.97\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH862673\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203125\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 104.13\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eActivated carbon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUnknown origin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203123\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFrisvad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 117626\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eArachis hypogaea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eArgentina\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNR_135325\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePildain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 133375\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIndoor air\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGermany\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH279382\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMH279885\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFrisvad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBD11\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835674\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839574\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT16\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835679\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839579\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eDT17\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835680\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839580\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ12\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835675\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839575\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ13\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835676\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839576\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ14\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835677\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839577\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ15\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835678\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839578\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTA18\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835681\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839581\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTA19\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835682\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839582\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. terricola\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 620.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSoil\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eUSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ531244\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ491476\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVarga et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. thomii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 120.51\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCulture contaminant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNot stated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH856773\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203133\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePildain et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 11732\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eKuro-koji\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGQ379912\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820012\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 42565\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNR_131293\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820009\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSamson et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2004\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 134.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUnknown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNot stated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNoonim et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBP47\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835690\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839589\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eBP48\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835691\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839590\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eCR41\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835684\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839583\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eJN44\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835687\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839586\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eJN45\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835688\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839587\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eJN46\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835689\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839588\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003ePJ42\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835685\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839584\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eTK43\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eTheobroma cacao\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMalaysia: Selangor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePQ835686\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePQ839585\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePresent study\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eA. sydowii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCBS 593.65\u003csup\u003eET\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDairy product\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGreece\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEU982025\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF652274\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKrimitzas et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003e\u003csup\u003eEN\u003c/sup\u003eEx-neotype culture, \u003csup\u003eET\u003c/sup\u003eEx-type culture\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eMycotoxin screening using coconut cream agar (CCA)\u003c/h3\u003e\n\u003cp\u003eAll fungal isolates obtained were screened for their potential to produce AF and OTA using coconut cream agar (CCA) (Maman et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Seven-day-old pure cultures grown on PDA were subcultured onto CCA plates. To prevent conidial dispersion, a drop of sterile paraffin oil was applied to the center of each plate. The inoculated plates were incubated at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C for 10 days. A CCA plate without fungal inoculation served as a negative control, while a plate inoculated with a non-mycotoxigenic fungal species (\u003cem\u003eColletotrichum\u003c/em\u003e sp.) was used as a positive control to validate the assay. After the incubation period, the reverse side of each CCA plate was examined under ultraviolet (UV) light at a wavelength of 365 nm. The presence fluorescence was used as an indicator of mycotoxin production.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eFungal isolation and enumeration\u003c/h2\u003e \u003cp\u003eA total of 43 \u003cem\u003eAspergillus\u003c/em\u003e isolates were recovered from dried cocoa bean samples collected from 12 farms across Peninsular Malaysia. To quantify fungal contamination, serial dilutions were performed on each sample, followed by plating on PDA to determine colony-forming units per gram (CFU/g). The fungal counts are summarized in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. Most samples exhibited fungal loads below the maximum permissible limit of 1 \u0026times; 10⁴ CFU/g, as stipulated by food hygiene standards (Good Manufacturing Practice \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). However, the sample from Pokok Jenerih, Penang, recorded a higher fungal count of 2.9 \u0026times; 10⁵ CFU/g, exceeding the recommended threshold and indicating potential post-harvest contamination issues.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cem\u003eAspergillus\u003c/em\u003e species isolated from dried cocoa beans collected across various locations in Peninsular Malaysia. The table includes fungal enumeration (CFU/g), sequence similarity to reference isolates in GenBank, and fluorescence intensity on coconut cream agar (CCA)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSampling location\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003csup\u003ea\u003c/sup\u003eFungal enumeration (CFU/g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eFungal species\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eFungal code\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eSequence similarity with GenBank\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003csup\u003eb\u003c/sup\u003eFluorescence on CCA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eITS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003etub2\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eBagan Datuk, Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e1.3 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBD21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBD22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBD11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eChemor, Perak\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1.1 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCR31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCR41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"7\" rowspan=\"8\"\u003e \u003cp\u003ePokok Jenerih, Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"7\" rowspan=\"8\"\u003e \u003cp\u003e2.9 x 10⁵\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePJ42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSungai Bakap, Penang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1.2 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSB24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSB25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTanjung Karang, Selangor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e1.8 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTK34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTK35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTK43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eParit Tujuh, Selangor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e2.2 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePT51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePT52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eTanjung Ipoh, Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e1.4 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTI53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTI54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTI55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTI36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eRembau, Negeri Sembilan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e2.4 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRU56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRU37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRU38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003eDurian Tunggal, Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003e1.8 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. aculeatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629320 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629271(99%)\u003c/p\u003e \u003cp\u003eAY585538 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDT17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"4\" rowspan=\"5\"\u003e \u003cp\u003eJasin, Melaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"4\" rowspan=\"5\"\u003e \u003cp\u003e2.0 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. flavus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJN29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJN210\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH864264 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eKJ175479 (99%)\u003c/p\u003e \u003cp\u003eKJ175478 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJN44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJN45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eJN46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eBatu Pahat, Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e1.8 x 10\u003csup\u003e3\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. niger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBP39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAY585553 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAY820002 (99%)\u003c/p\u003e \u003cp\u003eAY820001 (98%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e++\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. tubingensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBP47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBP48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFJ629354 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFJ629305 (98%)\u003c/p\u003e \u003cp\u003eAY820009 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTangkak, Johor\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e1.4 x 10\u0026sup3;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cem\u003eA. tamarii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTA18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTA19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMH854614 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eEF203122 (100%)\u003c/p\u003e \u003cp\u003eEF203123 (99%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eMorphological and molecular identification\u003c/h3\u003e\n\u003cp\u003eInitial identification based on colony morphology grouped the 43 \u003cem\u003eAspergillus\u003c/em\u003e isolates into five distinct species. Molecular confirmation was performed through DNA sequencing of the ITS region and the \u003cem\u003etub2\u003c/em\u003e gene. BLAST analysis against the GenBank database confirmed all isolates with 98\u0026ndash;100% sequence similarity to known \u003cem\u003eAspergillus\u003c/em\u003e species. The identified species included \u003cem\u003eA. aculeatus\u003c/em\u003e, \u003cem\u003eA. flavus\u003c/em\u003e, \u003cem\u003eA. niger\u003c/em\u003e, \u003cem\u003eA. tamarii\u003c/em\u003e, and \u003cem\u003eA. tubingensis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe morphological characteristics of each species were consistent with descriptions by Pitt and Hocking (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) and Samson et al. (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Among the \u003cem\u003eAspergillus\u003c/em\u003e species isolated, \u003cem\u003eA. aculeatus\u003c/em\u003e was represented by seven isolates, accounting for 16% of the total. These isolates were recovered from four different samples, resulting in a prevalence rate of 33.3% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). On both MEA and CYA, the colonies appeared dark brown and granular, with the reverse sides ranging from white to pale yellow. After three days of incubation, colony diameters measured between 3.0 and 3.2 cm on MEA and 3.1 to 3.3 cm on CYA. Microscopic examination revealed subspheroidal to ellipsoidal conidia measuring 1.4\u0026ndash;2.0 \u0026micro;m in diameter. The conidiophores were smooth, hyaline, and measured 85\u0026ndash;95 \u0026micro;m in length (Table\u0026nbsp;3).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAspergillus flavus\u003c/em\u003e exhibited the highest isolation frequency at 23%, with five isolates recovered from five different samples, corresponding to a prevalence rate of 41.7% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). On CYA, colonies were green-yellow, while on MEA, they appeared yellowish-white with a floccose, tufted texture. Colony diameters ranged from 1.6 to 1.8 cm on CYA and 2.2 to 2.4 cm on MEA. Over time, sclerotia developed on the colonies, initially white and gradually turning black. The reverse side of the colonies was cream-colored. Microscopically, the conidia were globose with rough walls, and the conidial heads were typically radiate, measuring 300\u0026ndash;400 \u0026micro;m in diameter. Vesicles were spherical to sub-spherical, and the conidiophores were hyaline with roughened walls (Table\u0026nbsp;3).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAspergillus niger\u003c/em\u003e was the most prevalent species, represented by nine isolates (21%) and detected in six samples, resulting in a prevalence rate of 50% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Colonies on CYA were dark brown, while those on MEA appeared black. The reverse sides of the colonies were cream to dull yellow on both media. The isolates exhibited rapid growth, with colony diameters ranging from 1.9 to 2.1 cm on CYA and 1.9 to 2.3 cm on MEA. Microscopically, the conidia were globose, brownish to black, and verruculose. The conidiophores were long, smooth, aseptate, and terminated in swollen apices forming vesicles. The conidial heads were dark brown to black, spherical to radiate, and measured 300\u0026ndash;500 \u0026micro;m in diameter. Vesicles were spherical and covered with short terminal branches (metulae) (Table\u0026nbsp;3).\u003c/p\u003e \u003cp\u003eSimilarly, \u003cem\u003eA. tamarii\u003c/em\u003e was represented by nine isolates (21%) and was recovered from four samples, indicating a prevalence rate of 33.3% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). On CYA, colonies were powdery olive-green, while on MEA, they appeared yellowish-green. A distinct white margin surrounded the colonies on both media. The reverse sides were green on CYA and brown on MEA. The isolates grew rapidly at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C, with colony diameters ranging from 1.4 to 1.7 cm on CYA and 1.7 to 1.9 cm on MEA. The conidia were globose to subglobose with rough surfaces. Conidial heads were globose to radiate, measuring 300\u0026ndash;500 \u0026micro;m in diameter. The conidiophores were long, unbranched, hyaline, and rough-walled, terminating in broadly clavate vesicles (Table\u0026nbsp;3).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAspergillus tubingensis\u003c/em\u003e was identified in eight isolates (19%) recovered from five samples, corresponding to a prevalence rate of 41.7% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Colonies on CYA were dark brown, while those on MEA appeared black. The reverse sides of the colonies were creamish dull yellow to yellow on both media. The isolates exhibited rapid growth, with colony diameters ranging from 2.4 to 2.6 cm on CYA and 2.7 to 3.5 cm on MEA. Microscopically, the conidia were globose and brownish to black, and the conidiophores were smooth and elongated. The conidial heads were globose to radiate, measuring 300\u0026ndash;500 \u0026micro;m in diameter (Table\u0026nbsp;3).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e \u003cp\u003ePhylogenetic trees constructed individually from ITS and \u003cem\u003etub2\u003c/em\u003e sequences were insufficient to clearly resolve the species-level relationships among the \u003cem\u003eAspergillus\u003c/em\u003e isolates (Supplementary Figs.\u0026nbsp;1 and 2). In both trees, several isolates clustered ambiguously with multiple reference species, indicating limited discriminatory power when using single-locus data.\u003c/p\u003e \u003cp\u003eTo overcome this limitation, a concatenated dataset combining ITS and \u003cem\u003etub2\u003c/em\u003e sequences was used to generate a ML phylogenetic tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The resulting tree provided significantly improved resolution, with all isolates forming well-supported clades (95\u0026ndash;99%) corresponding to their respective \u003cem\u003eAspergillus\u003c/em\u003e species. Bootstrap values at key nodes were high, indicating strong statistical support for the monophyly of each species group. Distinct clades were clearly delineated and visually highlighted using colored boxes, each representing a different \u003cem\u003eAspergillus\u003c/em\u003e species. This combined-locus approach successfully resolved the taxonomic placement of all isolates, confirming their identities as \u003cem\u003eA. aculeatus\u003c/em\u003e, \u003cem\u003eA. flavus\u003c/em\u003e, \u003cem\u003eA. niger\u003c/em\u003e, \u003cem\u003eA. tamarii\u003c/em\u003e, and \u003cem\u003eA. tubingensis\u003c/em\u003e. These results were fully consistent with BLAST analyses, reinforcing the reliability of the molecular identifications.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eMycotoxigenic potential of\u003c/b\u003e \u003cb\u003eAspergillus\u003c/b\u003e \u003cb\u003eisolates\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe mycotoxigenic potential of \u003cem\u003eAspergillus\u003c/em\u003e isolates was assessed using CCA, a qualitative medium that detect the presence of fluorescent secondary metabolites. All 43 isolates tested exhibited fluorescence, with varying levels of intensity. Low fluorescence intensity (\u0026lsquo;+\u0026rsquo;) was observed in all isolates of \u003cem\u003eA. flavus\u003c/em\u003e, \u003cem\u003eA. tamarii\u003c/em\u003e, \u003cem\u003eA. tubingensis\u003c/em\u003e, and \u003cem\u003eA. aculeatus\u003c/em\u003e. In contrast, moderate fluorescence intensity (\u0026lsquo;++\u0026rsquo;) was observed exclusively in \u003cem\u003eA. niger\u003c/em\u003e isolates. A total of nine \u003cem\u003eA. niger\u003c/em\u003e isolates from Chemor, Pokok Jenerih, Tanjung Karang, Tanjung Ipoh, Rembau, and Batu Pahat displayed this level of fluorescence. No isolates showed an absence of fluorescence or high fluorescence intensity (\u0026lsquo;+++\u0026rsquo;) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe recovery of 43 \u003cem\u003eAspergillus\u003c/em\u003e isolates from dried cocoa beans across 12 farms in Peninsular Malaysia underscores the ubiquitous presence of this fungal genus in post-harvest cocoa environments. Fungal load variation among samples can be attributed to differences in farm-level practices, including harvesting, fermentation, drying, and climatic factors. Notably, while most cocoa samples showed fungal counts below the generally accepted threshold of 1 \u0026times; 10⁴ CFU/g, a significantly elevated count of 2.9 \u0026times; 10⁵ CFU/g was recorded at Pokok Jenerih. This localized anomaly likely results from suboptimal drying conditions, a factor known to promote fungal proliferation (Mounjouenpou et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; S\u0026aacute;nchez-Herv\u0026aacute;s et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Additionally, biological vectors such as insects, birds, and rodents may introduce spores during the sun-drying phase, further exacerbating contamination risks (Delgado-Ospina et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). To mitigate such risks, strict adherence to the Codex Alimentarius Commission\u0026rsquo;s Code of Practice (2013) is essential. The guidelines recommend spreading cocoa beans in layers not exceeding 6 cm, ensuring uniform drying to a final moisture content of 6\u0026ndash;8%, and avoiding re-wetting by covering beans during inclement weather or nighttime (Copetti et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These post-harvest handling protocols are vital for minimizing fungal growth and reducing the risk of subsequent mycotoxin contamination.\u003c/p\u003e \u003cp\u003eMorphological and molecular analyses identified five \u003cem\u003eAspergillus\u003c/em\u003e species in dried cocoa beans namely \u003cem\u003eA. aculeatus\u003c/em\u003e, \u003cem\u003eA. flavus\u003c/em\u003e, \u003cem\u003eA. niger\u003c/em\u003e, \u003cem\u003eA. tamarii\u003c/em\u003e, and \u003cem\u003eA. tubingensis\u003c/em\u003e. The dual use of ITS and \u003cem\u003etub2\u003c/em\u003e region/gene sequences provided robust resolution at the species level, with BLAST and phylogenetic analysis showing high concordance. The predominance of \u003cem\u003eA. flavus\u003c/em\u003e (23%), \u003cem\u003eA. niger\u003c/em\u003e (21%), and \u003cem\u003eA. tamarii\u003c/em\u003e (21%) is particularly significant, as these species are frequently implicated in food spoilage and mycotoxin contamination. These findings align with previous studies reporting the presence of \u003cem\u003eAspergillus\u003c/em\u003e sections \u003cem\u003eFlavi\u003c/em\u003e and \u003cem\u003eNigri\u003c/em\u003e in cocoa (Guehi et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; S\u0026aacute;nchez-Herv\u0026aacute;s et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Nugroho et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The widespread presence of these species suggests they are well-adapted to the cocoa bean microenvironment, likely due to their thermotolerance and ability to survive in low-moisture substrates. Morphological characteristics were consistent with established taxonomic keys (Pitt and Hocking \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Samson et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), further validating the identifications.\u003c/p\u003e \u003cp\u003eAmong the isolates, \u003cem\u003eA. flavus\u003c/em\u003e was the most frequently recovered species, with 10 isolates identified. This species is a known producer of AFs, particularly aflatoxin B₁ (AFB\u003csub\u003e1\u003c/sub\u003e), one of the most potent naturally occurring carcinogens (Klich et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). However, accurate identification can be challenging due to its morphological and genetic similarity to \u003cem\u003eA. oryzae\u003c/em\u003e. Despite these challenges, the concatenated ITS\u0026ndash;\u003cem\u003etub2\u003c/em\u003e phylogenetic tree clearly resolved \u003cem\u003eA. flavus\u003c/em\u003e and \u003cem\u003eA. oryzae\u003c/em\u003e into distinct clades, confirming the accuracy of species-level discrimination in this study. This distinction is critical, given that \u003cem\u003eA. flavus\u003c/em\u003e is toxigenic, whereas \u003cem\u003eA. oryzae\u003c/em\u003e is generally non-aflatoxigenic (Wei and Jong \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Nakamura et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAspergillus tamarii\u003c/em\u003e, another member of section \u003cem\u003eFlavi\u003c/em\u003e, was also frequently isolated. Though generally considered weakly or non-aflatoxigenic, some isolates have been shown to produce aflatoxins under conducive conditions. Klich et al. (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) reported that \u003cem\u003eA. tamarii\u003c/em\u003e possesses genes homologous to the AF biosynthesis cluster and can produce both AFs and sterigmatocystin under specific environmental conditions. Thus, while not typically a major aflatoxin contributor, its toxigenic potential should not be entirely dismissed in tropical post-harvest environments.\u003c/p\u003e \u003cp\u003eSpecies in section \u003cem\u003eNigri\u003c/em\u003e, including \u003cem\u003eA. niger\u003c/em\u003e, \u003cem\u003eA. tubingensis\u003c/em\u003e, and \u003cem\u003eA. aculeatus\u003c/em\u003e, share overlapping morphological features such as dark conidia and similar colony textures, complicating species identification using phenotypic traits alone. These black Aspergilli are commonly found in food and agricultural products and are recognized both for their industrial applications and for their potential to produce ochratoxins (Nielsen et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDue to the limited resolution of ITS sequences alone, this study adopted a multi-locus approach combining ITS and \u003cem\u003etub2\u003c/em\u003e markers. This method significantly improved species-level resolution, as previously demonstrated by Zulkifli and Zakaria (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Yodsing et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Phylogenetic analysis revealed distinct clades for each species with strong bootstrap support (96\u0026ndash;99%), confirming the efficacy of this approach in resolving cryptic species within section \u003cem\u003eNigri\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe use of CCA medium provided preliminary insights into the mycotoxin-producing potential of the recovered isolates. Fluorescent metabolites were detected in all \u003cem\u003eAspergillus\u003c/em\u003e isolates, suggesting widespread biosynthetic capacity for mycotoxin production. The fluorescence observed under UV light, indicative of metabolic activity, varied among species. Notably, moderate fluorescence intensity was recorded only in \u003cem\u003eA. niger\u003c/em\u003e isolates, suggesting potential production of OTA or fumonisins (FBs), known secondary metabolites of this species.\u003c/p\u003e \u003cp\u003eInterestingly, \u003cem\u003eA. flavus\u003c/em\u003e isolates exhibited only low fluorescence. This unexpected result may reflect inter-isolate variability, environmental influences, or limitations of the CCA assay\u0026rsquo;s sensitivity to AFs. CCA\u0026rsquo;s high lipid content, particularly its medium-chain fatty acids like lauric acid, provides a conducive substrate for toxin biosynthesis (Dyer and McCammon \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). Nevertheless, as the assay is qualitative, confirmatory analysis using advanced techniques such as high-performance liquid chromatography (HPLC) or liquid chromatography\u0026ndash;mass spectrometry (LC-MS/MS) is strongly recommended to validate and quantify mycotoxin production (Mohamed et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis study confirms the diversity and prevalence of \u003cem\u003eAspergillus\u003c/em\u003e species in dried cocoa beans in Peninsular Malaysia, with species such as \u003cem\u003eA. flavus\u003c/em\u003e, \u003cem\u003eA. niger\u003c/em\u003e, and \u003cem\u003eA. tamarii\u003c/em\u003e dominating the post-harvest fungal community. The combination of morphological and molecular tools enabled accurate species-level identification, while preliminary assessments of toxigenic potential highlighted the need for further mycotoxin quantification. The findings reinforce the importance of good post-harvest practices and molecular surveillance to safeguard cocoa quality and food safety.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eConflict of interest:\u003c/h2\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eThis research was supported by Universiti Sains Malaysia through the Bridging Grant (Project No: R501-LR-RND003-0000000743-0000) and the Fundamental Research Grant Scheme (FRGS/1/2019/WAB01/USM/02/1) provided by the Ministry of Higher Education, Malaysia. The first author, Nuraina Syazana, also received financial support through the Postgraduate Research Attachment (PGRA) from the Institute of Postgraduate Studies (IPS), Universiti Sains Malaysia.\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eConceptualization, SN-S and MHM; Experimentation, data collection and formal analysis, SN-S, MZZ, and NSS; Writing\u0026mdash;original draft preparation, SN-S; Writing\u0026mdash;review and editing, SN-S, MZZ, NSS, Y-HL, and MHM; Supervision and resources, MHM and Y-HL. All authors have reviewed the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eThe authors extend their sincere gratitude to the Malaysian Cocoa Board (MCB) for granting permission to collect cocoa bean samples. Special appreciation is also given to the MCB staff for their invaluable assistance and support during fieldwork activities.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eData is provided within the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAgus BAP, Mohamad NN, Hussain N (2018) Composition of unfermented, unroasted, roasted cocoa beans and cocoa shells from Peninsular Malaysia. J Food Meas Charact 12:2581\u0026ndash;2589\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCodex Alimentarius Commission (2013) Proposed draft code of practice for the prevention and reduction of ochratoxin A contamination in cocoa. Joint FAO/WHO Food Standards Programme, FAO, Rome\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCopetti MV, Iamanaka BT, Pitt JI, Taniwaki MH (2014) Fungi and mycotoxins in cocoa: from farm to chocolate. 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PCR protocols: a guide to methods and applications. 18(1):315\u0026ndash;322\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYodsing N, Lekphrom R, Sangsopha W, Aimi T, Boonlue S (2018) Secondary metabolites and their biological activity from \u003cem\u003eAspergillus aculeatus\u003c/em\u003e KKU-CT2. Curr Microbiol 75:513\u0026ndash;518\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZulkifli NA, Zakaria L (2017) Morphological and molecular diversity of \u003cem\u003eAspergillus\u003c/em\u003e from corn grain used as livestock feed. HAYATI J Biosci 24(1):26\u0026ndash;34\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables 1 to 3 are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Theobroma cacao, toxigenic fungi, mycotoxins, post-harvest handling, phylogenetic analysis, food safety","lastPublishedDoi":"10.21203/rs.3.rs-6760467/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6760467/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eCocoa (\u003cem\u003eTheobroma cacao\u003c/em\u003e) is a key agricultural commodity that is highly susceptible to fungal and mycotoxin contamination, particularly during post-harvest processes such as fermentation, drying, and storage. This study aimed to characterize \u003cem\u003eAspergillus\u003c/em\u003e species associated with dried cocoa beans from 12 farms across six states in Peninsular Malaysia using morphological, molecular, phylogenetic, and toxigenicity analyses. Fungal isolation and enumeration were performed, followed by identification based on morphological characteristics, DNA sequencing, and phylogenetic analysis targeting the internal transcribed spacer (ITS) region and β-tubulin (\u003cem\u003etub2\u003c/em\u003e) gene. Mycotoxin-producing potential was assessed using coconut cream agar (CCA), where fluorescence under ultraviolet (UV) light indicated the presence of secondary metabolites. A total of 43 \u003cem\u003eAspergillus\u003c/em\u003e isolates were identified as \u003cem\u003eA. flavus\u003c/em\u003e (10 isolates), \u003cem\u003eA. niger\u003c/em\u003e (nine isolates), \u003cem\u003eA. tamarii\u003c/em\u003e (nine isolates), \u003cem\u003eA. tubingensis\u003c/em\u003e (eight isolates), and \u003cem\u003eA. aculeatus\u003c/em\u003e (seven isolates). One sample exceeded the hygienic threshold, with a fungal load of 2.9 \u0026times; 10⁵ CFU/g, surpassing the permissible limit of 1 \u0026times; 10⁴ CFU/g. All isolates exhibited fluorescence, indicating potential mycotoxin production. Moderate fluorescence intensity was observed exclusively in \u003cem\u003eA. niger\u003c/em\u003e, while all other species showed low fluorescence levels. These findings underscore the importance of improved post-harvest handling practices to mitigate fungal contamination and mycotoxin risks. The results offer valuable insights for enhancing food safety, supporting cocoa trade, and informing agricultural policy in cocoa-producing regions.\u003c/p\u003e","manuscriptTitle":"Characterization of Aspergillus species associated with dried cocoa beans in Peninsular Malaysia","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-06 13:54:15","doi":"10.21203/rs.3.rs-6760467/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"7d722f09-b419-4684-974b-516b96cf2fc3","owner":[],"postedDate":"June 6th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-06-27T19:53:20+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-06 13:54:15","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6760467","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6760467","identity":"rs-6760467","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}