Characterization of a <i>Paracoccidioides</i> spp. strain from southeastern Brazil genotyped as <i>Paracoccidioides restrepiensis</i> (PS3) and review of this phylogenetic species

Cocio, Tiago Alexandre; Nascimento, Erika; Kress, Marcia R. V. Z.; Bagagli, Eduardo; Martinez, Roberto

doi:10.1590/1678-4685-GMB-2019-0201

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Genomics and Bioinformatics • Genet. Mol. Biol. 43 (02) • 2020 • https://doi.org/10.1590/1678-4685-GMB-2019-0201 copy

Characterization of a Paracoccidioides spp. strain from southeastern Brazil genotyped as Paracoccidioides restrepiensis (PS3) and review of this phylogenetic species

Authorship SCIMAGO INSTITUTIONS RANKINGS

Abstract

Phylogenetic species of Paracoccidioides brasiliensis complex (S1a and S1b, PS2, PS3, and PS4) and Paracoccidioides lutzii are agents of paracoccidioidomycosis, an endemic fungal disease in Latin America. P. restrepiensis (PS3 genotype) was classified as monophyletic and geographically restricted to Colombia and neighboring territories. BAT (or Pb-327B) was isolated from a patient living in the southeast region of Brazil but with genotype similar to Colombian Paracoccidioides spp. strains. This study aimed to define the phylogenetic species of BAT isolate by using additional genotyping methods, as well as reviewing the epidemiological and clinical studies related to P. restrepiensis isolates. Genomic DNA of BAT isolate and reference strains of P. brasiliensis sensu stricto (S1b), P. americana (PS2), P. restrepiensis (PS3), and P. lutzii were analyzed by conventional polymerase chain reaction (PCR) of partial gp43 exon 2 loci, by PCR-RFLP technique of tub1 gene, and by sequencing of the whole gp43 exon 2 loci. Here, we show that BAT isolate belongs to P. restrepiensis species, which is an unusual identification in southeastern Brazil, where P. brasiliensis sensu stricto is the prevalent genotype. This identification has relevance for geographical distribution and propagation of the genus Paracoccidioides in South America.

Keywords:
Paracoccidioides restrepiensis; Paracoccidioides brasiliensis PS3; phylogenetic species; evolution; paracoccidioidomycosis epidemiology

Strain		Origin	Source	Identification method	Study Type	Reference
Original ID	Other ID
C1	P149	Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C2	P159	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C3	P163	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C4	ATCC 60855	Antioquia, Colombia	Chronic /PCM	MLST	Phenotypic	(*Bustamante-Simon et al., 1985*Bustamante-Simon B, McEwen JG, Tabares AM, Arango M and Restrepo-Moreno A (1985) Characteristics of the conidia produced by the mycelial form of Paracoccidioides brasiliensis Sabouraudia 23:407-414.)
					Phenotypic	(*Villar et al., 1988*Villar LA, Salazar ME and Restrepo A (1988) Morphological study of a variant of Paracoccidioides brasiliensis that exists in the yeast form at room temperature. J Med Vet Myco 26:269-276.)
					Virulence	(*Gomez et al., 2001*Gomez BL, Nosanchuk JD, Diez S, Youngchim S, Aisen P, Cano LE, Restrepo A, Casadevall A and Hamilton AJ (2001) Detection of melanin-like pigments in the dimorphic fungal pathogen Paracoccidioides brasiliensisin vitro and during infection. Infect Immun 69:5760-5767.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C5	P141	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C6	P196 / Higuita	Antioquia, Colombia	Chronic /PCM	MLST	Molecular Identification	(*Corredor et al., 2005*Corredor GG, Peralta LA, Castano JH, Zuluaga JS, Henao B, Arango M, Tabares AM, Matute DR, McEwen JG and Restrepo A (2005) The naked-tailed armadillo Cabassous centralis (Miller 1899): a new host to Paracoccidioides brasiliensis Molecular identification of the isolate. Med Mycol 43:275-280.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C7	P204	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C8	P292	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C9	P68	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C10	P72	Cordoba, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C11	P46	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C12	P161	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C13	76533	Antioquia, Colombia	Chronic /PCM	MLST	Susceptibility	(*Hoyos et al., 1984*Hoyos GL, McEwen JG, Brummer E, Castaneda E, Restrepo A and Stevens DA (1984) Chronic murine paracoccidioidomycosis: effect of ketoconazole on clearance of Paracoccidioides brasiliensis and immune response. Sabouraudia 22:419-426.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C14	H31	Boyaca, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C15	H45	Cundinamarca, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C16	H47	Arauca, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C17	P206	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C18	P151	Antioquia, Colombia	Chronic /PCM	MLST	Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C19	CIB44197	Caldas, Colombia	Armadillo	MLST	Molecular Identification	(*Corredor et al., 2005*Corredor GG, Peralta LA, Castano JH, Zuluaga JS, Henao B, Arango M, Tabares AM, Matute DR, McEwen JG and Restrepo A (2005) The naked-tailed armadillo Cabassous centralis (Miller 1899): a new host to Paracoccidioides brasiliensis Molecular identification of the isolate. Med Mycol 43:275-280.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C20	Pb73/ ATCC 32071	Antioquia, Colombia	Unknown	MLST	Serological tests	(Restrepo-Moreno 1967Restrepo-Moreno A and Schneidau JD (1967) Nature of the skin-reactive principle in culture filtrates prepared from Paracoccidioides brasiliensis J Bacteriol 93:1741-1748.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
C21	CIB40392	Caldas, Colombia	Armadillo	MLST	Epidemiological	(*Corredor et al., 1999*Corredor GG, Castano JH, Peralta LA, Diez S, Arango M, McEwen J and Restrepo A (1999) Isolation of Paracoccidioides brasiliensis from the nine-banded armadillo Dasypus novemcinctus in an endemic area for paracoccidioidomycosis in Colombia. Rev Iberoam Micol 16:216-220.)
					Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
T2	EPM54	Caracas, Venezuela	Soil	MLST, PCR – RFLP	Phylogenetic	(*Roberto et al., 2016*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
5598	EPM62	Caracas, Venezuela	Acute/PCM	MLST, PCR – RFLP	Phylogenetic	(*Roberto et al., 2016*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
JDA – 80	EPM77	Medellin, Colombia	Unknown	MLST, PCR – RFLP	Phylogenetic	(*Roberto et al., 2016*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
MSCol	EPM81	Medellin, Colombia	Chronic /PCM	MLST, PCR – RFLP	Phylogenetic	(*Roberto et al., 2016*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
I9	EPM83	Bogotá, Colombia	Chronic /PCM	MLST, PCR – RFLP	Phylogenetic	(*Theodoro et al., 2008*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
					Serological	(*Machado et al., 2013*Machado GC, Moris DV, Arantes TD, Silva LR, Theodoro RC, Mendes RP, Vicentini AP and Bagagli E (2013) Cryptic species of Paracoccidioides brasiliensis: impact on paracoccidioidomycosis immunodiagnosis. Mem Inst Oswaldo Cruz 108:637-643.)
					Proteome	(*Pigosso et al., 2013*Pigosso LL, Parente AF, Coelho AS, Silva LP, Borges CL, Bailao AM and Soares CM (2013) Comparative proteomics in the genus Paracoccidioides Fungal Genet Biol 60:87-100.)
					Secretome	(*de Oliveira et al., 2018*de Oliveira AR, Oliveira LN, Chaves EGA, Weber SS, Bailao AM, Parente-Rocha JA, Baeza LC, de Almeida Soares CM and Borges CL (2018) Characterization of extracellular proteins in members of the Paracoccidioides complex. Fungal Biol 122:738-751.)
*B18	Pb339 or	Brazil	Unknown	MLST, PCR - RFLP	Serological tests	(Restrepo-Moreno 1967Restrepo-Moreno A and Schneidau JD (1967) Nature of the skin-reactive principle in culture filtrates prepared from Paracoccidioides brasiliensis J Bacteriol 93:1741-1748.)
	ATCC 32069				Virulence	(Gomez et al.
	EPM01 or B339				Serological	(*Camargo et al., 2003*Camargo ZP, Berzaghi R, Amaral CC and Silva SH (2003) Simplified method for producing Paracoccidioides brasiliensis exoantigens for use in immunodiffusion tests. Med Mycol 41:539-542.)
	or ATCC 200273				Phylogenetic	(*Matute et al., 2006*Matute DR, McEwen JG, Puccia R, Montes BA, San-Blas G, Bagagli E, Rauscher JT, Restrepo A, Morais F, Nino-Vega G et al. (2006) Cryptic speciation and recombination in the fungus Paracoccidioides brasiliensis as revealed by gene genealogies. Mol Biol Evol 23:65-73.)
					Mitochondrial DNA	(*Salgado – Salazar et al., 2010*Salgado-Salazar C, Jones LR, Restrepo A and McEwen JG (2010) The human fungal pathogen Paracoccidioides brasiliensis (Onygenales: Ajellomycetaceae) is a complex of two species: phylogenetic evidence from five mitochondrial markers. Cladistics 26:613-624.)
					Phylogenetic	(*Roberto et al., 2016*Roberto TN, Rodrigues AM, Hahn RC and de Camargo ZP (2016) Identifying Paracoccidioides phylogenetic species by PCR-RFLP of the alpha-tubulin gene. Med Mycol 54:240-247.)
					Phylogenetic	(*Muñoz et al., 2016*Muñoz JF, Farrer RA, Desjardins CA, Gallo JE, Sykes S, Sakthikumar S, Misas E, Whiston EA, Bagagli E, Soares CM et al. (2016) Genome diversity, recombination, and virulence across the major lineages of Paracoccidioides mSphere 1:e00213-00216.)
PbBac	-	Colombia	PCM	Genomic sequencing	Phylogenetic	(*Muñoz et al., 2016*Muñoz JF, Farrer RA, Desjardins CA, Gallo JE, Sykes S, Sakthikumar S, Misas E, Whiston EA, Bagagli E, Soares CM et al. (2016) Genome diversity, recombination, and virulence across the major lineages of Paracoccidioides mSphere 1:e00213-00216.)
PbCnh	-	Colombia	Chronic /PCM	Genomic sequencing	Phylogenetic	(*Muñoz et al., 2016*Muñoz JF, Farrer RA, Desjardins CA, Gallo JE, Sykes S, Sakthikumar S, Misas E, Whiston EA, Bagagli E, Soares CM et al. (2016) Genome diversity, recombination, and virulence across the major lineages of Paracoccidioides mSphere 1:e00213-00216.)
PbJam	-	Colombia	Chronic /PCM	Genomic sequencing	Phylogenetic	(*Muñoz et al., 2016*Muñoz JF, Farrer RA, Desjardins CA, Gallo JE, Sykes S, Sakthikumar S, Misas E, Whiston EA, Bagagli E, Soares CM et al. (2016) Genome diversity, recombination, and virulence across the major lineages of Paracoccidioides mSphere 1:e00213-00216.)
BAT	Pb327 - B	Ribeirão Preto – São Paulo, Brazil	Subacute/PCM	MLST, PCR – RFLP	Serological	(*Silva – Vergara et al., 1998*Silva-Vergara ML, Martinez R, Chadu A, Madeira M, Freitas-Silva G and Leite Maffei CM (1998) Isolation of a Paracoccidioides brasiliensis strain from the soil of a coffee plantation in Ibia, State of Minas Gerais, Brazil. Med Mycol 36:37-42.)
					Mycological	(*Lacaz Cda, S et al., 1999*Lacaz CS, Vidal MS, Heins-Vaccari EM, de Melo NT, Del Negro GM, Arriagada GL and Freitas RS (1999) Paracoccidioides brasiliensis A mycologic and immunochemical study of two strains. Rev Inst Med Trop Sao Paulo 41:79-86.)
					Phylogenetic	(*Calcagno et al., 1998*Calcagno AM, Nino-Vega G, San-Blas F and San-Blas G (1998) Geographic discrimination of Paracoccidioides brasiliensis strains by randomly amplified polymorphic DNA analysis. J Clin Microbiol 36:1733-1736.)
					Phylogenetic	(*Niño – Vega et al., 2000*Nino-Vega GA, Calcagno AM, San-Blas G, San-Blas F, Gooday GW and Gow NA (2000) RFLP analysis reveals marked geographical isolation between strains of Paracoccidioides brasiliensis Med Mycol 38:437-441.)
					Serological	(*Panuto – Castelo et al., 2003*Panunto-Castelo A, Freitas-da-Silva G, Bragheto IC, Martinez R and Roque-Barreira MC (2003) Paracoccidioides brasiliensis exoantigens: recognition by IgG from patients with different clinical forms of paracoccidioidomycosis. Microbes Infect 5:1205-1211.)
					Immunologic	(*Coltri et al., 2006*Coltri KC, Casabona-Fortunato AS, Gennari-Cardoso ML, Pinzan CF, Ruas LP, Mariano VS, Martinez R, Rosa JC, Panunto-Castelo A and Roque-Barreira MC (2006) Paracoccin, a GlcNAc-binding lectin from Paracoccidioides brasiliensis binds to laminin and induces TNF-alpha production by macrophages. Microbes Infect 8:704-713.)
					Molecular Identification	This Study

P. restrepiensis isolate	P. brasiliensis complex or P. lutzii isolate	Evaluated trait	Observed differences	Study
C4, BAC, CNH	Pb18, T15LN1, T4B14 - P. brasiliensis sensu stricto (S1)	Yeast cell size	P. restrepiensis (PS3) shows larger yeast area than P. brasiliensis sensu stricto (S1) and P. americana (PS2) isolates but similar to P. venezuelensis (PS4)	(*Turissini et al., 2017*Turissini DA, Gomez OM, Teixeira MM, McEwen JG and Matute DR (2017) Species boundaries in the human pathogen Paracoccidioides Fungal Genet Biol 106:9-25.)
	Pb3 and T10B1 - P. americana (PS2)
	Pb300 and Pb305 - P. venezuelensis (PS4)
	Pb66, Pb01, EE - P. lutzii
BAT	Pb18 - P. brasiliensis sensu stricto (S1)	Exoantigen production	Higher number of BAT exoantigens recognized by patient sera	(*Panuto-Castelo et al., 2003*Panunto-Castelo A, Freitas-da-Silva G, Bragheto IC, Martinez R and Roque-Barreira MC (2003) Paracoccidioides brasiliensis exoantigens: recognition by IgG from patients with different clinical forms of paracoccidioidomycosis. Microbes Infect 5:1205-1211.)
	DGO and C – 9 - Unknown Genotype
	B_339 – P. brasiliensis sensu stricto (S1)
EPM83	Pb265 - P. brasiliensis sensu stricto (S1)	GP43 production and antigenicity	EPM83 showed higher GP43 production and antigenicity in the immunodiffusion test with PCM sera	(*Machado et al., 2013*Machado GC, Moris DV, Arantes TD, Silva LR, Theodoro RC, Mendes RP, Vicentini AP and Bagagli E (2013) Cryptic species of Paracoccidioides brasiliensis: impact on paracoccidioidomycosis immunodiagnosis. Mem Inst Oswaldo Cruz 108:637-643.)
	Pb01, Pb8334, Pb66 - P. lutzii
EPM83	Pb339 – P. brasiliensis sensu stricto (S1)	Proteome	EPM83 presented higher expression of 38 proteins, including HSP and GP43	(*Pigosso et al., 2013*Pigosso LL, Parente AF, Coelho AS, Silva LP, Borges CL, Bailao AM and Soares CM (2013) Comparative proteomics in the genus Paracoccidioides Fungal Genet Biol 60:87-100.)
	Pb2 – P.americana (PS2)
	Pb01 – P. lutzii
EPM83	Pb01 - P. lutzii	Secretome	EPM83 production of 36 specific proteins; 21 proteins shared with Pb01	(*de Oliveira et al., 2018*de Oliveira AR, Oliveira LN, Chaves EGA, Weber SS, Bailao AM, Parente-Rocha JA, Baeza LC, de Almeida Soares CM and Borges CL (2018) Characterization of extracellular proteins in members of the Paracoccidioides complex. Fungal Biol 122:738-751.)

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[1] Send correspondence to: Tiago Alexandre Cocio. Universidade de São Paulo (USP), Faculdade de Medicina de Ribeirão Preto, Departamento de Clínica Médica. Av. Bandeirantes, 3900, Monte Alegre, Ribeirão Preto, São Paulo, Brazil, Postal Code: 14049-900. Phone/Fax: 55-16-3602-2468; E-mail: alexcocio@gmail.com