1. Cátedra de Fitopatología. Fa c u l t a d d e A g ro n o m í a . Universidad de Buenos Aires. Av. San Martin 4453. CP 1417. Buenos Aires, Argentina.

2.CIDEFI – UNLP - CICBA, Facultad de Ciencias Agrarias y Forestales, UNLP, 60 y 119, CP 1900. La Plata, Buenos Aires, Argentina.

*palmucci@agro.uba.ar

Citar este artículo Palmucci, H. E. and S. M. Wolcan. 2020. The Genus Phytophthora (Peronosporales) in Argentina. Bol. Soc. Argent. Bot. 55: 161-193.

DOI: https://doi. org/10.31055/1851.2372.v55. n2.25537

Recibido: 4 Octubre 2019

Aceptado: 20 Mayo 2020

Publicado: 30 Junio 2020

Editora: María Victoria Vignale

ISSN versión impresa 0373-580X ISSN versión on-line 1851-2372

SUMMARY

Background and aims: The genus Phytophthora includes plant pathogens that affect a wide host range and cause severe damage and economic losses. The aim of this study was to achieve a more comprehensive knowledge of Phytophthora in Argentina. To this end, a review was carried out from the first reports in the late nineteenth century until March 2019.

M&M: Information was taken from printed and on-line primary and secondary sources such as Proceedings of National and International Scientific Meetings, Bulletins from National Institutions and Universities, periodical Journals, books and data bases, and then analyzed and categorized.

Results: The revision allowed updating the status of Phytophthora species recorded in the country, considering their geographical distribution, groups of crops affected, host-pathogen relationships, symptoms and nomenclature changes, as well as presenting a quick and comparative access to different subjects related to these pathogens. The results showed that, to date, 20 Phytophthora spp., one species affinis and one taxon affect 223 host-pathogen relationships in Argentina. The diversity of Phytophthora species in the world suggests that a larger number of species, still not cited, could be present in Argentina.

Conclusions: Researchers specialized in the genus Phytophthora, molecular techniques and phylogenetic studies, may allow progressing in the accurate identification of the species and knowledge of their genetic variability.

KEY WORDS

Argentina, host range, Oomycetes, Peronosporomycetes, Phytophthora, Pseudofungi, Stramenopila.

RESUMEN

Introducción: El género Phytophthora incluye patógenos de plantas que afectan un amplio rango de hospedantes, causando severos daños y pérdidas económicas. Con la finalidad de conocer el estado actual de la información disponible sobre este género en Argentina, se llevó a cabo una revisión desde fines del siglo XIX hasta marzo de 2019.

M&M: Se consultaron fuentes primarias y secundarias de información escrita y electrónica: presentaciones en congresos nacionales e internacionales, publicaciones periódicas nacionales e internacionales, boletines de instituciones y universidades, libros y bases de datos. La información obtenida fue analizada y categorizada.

Resultados: La presente revisión permitió actualizar las especies presentes en el país, considerando su distribución geográfica y por grupos de cultivos, relaciones hospedante-patógeno, sintomatología y los cambios en su nomenclatura. Se presenta un rápido y comparativo acceso a los diferentes tópicos relacionados con estas patologías. Se encontraron citadas 20 Phytophthora spp., 1 especie affinis y 1 taxon que afectan a 223 relaciones hospedante-patógeno. La diversidad de Phytophthora spp. citadas en el mundo sugiere que en el país podría haber una mayor cantidad de especies aún no registradas.

Conclusión: Investigadores especializados en el género Phytophthora, técnicas moleculares y estudios filogenéticos permitirán avanzar en la certera identificación de las especies y el conocimiento de su variabilidad genética.

PALABRAS CLAVE

Argentina, Oomycetes, Peronosporomycetes Phytophthora, Pseudofungi, rango de hospedantes, Stramenopila.

161

Bol. Soc. Argent. Bot. 55 (2) 2020

INTRODUCTION

The genus Phytophthora de Bary includes pathogen species that cause diseases in both natural (grasslands or forests) and agricultural ecosystems worldwide, producing severe damage and economic losses. These species can infect a wide variety of ornamental, forest, oleaginous, vegetable and fruit crops, producing symptoms as root, stem and fruit rots, blights, cankers and wilting (Erwin & Ribeiro, 1996). Most Phytophthora species, such as P. nicotianae Breda de Haan, P. cinnamomi R.D. Rands and P. citrophthora (R.E. Sm. & E.H. Sm.) Leonian, affect a wide host range, while others such as P. infestans and P. sojae Kaufm. & Gerd. are host-specific (Erwin

&Ribeiro, 1996; Agrios, 2005). Due to their high virulence and ability to spread disease rapidly, the species of this genus are some of the most important plant pathogens worldwide (Erwin & Ribeiro, 1996).

Regarding its taxonomic location, the genus Phytophthora has been subject of successive changes. At first, it was considered to belong to the

Kingdom Fungi (Whittaker, 1969), whereas, later, due to the existence of numerous morphological and physiological differences with true fungi, it was relocated in the Kingdom Chromista (Cavalier-Smith, 1986) and in the Kingdom Protoctist (Margulis et al., 1989). In this context, these organisms were denominated “pseudofungi” and included into the Subdivision Pseudofungi, Phylum Heterokonta (Kirk et al., 2011). In 2001, Dick placed them within the new Kingdom Stramenopila. This position was supported by characteristics such as the composition of the cellular wall, which is composed of cellulose and beta glucans instead of chitin as in true fungi (Bartnicki-García, 1969); the presence of heterokont spores with two flagella (Desjardins et al., 1969), which allow the zoospores to move in liquid media; the coenocytic mycelia, and different sensitivity to conventional fungicides (Latijnhouwers et al., 2003) among other differences.

Currently, the two classifications that place pseudofungi in the Kingdom Chromista (Class Oomycetes) and the Kingdom Stramenopila (Class Peronosporomycetes) are accepted, although the tendency is to consider them as belonging to the Kingdom Stramenopila (Lamour, 2013).

Based on its morphological and nutritional characteristics, the genus Phytophthora was located in the order Pythiales, family Pythiaceae, along

162

with Pythium Pringsh and related genera for many years (Dick, 2001). Later, molecular studies allowed proposing a close Phytophthora affiliation with the downy mildews in the order Peronosporales and family Peronosporaceae (Hulvey et al., 2010; Beakes et al., 2014; Thines & Choi, 2016; Beakes & Thines, 2017).

Molecular-based phylogenies with up to 15 nuclear and mitochondrial genes have been implemented since 2000 (Cooke et al., 2000; Kroon et al., 2004; Blair et al., 2008; Martin et al., 2014; Yang et al., 2017). These studies have placed Phytophthora species in ten well- defined clades.

Several surveys, reviews and taxonomic studies about the genus Phytophthora, including parasitic and/or saprophytic species obtained from soil or water, have been carried out in different countries, including Argentina (Frezzi, 1950, 1977), China (Ho, 1996), England (Brasier & Strouts, 1976 cited by Lodhi, 2007), New Zealand (Boesewinkel, 1974 cited by Lodhi, 2007), Taiwan (Ann, 1992; Ho et al., 1995) and the USA (Campbell & Hendrix, 1967; Hendrix & Campbell, 1970; Erwin & Ribeiro, 1996). The results of all these studies have greatly contributed to the knowledge of the biodiversity of the genus in different regions.

The most complete review was that published by Erwin & Ribeiro (1996) in the USA. This review provided extensive information about Phytophthora spp. and deepened different aspects about their worldwide distribution, morphology, physiology, taxonomic evolution, pathogenesis, disease control, etc. Later, different surveys were conducted in Sindhum, Pakistan (Lodhi, 2007), Hainan Island, China (Zeng et al., 2009), Spain (Moralejo et al., 2009), the United Kingdom (Denton et al., 2008) and Italy (Cacciola et al., 2008). Other studies were carried out in different regions of the USA by Schwingle et al. (2007), Warfield et al. (2008), Yakabe et al. (2009) and Olson & Benson (2011), in Mexico by Perez Sierra et al. (2012), and in South America only in Brazil by Luz et al. (2001).

The information referred to Phytophthora is scattered in on-line or printed scientific articles published by National Universities, National or International Scientific Journals or Meetings and books. The aim of this study was to collect, analyze, and organize the information about Phytophthora spp. recorded in Argentina and update their taxonomic nomenclature.

MATERIALS AND METHODS

The revision of the Phytophthora spp. recorded in Argentina here performed covered the period from the late nineteenth century to March 2019, and considered only the records of Phytophthora spp. affecting plants or related to plant diseases. Both printed and electronic primary and secondary sources of information were consulted. Primary sources included a) Proceedings of National and International Scientific Meetings; b) National Journals such as Boletín de la Sociedad Argentina de Botánica, Revista de la Facultad de Agronomía de la Universidad de Buenos Aires (FAUBA), Revista de la Facultad de Agronomía de la Universidad de La Plata (UNLP), Revista Argentina de Agronomía, and Horticultura Argentina, among others; and c) International Journals such as Australasian Plant Disease Notes, Journal of Plant Pathology, New DiseaseReports,PlantDisease,Phytopathology,Plant Pathology, Tropical Plant Pathology, and Revista Fitopatología de la Asociación Latinoamericana de Fitopatología (ALF), among others. Secondary sources included a) books (Marchionatto, 1948, 1950; Fernández Valiela, 1978, 1979; Erwin & Ribeiro, 1996; b) databases: AGRIS (International Information System for the Agricultural Science

&Technology) 1991-2001 (http://agris.fao.org/); CABPEST CD 1973-1988; 1989-1999; 2000-2002; CABPEST or (CAB International on SilverPlatter, Search and Retrieval Software) 1990-2013; EBSCO Host Research Databases, 1975-2019 (http://search.ebscohost.com); CAB ABSTRACT, 1990-2013 (www.cabi.org/publishing-products/cab- abstracts); ScienceDirect (http://www.sciencedirect. com); SCOPUS, 1975-2019 (www.scopus.com); Biblioteca Electrónica de Ciencia y Tecnología (www.biblioteca.mincyt.gob.ar) and c) electronic publications: Atlas Fitopatológico de Argentina (Nome, et al. 2019).

The botanical nomenclature of the hosts was based on the International Plant Name Index (www. ipni.org). For the nomenclature of pseudofungi, different databases were consulted: Farr, D.F & Rossman, A.Y. Fungal Databases; U.S. National Fungus Collections; ARS, USDA, (https://nt.ars- grin.gov/fungaldatabases); Phytophthora Database (www.phytophthoradb.org/species.php) and Index Fungorum (www.indexfungorum.org/Names/ Names.asp).

RESULTS

The information obtained as a result of the revision was analyzed and categorized and is presented in three tables and one figure.

The species of Phytophthora so far recorded in Argentina, together with their host range, symptoms caused and geographical distribution are shown in Table 1. This Table also includes references of the first report in each province, which in turn include some other significant data about the pathogens or disease descriptions and references of the first molecular characterization of the species.

According to our search, the first reference of Phytophthora in Argentina was P. infestans on potato, cited by Rieder (1887). The following reference was Girola (1898), who wrote about this pathogen. After that time, Lindquist made new findings and detailed descriptions (Lindquist, 1932, 1935, 1938), whereas Frezzi made several reports and two compilations of Phytophthora in Argentina (Frezzi, 1950, 1977). The first compilation (Frezzi, 1950) included an exhaustive morphological characterization of the 11 species known until that time, with their host range, location, and detailed drawings of their morphology, and served as later reference for foreign papers. The second compilation (Frezzi, 1977) included the new report of P. cryptogea (Frezzi, 1957) and a wider range of host-pathogen relations and their locations.

Most of the taxonomic identification of Phytophthora spp. has been based on their morphological features and colony patterns on different culture media. Some species such P. aff. boehmeriae, P. citricola complex, P medicaginis, P. megasperma and P. syringae were reported only 50 – 70 years ago on a few hosts and were not cited again (Table 1). Later, in the 2000s, molecular diagnostic methods to identify Phytophthora spp. began to be applied. This allowed making taxonomic identifications with greater certainty. The first ITS sequence identification was in

P.austrocedri (Greslebin et al., 2007). Then, molecular techniques allowed identifying P. sojae (Gally et al., 2007), P. palmivora (Lucero et al., 2007), P. cinnamomi (Palmucci et al., 2008), P. nicotianae on different hosts (Lucero et al., 2006; Grijalba et al., 2008; Vetraino et al., 2009; Palmucci et al., 2011a, b, c; 2012, 2013, Palmucci, 2015), P. lacustris (ex P. taxon salixsoil) (Dobra et al.,

	Species	Host	Symptoms	Province
	P. austrocedri	Austrocedrus	Necrosis of roots and	Río Negro
				Chubut
	Greslebin & Hansen	chilensis	stems, tree decline
				Neuquén
	P. aff. boehmeriae	Citrus sinensis	Fruit rot	Corrientes
	(P. boehmeriae	Chamelauciun	Basal rot	Buenos Aires
	Sawada)	uncinatum
		Citrus limon	Brown rot of fruits	Entre Ríos
				Jujuy
		C. maxima=paradisi	Brown rot of fruits	Buenos Aires
		Eucalyptus grandis	Dieback	Corrientes
			Root rot, trunk
		Fatsia japonica	necrosis, death of	Córdoba
			adult plants, basal rot

References1

Greslebin et al. (2007*, 2014);

Greslebin & Hansen (2010); Vélez

et al. (2012, 2014*); La Manna (2013); Taccari et al. (2019)

Frezzi (1941,1942, 1950)

Wolcan & Lori (2001)

Fawcett 1937 In: Frezzi (1950)

Bejarano & Catacata (2009)

Frezzi 1949 In: Frezzi (1950)

Colombo et al. (2011)

Frezzi 1947 In: Frezzi (1950)

	Fragaria ananassa	Root rot, dry rot of
		fruits, crown rot

Corrientes	Colombo (1993); Colombo et al.(2001)
Jujuy	Alcoba et al.(2005)
Tucumán	Baino (2008)
Santa Fe	Mitidieri & Scandiani (1989)

Vigliola et al. (1984); Mitidieri &

Buenos Aires Scandiani (1989); Gonzalez et al.(2003); Irribarren et al.(2012e)

P. cactorum (Lebert
& Cohn) Schroet.
Malus domestica	Root and crown rot
Olea europaea	Root rot

Buenos Aires

Córdoba

Río Negro

Mendoza

Neuquén

Catamarca

La Rioja

Mendoza

San Juan

Lindquist (1938)

Frezzi 1942 In: Frezzi (1950)

Frezzi 1945 In: Frezzi (1950); Bergna

et al.(1979); Rossini et al.(1997)

Pontis & Feldman (1960)

Bergna (1963); Rossini et al.(1997)

Velasco et al. (1997)

	Persea americana	Root rot
	Prunus persica	Basal rot
	Pyrus communis	Foot rot, trunk
		gummosis

	Jujuy	Alcoba et.al. (2005)
	Jujuy	Alcoba et.al. (2005)
		Frezzi 1946 In: Frezzi (1950); Bergna
	Río Negro	et al. (1979); Bergna (1983, 1985);
		Rossini et al. (1997); Dobra et al.
		(2007); Sánchez et al. (2017)
	Jujuy	Alcoba et al. (2005)
	Mendoza	Feldman & Gracia (1981)
	Neuquén	Rossini et al. (1997)

SpeciesHostSymptoms

Asparagus officinalis

Damping-off, root

and crown rot, fruit rot, blight

Province	References1
Mendoza	Velasco & Camargo (1994)
	Lindquist (1932); Godoy (1940);
	Carrera (1942); Palmucci et al. (2011
Buenos Aires	a*b*); Palmucci (2015); Gobena
	et al. (2012); Iribarren et al. (2012
	a*b*, 2013*, 2015*, 2016*, 2019*)

Capsicum annuum

P.capsici Leonian

	Coronopus didimus	Yellowing
	Cucurbita spp.	Root and stem
		rot, fruit rot.
	C. pepo	Root and crown
		rot, fruit rot

	C. maxima var.	Leaves and fruit
		necrosis, root and
	zapallito
		crown rot, fruit rot

Córdoba

Chaco

Entre Ríos

Tucumán

Jujuy

Corrientes

Catamarca

La Rioja

La Pampa

Salta

San Luis

San Juan

Santa Fe

Formosa

Mendoza

Río Negro

Neuquén

Neuquén

Buenos Aires

Córdoba

Río Negro

Neuquén

Buenos Aires

Mendoza

Córdoba

Río Negro

Neuquén

Buenos Aires

Carrera (1941a) FF; Frezzi

1942 In: Frezzi (1950)

Bonacic Krecic & Campagnat (1982)

Hauman Merck & Parodi (1921)

Traversi (1948) FF

Godoy (1940); Alcoba et al. (2005)

Colombo (1988); Mazzanti

de Castañon (1972)

Raggi (1945) FF

Carrera (1940) FF

Vitoria (1941) FF

Godoy (1940)

Carranza (1945) RF

Oriolani & Gatica (1980)

Herzog & Rista (1989)

Mazzanti de Castañon et al. (1994); Bonacic Krecic & Campagnat (1982)

Godoy (1940); Bergna (1963); Oriolani

& Gatica (1980); Roig et al. (2009)

Bergna (1963); Rossini et al. (1992)

Cipolla (1948a) FF

Iribarren et al. (2017);

Yabar et al. (2017)

Frezzi 1945 In: Frezzi (1950)

Rossini et al. (1992)

Irribarren et al. (2012a*, 2016)

Pontis (1945); Felman & Pontis (1960)

Frezzi 1946 In: Frezzi (1950)

Rossini et al. (1992)

Mitidieri (1976);Vigliola &

Calot (1982); Irribarren et al.

(2012a*b*, 2015, 2019*)

Leaves and fruit

C. moschata necrosis, root and crown rot, fruit rot

	Río Negro	Rossini et al. (1992)
	Neuquén
	Buenos Aires	Palmucci et al. (2015*)

Species

Host

Symptoms

Province

Mendoza

References1

Oriolani & Gatica (1980);

Velasco (1981)

P.capsici Leonian

P.cinnamomi Rands

C. maxima

Cucumis melo

subsp. melo

Cucumis sativus

Ficus carica

Phaseolus lunatus

Phaseolus vulgaris

Philodendron

scandens subsp.

oxycardium

Solanum

lycopersicum

Solanum melongena

Actinidida deliciosa

Calycanthus floridus

Casuarina sp.

Casuarina

cunningamiana

Leaves, fruit and

seed necrosis.

Root and crown

rot, fruit rot

Root and crown

rot, fruit rot

Fruit rot

Necrosis of branches,

pods and seeds

Root and crown

rot, fruit rot

Leaf spots

Damping- off, fruit rot

Root and crown

rot, fruit rot

Basal rot

Root and trunk rot

Crown and trunk

cankers

Root and basal rot,

trunk cankers

San Juan

Río Negro

Neuquén

Buenos Aires

Río Negro

Neuquén

Mendoza

Córdoba

Río Negro

Neuquén

Jujuy

Córdoba

Río Negro

Neuquén

Corrientes

Mendoza

Corrientes San Juan

Buenos Aires

Río Negro

Neuquén

Córdoba

Buenos Aires

Mendoza

Río Negro

Neuquén

Buenos Aires

Córdoba

Buenos Aires

Entre Ríos

Buenos Aires

Oriolani & Gatica (1980)

Rossini et al. (1992)

Iribarren (2016)

Rossini et al. (1992); Velasco

& Perinetti (1990)

Frezzi 1947 In: Frezzi (1950)

Rossini et al. (1992)

Alcoba et al. (2005)

Frezzi 1947 In: Frezzi (1950)

Rossini et.al. (1992)

Cúndom & Cabrera (2004)

Frezzi 1948 In: Frezzi (1950);

Pontis & Feldman (1958);

Felman & Pontis (1960)

Colombo & Obregón (2009)

Oriolani & Gatica (1980)

Irribarren et al. (2012c*);

Iribarren et al. (2019*)

Rossini et.al. (1992)

Frezzi 1949 In: Frezzi (1950)

Irribarren et al. (2012d*)

Iribarren et al 2019*

Velasco & Pontis (1977)

Rossini et al. (1992)

Palmucci et al. (2015*)

Frezzi 1946 In: Frezzi (1950)

Frezzi 1952 In: Frezzi (1977)

Palmucci et al. (2008); Palmucci et al. (2011a*b*); Palmucci (2015*)

Bakarcic (1961)

Species

Host	Symptoms
Castanea sativa	Basal rot
Cupressus sp.	Root rot
Echeveria gibbiflora	Root rot
Eucalyptus
camaldulensis

Province	References1
Buenos Aires	Marchionatto (1953)
Córdoba	Frezzi 1946 In: Frezzi (1950)
Córdoba	Frezzi 1949 In: Frezzi (1950)
	Sarasola & de Sarasola (1959)

P.cinnamomi Rands

E. rostrata

Iresine sp.

Juglans regia

Ligustrum lucidum

Persea americana

Phaseolus lunatus

Pinus radiata-

insigne

Pittosporum tobira

Platanus orientalis

Prunus persica

Quercus robur

Rhododendron sp.

Rhododendron

indicum

Salix caprea

Schinus molle

Sedum sp.

Solanum

lycopesicum

Spiraea

cantoniensis

Thuja sp.

Root rot, necrosis of

leaves and twigs Necrosis in leaves Crown rot

Root and crown rot

Root and crown rot

Necrosis of pods

and seeds

Root rot

Necrosis of leaves

and twigs

Cankers of trunk

Root and crown rot

Root and crown rot

Root rot and

necrosis of trunk

Root rot and

necrosis of trunk

Root rot and

necrosis of trunk

Root rot and necrosis

of leaves and twigs

Root rot and necrosis

of leaves and twigs

Fruit rot

Root rot

Root rot

	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Jujuy	Alcoba et al. (2005)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1954 In: Frezzi (1977)
	Salta	Frezzi (1952); 1951 In: Frezzi
		(1977); Vazquez de Ramallo (1974)
	Jujuy	Vazquez de Ramallo (1974);
		Flores et al. (2006)
	Tucumán	Frezzi 1972 In: Frezzi (1977)
	Córdoba	Frezzi (1950)
	Córdoba	Frezzi 1946 In: Frezzi (1950)
	Buenos Aires	Bakarcic (1961)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Buenos Aires	Frezzi 1949 In: Frezzi
		(1950); Bakarcic (1951)
	Buenos Aires	Frezzi 1952 In: Frezzi
		(1977); Bakarcic (1961)
	Buenos Aires	Bakarcic (1961)
	Buenos Aires	Frezzi 1949 In: Frezzi (1950)
		Frezzi 1949 In: Frezzi
	Buenos Aires	(1950); Palmucci et al. (2011
		a*b*); Palmucci (2015*)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1946 In: Frezzi (1950)

SpeciesHost

Symptoms

Province	References1
Salta	Hongn et al. (2003)

P.cinnamomi Rands

P.citricola complex (Syn:

P.cactorum (Leb. & Cohn) Schroet. var. applanata Chest.)

P.citrophthora (R.E. Sm. & E.H.

Sm.) Leonian

Vaccinium

corymbosum

Brachytchiton

populneum

Populus alba

Schinus molle

Acer negundo

Araucaria

angustifolia

Antirrhinum majus

Bauhinia forficata

(=B.candicans)

Bignonia sp.

Brachychiton

acerifolia

B. populneum

Capsicum annuum

C. pubescens

Root and crown rot

Necrosis of the apex

Necrosis of

basal leaves

Root rot and necrosis of stem, leaves and apex of potted plants

Crown and trunk rot

Nurseries: necrosis

of the plant apex and plant death

Root rot

Root rot in

young plants

Root and stem rot

Root rot and death of plants (two-years-old)

Root rot, necrosis of

stem and death of

plants (two-years-old)

Damping off, root

and basal rot, fruit rot and death of plants

Root rot, necrosis of stem and death of plants.

	Tucumán	Hongn et al. (2003); Palmucci et
		al. (2011a*b*); Palmucci (2015*)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1951 In: Frezzi (1977)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1946 In: Frezzi (1950)
	Córdoba	Frezzi 1943 In: Frezzi (1950)
	Córdoba	Frezzi 1953 In: Frezzi (1977)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1956 In: Frezzi (1970)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1943 In: Frezzi (1950)
	Mendoza	Feldman 1976 In: Frezzi (1977);
		Felman & Gracia (1979)
	Salta	Frezzi 1952 In: Frezzi (1977)
	Córdoba	Frezzi 1949 In: Frezzi (1950)

Castanea sativa	Nurseries: root rot
Catharanthus	Root rot and necrosis
roseus	of the apex
Celtis australis	Root and crown rot
Cicer arietinum	Root rot
	Gummosis, damping
Citrus aurantifolia	off, foot rot and
	brown rot of fruits

Mendoza	Frezzi 1955 In: Frezzi (1977)
Córdoba	Frezzi 1947 In: Frezzi (1950)
Córdoba	Frezzi 1942 In: Frezzi (1950)
Córdoba	Frezzi 1946 In: Frezzi (1950)
Córdoba	Frezzi 1943 In: Frezzi (1950)

SpeciesHost

Citrus aurantifolia

C. limon

C. limetioides

Citrus jambhiri

P.citrophthora (R.E. Sm. & E.H.

Sm.) Leonian

C. sinensis

C. aurantium

C. nobilis

Symptoms

Gummosis, damping

off, foot rot and

brown rot of fruits

Damping off, foot

rot, gummosis, and

brown rot of fruits

Damping-off, crown

rot of fruits

Damping-off

Damping off, foot rot

Gummosis of twigs,

Brown rot of fruits

Brown rot of fruits

Province

Corrientes

Corrientes

Córdoba

Tucumán

Entre Ríos

Buenos Aires

Jujuy

Corrientes

Tucumán

Córdoba

Corrientes

Corrientes

Córdoba

Entre Ríos

Buenos Aires

Tucumán

Salta

Jujuy

Corrientes

Corrientes

Corrientes

References1

Frezzi (1942); Frezzi 1941 In: Frezzi (1950); Mazanti de Castañón (1972)

Frezzi 1939 In: Frezzi 1950; Frezzi (1940; Frezzi (1942

Frezzi 1950 In: Frezzi (1977)

Stein & Torres Leal (1991)

Cipolla (1948b) FF

Frezzi (1950)

Marchionatto (1948);

Carranza (1954) RF

Bejarano & Catacata (2009)

Frezzi (1942); Frezzi 1941 In: Frezzi (1950); Mazanti de Castañón (1972)

Frezzi (1940)

Frezzi & Mácola (1945)

Frezzi 1941 In: Frezzi (1950); Mazanti de Castañón (1972)

Frezzi (1938); Frezzi (1940); Frezzi

(1942); Frezzi 1939 In: Frezzi

(1950); Marchionatto (1948);

Mazanti de Castañón (1972)

Frezzi 1942 In: Frezzi (1950);

Frezzi & Mácola (1945)

Marchionatto (1948)

Marchionatto (1948); Frezzi

(1960 In: Frezzi (1977)

Frezzi 1967 In: Frezzi (1977)

Frezzi 1944 In: Frezzi (1950)

Bejarano & Catacata (2009)

Frezzi 1939 In: Frezzi (1950);

Frezzi (1940, 1942); Mazanti

de Castañón (1972)

Frezzi 1941 In: Frezzi (1950)

Frezzi 1940 In: Frezzi (1950); Frezzi 1942; Mazanti de Castañón (1972)

	C. paradisi	Brown rot of fruits	Córdoba
			Buenos Aires
			Tucumán
	C. reticulata	Brown rot of fruits	Corrientes
	var. austera
	C. reticulata	Damping-off, brown	Corrientes
		rot of fruits

Frezzi 1951 In: Frezzi (1977)

Frezzi 1960 In: Frezzi (1977)

Frezzi (1940)

Frezzi 1941 In: Frezzi

(1950); Frezzi (1942)

Frezzi 1940 In: Frezzi (1950);

Marchionatto (1948); Mazanti

de Castañón (1972)

Species

HostSymptoms

Province	References1
Entre Ríos	Marchionatto (1948)

P.citrophthora (R.E. Sm. & E.H.

Sm.) Leonian

C. reticulata

Clerodendron sp.

Cucumis sativum

Cucurbita maxima

Cucurbita maxima

var. zapallito

Dianthus

caryophyllus

D.sinensis Fagara coco

Ficus carica

Fragaria vesca

Fortunella sp.

Gloxinia sp.

Gomphrena sp.

Hedera helix

Juglans nigra

Laurus nobilis

Damping-off, brown

rot of fruits

Necrotic lesions

on leaves

Fruit rot

Root rot

Root rot, necrosis

of leaves

Root rot

Root rot

Necrosis of apex, lesions on leaves and stem

Necrosis on leaves

Root rot

Necrosis of buds

near the soil

Stem and

leaves blight

Root rot and

death of plants

Necrosis of stem

Necrosis of leaves

(one-year-old

plants), crown rot

(adult plants)

Necrosis of leaves

and twig blight

	Salta	Alcoba et al. (2005)
	Buenos Aires	Marchionatto (1948)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1944 In:Frezzi (1950)
	Córdoba	Frezzi 1946 In: Frezzi (1950)
	Salta	Frezzi 1953 In: Frezzi (1977)
	Buenos Aires	Mitidieri (1976)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1943 In: Frezzi (1950)
	Córdoba	Frezzi 1950 In: Frezzi (1977)
	Córdoba	Frezzi 1959 In: Frezzi (1977)
	Argentina	Marchionatto (1950)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Pontis (1941); Frezzi 1948 In:Frezzi
		(1950); Marchionatto (1948)
	Catamarca	Pontis (1941); Carrera (1951) RF
	Jujuy	Alcoba et al. (2005)
		Pontis (1941); Marchionatto (1948);
	Mendoza	Frezzi 1947 In: Frezzi (1950);
		Feldman & Pontis (1960)
	San Juan	Pontis (1941); Marchionatto (1948)
	La Rioja	Pontis (1941); Carrera (1947) RF
	Río Negro	Bergna (1963)
	Córdoba	Frezzi 1961 In: Frezzi (1977)

Species

HostSymptoms

Province

References1

P.citrophthora (R.E. Sm. & E.H.

Sm.) Leonian

P.cryptogea Pethybr. & Laff

Magnolia grandiflora

Melia azedarach

Nerium oleander

Olea europea

Petroselinum

crispum

Pinus halepensis

P.pinea P. radiata

Poncirus trifoliata

Psidium guayaba

Populus simonii Punica granatum

Robinia

pseudoacacia

Ruta sp.

Salvia splendens

Solanum

lycopersicum

Acer negundo

Actinidida deliciosa

Callistephus

chinensis

Gerbera jamesonii

Helianthus annus

Ipomea batatas

Ocimum basilicum

Prunus avium

Populus simonii

Necrosis of twigs

and plant apex

Root rot, necrosis of the apex, defoliation

Necrosis basal

of new stems

Root rot, wilt,

defoliation

Necrosis of roots

and leaves

Necrosis of the apex and twigs

Necrosis of the apex

Necrosis of apex

and twigs

Damping-off

Fruit rot

Root rot

Fruit rot

Root and trunk rot

Root rot

Root rot and necrosis

of branches

Fruit rot

Root rot, basal

canker and death

of adult plants

Root and crown rot

Root and crown rot,

death of plants

Basal rot

Root and crown rot

Root rot and

death of plants

Basal rot

Leaf yellowing

and decline

Root rot and

cankers on trunk

	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1956 In: Frezzi (1977)
	Mendoza	Lucero et al.(2011b*)
	Córdoba	Frezzi 1958 In: Frezzi (1977)
	Córdoba	Frezzi 1943 In: Frezzi (1950)
	Córdoba	Frezzi 1947 In: Frezzi, (1950)
	Córdoba	Frezzi 1947 In: Frezzi, (1950)
	Corrientes	Frezzi (1950); Mazanti de
		Castañón (1972)
	Córdoba	Frezzi 1953 In: Frezzi (1977)
	Mendoza	Feldman & Pontis (1960)
	Córdoba	Frezzi 1953 In: Frezzi (1977)
		Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1951 In: Frezzi (1977)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Córdoba	Frezzi 1952 In: Frezzi (1977)
	Buenos Aires	Wolcan & Ronco (2006); Palmucci
		et al. (2011 a*b*); Palmucci (2015*)
	Mendoza	Pontis & Feldman (1959)
	Buenos Aires	Wolcan & Ronco (2006)
	Buenos Aires	Wolcan & Ronco (2006); Palmucci
		et al. (2011a*b*); Palmucci (2015*)
	Buenos Aires	Kiehr et al. (2002); Paoloni et al.(2002)
	Buenos Aires	Mitidieri 1971 In: Frezzi (1977)
	Buenos Aires	Wolcan & Ronco (2006)
	Mendoza	Lucero et al. (2011a*)
	Mendoza	Frezzi (1957)

Species

Host

Symptoms

Province

References1

P.cryptogea Pethybr. & Laff

P.drechsleri Tucker

P.inundata

Brasier, Sánch.

Hern. & S.A. Kirk

Salvia officinalis

Solanum

lycopersicum

Brassica oleracea

var. capitata

Capsicum annuum

C. plumosa

Chrysanthemum cinerariaefolium

Coleus spp.

Pelargonium zonale

Pyrus communis

Salvia officinalis

Schinus molle

Senecio cruentus

Solanum

lycopersicum

Solanum melongena

Pyrus communis

Capsicum annum

Solanum melongena

Basal rot

Damping off

Root and crown rot

Fruit rot and

necrosis of stem

Root rot and necrosis of stem and branches

Root and basal rot

Root rot and death of plant

Necrosis of roots

Fruit rot (in plant)

Root rot

Necrosis of leaves

and stem

Root and crown rot

Rot of green fruits

Necrosis of stems

and fruit rot

Preharvest fruit rot and root rot

Late blight and

fruit rot

Fruit rot and wilt

	Buenos Aires	Wolcan & Ronco (2006)
	Buenos Aires	Mitidieri (1995)
	Buenos Aires	Iribarren et al. (2012d*)
	Buenos Aires	Iribarren et al. (2019*)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1945 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Río Negro	Sosa et al. (2015*)
	Neuquén
	Mendoza	Klingner (1969); Frezzi (1977)
	Córdoba	Frezzi 1948 In: Frezzi (1950)
	Buenos Aires	Carranza (1963)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Buenos Aires	Iribarren et al. (2017*)
	Río Negro	Sánchez et al. (2017*; 2019*)
	Buenos Aires	Prioletta (2006)
	Northwest	Ganem et al. (1992)
	Region
	Jujuy	Godoy (1941) FF; Alcoba et al. (2005)

SpeciesHost

	P. infestans	S. tuberosum
	(Mont.) de Bary

		Gerbera jamesonii
	P. taxon kelmania	Gypsophilla
	Abad & Abad
		paniculata
		Actinidia deliciosa

P.lacustris

Brasier, Cacciola,

Nechwatal, Jung

Pyrus communis

&Bakonyi (ex P. taxon “Salixsoil”)

Symptoms	Province
	Buenos Aires
	Entre Ríos
	Chaco
	Santa Fe
	Córdoba
Late blight and	Misiones
downy mildew	Jujuy
	Río Negro
	Corrientes
	San Juan
	Mendoza
	Salta
	Catamarca
Root and basal rot	Buenos Aires
Root and basal rot	Buenos Aires
Root and basal rot	Buenos Aires
Postharvest fruit	Rio Negro
rot, fruit rotin plant	Neuquén

References1

Rieder (1887); Girola (1898);

Spegazzini (1902); Marchionatto

(1939); Godoy (1943); Frezzi (1950);

Van Damme & Ridao (1994); Lucca &

Huarte (2012); Servici et al. (2017*)

Cipolla (1948c) FF

Calderoni & Malamud (1966)

Hauman-Merck (1915);

Carrera (1941b) FF

Frezzi (1950)

Carrera (1949) FF

Alcoba et al. (2005)

Pelayo (1935) FF; Bergna (1963); Colavita et al. (2005)

Mazanti de Castañón (1972)

Oriolani & Gatica (1980)

Feldman & Pontis (1960);

Oriolani & Gatica (1980)

Frezzi 1950 In: Frezzi (1977);

Alcoba et al. (2005)

Schechaj (1954)

Palmucci et al. (2011a*b*);

Palmucci (2015*)

Palmucci et al. (2015)

Palmucci et al. (2015*)

Dobra et al. (2011*);

Sosa et al. (2015*)

	P. medicaginis	Medicago sativa	Root rot	Santa Fe
	Hansen &
	Maxwel (ex P.	Medicago sativa	Root rot	Buenos Aires
	megasperma
	f.sp. medicaginis
	Kuan & Erwin	Medicago sativa	Root rot	Córdoba
		Cicer arietinum	Root rot	Córdoba
	P. megasperma	Citrus limon	Brown rot of fruits	Buenos Aires
	Drechsler	C. paradisi	Brown rot of fruits	Buenos Aires
		C. sinensis	Brown rot of fruits	Corrientes

Hijano et al. (1986); Gieco et al. (2007)

Hijano et al. (1986)

Hijano et al. (1986); Hijano

1979 In: Hijano (1987)

Frezzi (1950)

Lindquist 1936 In: Frezzi (1950)

Frezzi (1950)

Frezzi (1942); Mazanti de

Castañón (1972)

Species

HostSymptoms

Province

References1

P.multivora Scott & Jung

P.nicotianae

Breda de Haan

(Syn: P. parasitica)

Dastur

Rhododendron spp.

Begonia sp.

Bignonia sp.

Brachychiton

populneum

Callistephus

chinensis

Carthamus tinctorius

Celosia plumosa

Catharanthus

roseus

Citrus aurantium

Citrus aurantifolia

Citrus limettioides

Citrus limon

Citrus nobilis

Citrus paradisi

Citrus reticulata

var. austera

Citrus sinensis

Dark spots from

the leaf apex

Necrotic lesions

on leaves

Root and stem rot

Root rot and

necrosis of stem

Root rot

Root rot

Root rot and

necrosis of stem

Apical necrosis

and root rot

Damping-off and foot rot

Damping off, foot rot and brown rot of fruits

Damping off

Foot rot, trunk gummosis and brown rot of fruits

Foot rot and brown

rot of fruits

Foot rot and brown

rot of fruits

Foot rot and brown

rot of fruits

Damping.off, foot rot and brown rot of fruits

Buenos Aires

Córdoba

Córdoba

Córdoba

Córdoba

Córdoba

Córdoba

Córdoba

Buenos Aires

Corrientes

Córdoba

Corrientes

Buenos Aires

Corrientes

Corrientes

Santiago

del Estero

Tucumán

Salta

Buenos Aires

Jujuy

Corrientes

Córdoba

Buenos Aires

Corrientes

Corrientes

Corrientes

Grijalba et al. (2012*, 2013*); Grijalba & Palmucci (2014)

Frezzi 1949 In: Frezzi (1950)

Frezzi 1956 In: Frezzi (1977)

Frezzi 1948 In: Frezzi (1950)

Frezzi 1947 In: Frezzi (1950)

Frezzi 1949 In: Frezzi (1950)

Frezzi 1947 In: Frezzi (1950)

Frezzi 1946 In: Frezzi (1950)

Rivera et al. (2000); Palmucci et al. (2011a*b*); Palmucci (2015)

Frezzi, 1939 In: Frezzi (1950); Mazanti de Castañón (1972)

Frezzi 1942 In: Frezzi (1950)

Frezzi (1938); Frezzi 1937 In: Frezzi (1950); Mazanti de Castañón (1972)

Frezzi 1960 In: Frezzi (1977) Mazanti de Castañón (1972)

Frezzi (1938); Frezzi 1937 In: Frezzi

(1950); Frezzi (1940); Frezzi (1942);

Mazanti de Castañón (1972)

Frezzi 1948 In: Frezzi (1950)

Frezzi (1940); Frezzi 1939 In: Frezzi (1950); Stein & Torres Leal (1991)

Frezzi 1944 In: Frezzi (1950)

Cipolla (1948d) FF; Frezzi

1960 In: Frezzi (1977)

Bejarano & Catacata (2009)

Frezzi (1938); Frezzi 1939 In:

Frezzi (1950); Frezzi (1942)

Frezzi 1942 In: Frezzi (1950);

Frezzi & Mácola (1945)

Frezzi 1960 In: Frezzi (1977)

Frezzi (1938); Frezzi (1942); Frezzi (1950); Mazanti de Castañón (1972)

Frezzi 1938 In: Frezzi (1950);

Mazanti de Castañón (1972)

Fawcett (1936); Frezzi 1937 In: Frezzi (1950); Frezzi (1942); Mazanti de Castañón (1972)

SpeciesHostSymptoms

Province

Córdoba

References1

Carrera (1935) FF; Frezzi (1938);

Frezzi 1942 In: Frezzi (1950);

P.nicotianae

Breda de Haan

(Syn: P. parasitica)

Dastur

Citrus sinensis

Chamelaucium

uncinatum

Cryptomeria

japonica

Cucumis melo

Cucurbita sp.

Dianthus

caryophillus

Dianthus sinensis

Diefembachia picta

Equinucactus sp.

Eucalyptus viminalis

Fragaria sp.

Fucsia sp.

Gypsophila paniculata

Hedera helix

Heliotropium peruvianum

Hebe speciosa

Hibiscus cannabis

Damping.off, foot rot and brown rot of fruits

Root and basal rot

Apical necrosis

Damping off, fruit rot

Fruit rot

Root rot

Root rot

Root and basal rot

Root and crown

necrosis

Apical necrosis

Death of adult plants

Root rot

Root and basal rot

Necrotic spots

on leaves

Necrotic spots

on leaves

Root and basal rot

Stem bligth

Santiago

del Estero

Salta

Buenos Aires

Córdoba

Córdoba

Buenos Aires

Buenos Aires

Córdoba

Córdoba

Buenos Aires

Córdoba

Córdoba

Corrientes

Córdoba

Buenos Aires

Córdoba

Córdoba

Buenos Aires

Salta

Frezzi & Mácola (1945)

Frezzi 1948 In: Frezzi (1950)

Frezzi 1952 In: Frezzi (1977)

Wolcan et al. (2005);

Palmucci et al. (2013*)

Frezzi 1947 In: Frezzi (1950)

Frezzi 1947 In: Frezzi (1950)

Mitidieri 1973 In: Frezzi (1977)

Mitidieri (1973)

Frezzi 1944 In: Frezzi (1950)

Frezzi 1949 In: Frezzi (1950)

Grijalba et al. (2007, 2008*);

Palmucci et al. (2011a*b*c*)

Frezzi 1953 In: Frezzi (1977)

Frezzi 1949 In: Frezzi

(1950); Merlo (1981)

Frezzi, 1940 In: Frezzi (1950)

Frezzi 1947 In: Frezzi (1950)

Wolcan et al. (2007); Palmucci (2015*)

Frezzi 1949 In: Frezzi (1950)

Frezzi 1948 In: Frezzi (1950)

Palmucci (2015*)

Vecchietti & Zapata (2005)

Impatiens balsamina Root rot and necrosis of branches

	Ipomea batatas	Root rot
	Magnolia grandiflora	Necrosis of apex
		and stem
	Melia azedarach	Necrosis of apex
		and stem
	Nerium oleander	Necrosis of crown
	Nicotiana tabacum	Black collar

Córdoba

Buenos Aires

Córdoba

Córdoba

Córdoba

Salta

Jujuy

Frezzi 1948 In: Frezzi (1950)

Frezzi (1977)

Frezzi 1947 In: Frezzi, (1950

Frezzi 1947 In: Frezzi (1950)

Frezzi 1956 In: Frezzi (1977)

Castelló (1986)

Species	Host	Symptoms
	Olea europea	Root rot and
		twigs wilting
	Oncidium sp.	Necrosis of base
		of leaves
	Phaseolus lunatus	Necrosis of pod
		and green seed
	Phaseolus vulgaris

Province	References1
Mendoza	Vettraino et al. (2009*)
Córdoba	Frezzi 1958 In: Frezzi (1977)
Córdoba	Frezzi 1947 In: Frezzi (1950)
Salta	Alcoba et al. (2005)

P.nicotianae

Breda de Haan

(Syn: P. parasitica)

Dastur

P. palmivora (E.J. Butler) E.J. Butler

Pinus halepensis

Punica granatum

Primula obconica

Rheum rhaponticum

Ricinus communis

Salvia splendens

Schinus molle

Sesamum indicum

Simmondsia

chinensis

Solanum

lycopersicum

Solanum melongena

Tradescantia

virginiana

Schlumbergera

truncata

Brachychiton

populneum

Chamaedorea

elegans

Chamaerodea spp.

Citrus limon

Citrus sinensis

Root rot

Necrotic lesions

of leaves

Root and crown rot

Crown rot

Leaf bligth

Root rot and necrosis

of branches

Apical necrosis of twigs and branches

Root rot

Crown necrosis

and wilt

Fruit rot

Fruit rot

Root rot and

necrosis of leaves

Root and cladodes rot

Root rot

Leaf spots

Leaf spots

Brown rot of fruits

Foot rot, brown

rot of fruits

	Córdoba	Frezzi 1945 In: Frezzi (1950)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Buenos Aires	Palmucci et al. (2015)
	Buenos Aires	Lindquist (1935) RF
	Tucumán	Vazquez de Ramallo (1989)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Córdoba	Frezzi 1943 In: Frezzi (1950)
	La Rioja	Lucero et al. (2006)
	Córdoba	Frezzi 1949 In: Frezzi (1950)
	Buenos Aires	Irribarren et al. (2014*, 2019*)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Buenos Aires	Irribarren et al. (2014*, 2019*)
	Córdoba	Frezzi 1947 In: Frezzi (1950)
	Buenos Aires	Palmucci et al. (2012*),
		Palmucci (2015*)
	Córdoba	Frezzi 1946 In: Frezzi (1950)
	Corrientes	Cúndom et al. (2006)
	Chaco	Cúndom et al. (2004)
	Corrientes
	Córdoba	Frezzi & Mácola (1943)
	Córdoba	Frezzi 1942 In: Frezzi (1950);
		Frezzi & Mácola (1943)

	Species	Host	Symptoms
		Citrus sinensis	Foot rot, brown
			rot of fruits
		Eucalyptus viminalis	Damping off
	P. palmivora (E.J.	Ficus carica	Fruit rot
	Butler) E.J. Butler
		Olea europea	Root rot, die-
			back, wilt
		Phoenix dactylifera	Damping off
	P. rosacearum		Preharvest fruit
	E.M. Hansen &	Pyrus communis
			rot and root rot
	W.F. Wilcox

	P. sojae	Glyicine max	Root and basal rot
	Kaufm. & Gerd.

Province

Misiones

Argentina

Tucumán

Catamarca

San Juan

La Rioja

Córdoba

Río Negro

Tucumán

Salta

Buenos Aires

Córdoba

References1

Fawcett 1937 In: Fawcett

& Bitancourt (1940)

Sarasola & de Sarasola (1959)

González et al. (1997, 1998)

Prioletta (2006); Lucero et al. (2007*)

Carrera (1954) FF

Sánchez et al. (2017*; 2019*)

Ploper et al. (1998, 1999); Distéfano

de Vallone et al. (1999)

Martinez & Ivancovich (1979); Barreto

et al. (1991, 1995, 1998ab; 2005); Gally et al. (2007*); Distéfano de Vallone et al. (1999); Palmucci et al. (2011 a*b*; Grijalba et

al. (2011*); Grijalba & Ridao

(2013*); Grijalba & Gally (2015);

Grijalba et al. (2014*, 2018*)

Barreto et al. (1997, 1998ab; 2005); Distéfano de Vallone et al. (1999); Gally et al. (2007*); Grijalba & Gally (2015)

				Entre Ríos
	P. syringae	Citrus limon	Brown rot of fruits	Buenos Aires
	Klebahn	Citrus sinensis	Brown rot of fruits	Buenos Aires
				Entre Ríos
	P. thermophila		Preharvest fruit
	T. Jung, M.J.C.	Pyrus communis		Río Negro
			rot and root rot
	Stukely & T. Burgess

Barreto et al. (1997, 1998ab); Gally et al.(2007*); Grijalba & Gally (2015)

Frezzi (1950)

Frezzi (1950)

Pereda (1955) FF

Sánchez et al. (2017*, 2019*)

Bol. Soc. Argent. Bot. 55 (2) 2020

2011), P. cryptogea on two hosts (Lucero et al., 2011a; Palmucci et al., 2011a, b; Palmucci, 2015), P. citrophthora (Lucero et al., 2011b), P. taxon kelmania (Palmucci et al., 2011a, b; Palmucci, 2015), P. sojae (Grijalba et al., 2011; Palmucci et al., 2011a, b), P. capsici (Palmucci et al., 2011a, b; Palmucci, 2015; Gobena et al., 2012; Iribarren et al., 2012a, b; 2013), P. drechsleri (Iribarren et al., 2012b, e; Sosa et al., 2015; Iribarren et al., 2017), P. multivora (Grijalba et al., 2013; Grijalba

&Palmucci, 2014), and P. inundata, P. rosacearum, and P. termophila (Sanchez et al., 2017, 2019).

During the last few years, some studies about the genetic variability of populations of P. austrocedri (Velez et al., 2014), P. capsici (Gobena et al.,

2012; Iribarren et al., 2013, 2015, 2016) and P. infestans (Lucca & Huarte, 2012) were carried out. Also others about the virulence and races of Phytophthora sojae were carried out (Grijalba & Ridao, 2013; Grijalba et al., 2014, 2018; Grijalba & Gally, 2015).

The species affecting a greater number of hosts are P. citrophthora (55), P. nicotianae (54) and P. cinnamomi (31) (Table 2). On the other hand, other species as P. austrocedri (1), P. medicaginis (1) and P. sojae (1) have specific hosts, and some others, which have been recently identified, as P. inundata,

P.lacustris, P. multivora, P. rosacearum and P. thermophila, have been reported only on a single host to date (Table 2).

The present inventory shows that there are 20 Phytophthora spp., one species affinis and one taxon reported in Argentina, affecting a total of 223 hosts. Considering the percentage of host- pathogen relations of Phytophthora species, P. nicotianae and P. citrophthora are the main species with 24%, followed by P. cinnamomi with 14% (Fig. 1).

The distribution of Phytophthora species in the different provinces and regions of Argentina are shown in Table 3. Córdoba and Buenos Aires provinces present the greatest number of species, followed by Tucumán, Río Negro and Corrientes. According to the results, P. capsici presents the highest distribution along the country, followed by P. cactorum, P. infestans, and P. citrophthora (Table 3).

Phytophthora spp

P.austrocedri P. aff. boehmeriae

P. cactorum

P. capsici

P. cinnamomi

P. citricola

P. citrophthora

P. cryptogea

P.drechsleri

P.infestans

P.inundata P. taxon kelmania

P.lacustris

P.medicaginis P. megasperma

P.multivora P. nicotianae

P.palmivora

P.rosacearum P. sojae

P.syringae P. termophila

P. spp./ province

Northwest

Northeast

Cuyo

Pampas

Patagonia

/

Juj Sal Tuc Cat SE Mis ChaCtes ER FsaMza LR SJ SL BASFCbaLP RN Chu SC Neu TF

Provinces P. sp.

X

X

X

3

X

1

X

X X

X X

X X X

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DISCUSSION

So far, the most complete international review regarding Phytophthora spp. was the one published by Erwin & Ribeiro in the USAin 1996. These authors provided broad information about Phytophthora spp. worldwide, and presented different types of tables about their worldwide distribution, morphology, physiology, and taxonomic evolution, as well as disease control measures.

The present review allowed the updating of the previous status and inventory of the Phytophthora species present in Argentina (Frezzi, 1950, 1977) as well as of the nomenclature of Phytophthora spp. in relation to that of Frezzi’s compilations (Frezzi, 1950, 1977).

There are some issues to be highlighted regarding some species. P. boehmeriae was first mentioned affecting Citrus sinensis in Corrientes province (Frezzi, 1941) (Table 1). Sequences of some of these isolates have been recently used in phylogenetic studies (G. Abad, USDA-APHIS-PPQ, Pers. comm.), in which it was observed that they were not grouped with the isolate sequence Type of P. boehmeriae, whereby the isolates obtained by

Frezzi were considered P. aff. boehmeriae (Abad, Pers. comm.). In addition, after 1941, P. boehmeriae has not been cited in Argentina again, except by Wolcan & Lori (2001), who report it causing basal and root rot of Chamelaucium uncinatum. However, more detailed morphobiometric and cultural studies that were made later allowed identifying this oomycete as P. nicotianae (Wolcan et al., 2005), a fact confirmed with the application of molecular analyses and ITS sequencing (Palmucci et al., 2011a, b; Palmucci, 2015). Thus, P. boehmeriae was deleted from the list of Phytophthora spp. in Argentina and the record by Frezzi was replaced by P. aff. boehmeriae.

Frezzi also cited P. cactorum var. applanata on three trees (Frezzi, 1950). Waterhouse (1957) later compared the P. citricola original isolates with P. cactorum var. applanata and concluded that they were the same species, prioritizing the name P. citricola. Currently, P. citricola is recognized as a “species complex” The name P. citricola is retained only for isolates whose morphology and sequence information has an exact match with the isolate Type (CBS 221.88) (Lamour, 2013). Since there are no molecular studies of the Argentinian

isolates cited in Table 1, they were considered as “P. citricola complex”.

Another species that was not taken into account in Table 1 was P. fragariae. Although there are two reports mentioning this species in the country, neither of them describes the pathogen. In his book about the culture of Fragaria sp., Folquer (1986) named the Phytophthora spp. that caused basal rot on this crop as P. fragariae. Later, Alcoba et al. (2005) cited P. fragaria again, in Tucumán province, but based only on the symptoms of the plants.

During the first decades of the past century, plant diseases of different crops were recorded by phytopathologists from the Argentine Ministry of Agrarian Affairs along the country. The records were noted in phytopathologycal record cards, which were later compiled by the Institute of Agricultural Technology (INTA). These records about Phytophthora spp. were consulted from the Atlas Fitopatológico de Argentina (Nome et al., 2019). Because of their historical value, some of them were included in Table 1 when it corresponded.

According to the recent online resource “IDphy” (https://idtools.org/id/phytophthora/index.php) there were 182 species described worldwide until May 2018 (Abad et al., 2019). Six new species recently described were not included in IDphy yet. Of the 182 species, 167 can be cultivated.

Taking into account that P. boehmeriae was deleted from the list of species, in Argentina, there are currently 20 Phytophthora species, P. aff. boehmeriae and P. taxon kelmania, cited on 223 host-pathogen relationships. The diversity of Phytophthora species in the world suggests that a higher number of species, still not cited here, could be present in Argentina. Further studies with an extended range of hosts would be necessary to determine in detail the diversity of the Phytophthora population in different cropped areas. It is hoped that molecular and secuentiation techniques will allow a better knowledge of the diversity of Phytophthora species present in Argentina.

CONCLUSIONS

From the 19th century until now, 20 Phytophthora spp., one species affinis and one taxon of Phytophthora have been recorded in Argentina. In recent years, 30% of them have been identified. Researchers specialized

in the genus Phytophthora, molecular techniques and phylogenetic studies, may allow progressing in the accurate identification of the species and knowledge of their genetic variability.

In the context of global climate change, which predisposes the emergence and spread of biotical diseases, this work suggests the need to maximize the study of ecology, species richness, dispersion, pathogenesis, new host-pathogen relations (host switching ability) and the host-Phytophthora- environment interactions. This applies both to climate change as a whole as well as to individual factors such as temperature, rainfall, humidity, among others.

AUTHORS’ CONTRIBUTION

The authors shared the tasks of gathering and interpretation of the information, the preparation of tables and figure and the writing of the manuscript.

ACKNOWLEDGEMENTS

We thank Dr. Gloria Abad for her remarkable comments and suggestions and Ing. Agr. Carlos Carloni for providing some valuable bibliographic references. This paper was written as part of the Ubacyt 033BA Project.

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