Curvularia
Boedijn Bull. Jard. Bot. Buitenzorg, 3 Sér. 13: 123. 1933.
- Synonyms: Malustela Bat. & J.A. Lima, Publ. Inst. Micol. Recife 263: 5. 1960.
- Curvusporium Corbetta as “Curvosporium”, Riso 12: 28, 30. 1963.
- Pseudocochliobolus Tsuda, et al., Mycologia 69: 1117. 1978.
- Classification: Dothideomycetes, Pleosporomycetidae, Pleosporales, Pleosporaceae
- Type species: Curvularia lunata (Wakker) Boedijn. Ex-neotype culture: CBS 730.96
- DNA barcodes (genus): LSU, ITS.
- DNA barcodes (species): ITS, gapdh, tef1.
Ascomata pseudothecial, mostly globose to ellipsoidal, sometimes flask-shaped or flattened on hard substrata, brown or black, immersed, erumpent, partially embedded or superficial, free or developing on a basal columnar or flat stroma, smooth or covered with vegetative filaments; ostiole central, papillate or with a sub-conical, conical, paraboloid or cylindrical neck; ascomatal wall comprising pseudoparenchymatous cells of equal thickness or slightly thickened at apex of the ascoma. Hamathecium comprising septate, filiform, branched pseudoparaphyses. Asci bitunicate, clavate, cylindrical-clavate or broadly fusoid, straight or slightly curved, thin-walled, fissitunicate, often becoming more or less distended prior to dehiscence, short pedicellate, rounded at the apex. Ascospores multiseriate, filiform or flagelliform, hyaline or sometimes pale yellow or pale brown at maturity, septate, helically coiled within ascus, degree of ascospore coiling moderate to very strongly coiled, often with a mucilaginous sheath. Conidiophores straight to flexuous, often geniculate, multiseptate, usually simple, sometimes branched, smooth to verruculose, macronematous, mononematous, sometimes nodose, cylindrical. Conidiogenous nodes cylindrical, integrated, terminal and intercalary, proliferating sympodially, cicatrized. Conidia solitary, often curved, acropleurogenous, broadly fusoid, elliptical, obovoid or obpyriform, mostly smooth, sometimes verruculose, echinulate or tuberculate, 3 or more distoseptate, with or without an unequally swollen cell which is more pigmented than the other cells, septa sometimes accentuated with a dark band in some or all the cells, germinating mainly from one or both polar cells with the basal germ tube growing semiaxially, hilum in a slightly protruding truncate basal section of the conidial wall and often visible as two dark lenticular spots in optical section arranged close together with a small obscure narrow separating canal between them or distinctly protuberant, first conidial septum median or submedian, second septum often delimiting the basal cell of the mature conidium, third septum then distal. Microconidiation not common, producing conidia 1–2-celled, pale brown, globose to subglobose (adapted from Sivanesan 1987).
Culture characteristics:
Colonies on PDA white or pale grey when young, orange to brown or different shades of grey (mainly dark olivaceous grey) when mature, fluffy, cottony, raised or convex with papillate surface, margin lobate, undulate, entire or sometimes rhizoid.
Optimal media and cultivation conditions: Sterilised Zea mays leaves placed on 1.5 % WA or slide cultures of half-strength PDA under near-ultraviolet light (12 h light, 12 h dark) to induce sporulation of the asexual morph, while for the sexual morph Sach's agar with sterilised rice or wheat straw at 25 °C is used.
Distribution:
Worldwide.
Hosts:
Wide host range, occurring as pathogens, saprobes or endophytes. Mainly found on members of the Poaceae, being pathogens of grass and staple crops, including rice, maize, wheat and sorghum. This genus also occurs on genera belonging to Actinidiaceae, Aizoaceae, Caricaceae, Convolvulaceae, Fabaceae, Iridaceae, Lamiaceae, Lythraceae, Oleaceae, Polygonaceae and Rubiaceae.
Disease symptoms:
Leaf spots, leaf blight, melting out, root rot, foot rot, among others.
Notes:
Species delimitation in Curvularia based on morphology only is difficult due to the morphological complexity within this genus, as also observed in Bipolaris. Furthermore, the distinction of both genera based on morphology alone is sometimes complicated (see Bipolaris notes for morphological differences between Bipolaris and Curvularia). Therefore, molecular data are essential for an accurate identification of species within these genera, ITS, gapdh and tef1 being the loci selected for this purpose (Manamgoda et al. 2014, 2015).
Curvularia is a rich genus in host range and geographic distribution compared to Bipolaris. Apart from phytopathogenic species, this genus comprises species that are pathogens of humans and other animals, causing respiratory tract, cutaneous, cerebral and corneal infections, mainly in immunocompromised patients (Carter & Boudreaux 2004). Some species can be found in association with both humans and plants, such as Cu. hawaiiensis, Cu. lunata and Cu. spicifera (Manamgoda et al. 2015).
References:
- Sivanesan 1987 (morphology and pathogenicity); Manamgoda et al. 2011 (pathogenicity), Manamgoda et al. 2015 (morphology, pathogenicity and phylogeny).
- Carter E, Boudreaux C (2004). Fatal cerebral phaeohyphomycosis due to Curvularia lunata in an immunocompetent patient. Journal of Clinical Microbiology 42: 5419–5423.
- Manamgoda DS, Cai L, Bahkali AH, et al. (2011). Cochliobolus: an overview and current status of species. Fungal Diversity 51: 3–42.
- Manamgoda DS, Rossman AY, Castlebury LA, et al. (2014). The genus Bipolaris. Studies in Mycology 79: 221–288.
- Manamgoda DS, Rossman AY, Castlebury LA, et al. (2015). A taxonomic and phylogenetic re-appraisal of the genus Curvularia (Pleosporaceae): human and plant pathogens. Phytotaxa 212: 175–198.
- Sivanesan A (1987). Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs. Mycological Papers 158: 1–261.
Table 9. DNA barcodes of accepted Curvularia spp.
Species |
Isolates1 |
|
GenBank accession numbers2 |
|
References |
|||
|
|
ITS |
gapdh |
tef1 |
|
|
||
Cu. aeria |
CBS 294.61T |
HE861850 |
HF565450 |
- |
da Cunha et al. (2013) |
|
||
Cu. affinis |
CBS 154.34SynT |
KJ909780 |
KM230401 |
KM196566 |
Manamgoda et al. (2015) |
|
||
Cu. akaii |
CBS 317.86 |
KJ909782 |
KM230402 |
KM196569 |
Manamgoda et al. (2015) |
|
||
Cu. akaiiensis |
BRIP 16080IsoT |
KJ415539 |
KJ415407 |
KJ415453 |
Tan et al. (2014) |
|
||
Cu. alcornii |
MFLUCC 10-0703T |
JX256420 |
JX276433 |
JX266589 |
Manamgoda et al. (2012a) |
|
||
Cu. americana |
UTHSC 08-3414T |
HE861833 |
HF565488 |
- |
Madrid et al. (2014) |
|
||
Cu. asiatica |
MFLUCC 10-0711T |
JX256424 |
JX276436 |
JX266593 |
Manamgoda et al. (2012a) |
|
||
Cu. australiensis |
BRIP 12044T |
KJ415540 |
KJ415406 |
KJ415452 |
Tan et al. (2014) |
|
||
Cu. australis |
BRIP 12521T |
KJ415541 |
KJ415405 |
KJ415451 |
Tan et al. (2014) |
|
||
Cu. bannonii |
BRIP 16732IsoT |
KJ415542 |
KJ415404 |
KJ415450 |
Tan et al. (2014) |
|
||
Cu. borreriae |
CBS 859.73 |
HE861848 |
HF565455 |
- |
da Cunha et al. (2013) |
|
||
Cu. bothriochloae |
BRIP 12522T |
KJ415543 |
KJ415403 |
KJ415449 |
Tan et al. (2014) |
|
||
Cu. brachyspora |
CBS 186.50 |
KJ922372 |
KM061784 |
KM230405 |
Manamgoda et al. (2014) |
|
||
Cu. buchloës |
CBS 246.49T |
KJ909765 |
KM061789 |
KM196588 |
Manamgoda et al. (2014) |
|
||
Cu. carica-papayae |
CBS 135941T |
HG778984 |
HG779146 |
- |
Madrid et al. (2014) |
|
||
Cu. chlamydospora |
UTHSC 07-2764T |
HG779021 |
HG779151 |
- |
Madrid et al. (2014) |
|
||
Cu. clavata |
BRIP 61680b |
KU552205 |
KU552167 |
KU552159 |
Khemmuk et al. (2016) |
|
||
Cu. coicis |
CBS 192.29SynT |
JN192373 |
JN600962 |
JN601006 |
Manamgoda et al. (2015) |
|
||
Cu. crustacea |
BRIP 13524ET |
KJ415544 |
KJ415402 |
KJ415448 |
Tan et al. (2014) |
|
||
Cu. cymbopogonis |
CBS 419.78 |
HG778985 |
HG779129 |
HG779163 |
Madrid et al. (2014) |
|
||
Cu. dactyloctenii |
BRIP 12846T |
KJ415545 |
KJ415401 |
KJ415447 |
Tan et al. (2014) |
|
||
Cu. ellisii |
CBS 193.62T |
JN192375 |
JN600963 |
JN601007 |
Manamgoda et al. (2011) |
|
||
Cu. eragrostidis |
CBS 189.48 |
HG778986 |
HG779154 |
HG779164 |
Madrid et al. (2014) |
|
||
Cu. geniculata |
CBS 187.50 |
KJ909781 |
KM083609 |
KM230410 |
Manamgoda et al. (2015) |
|
||
Cu. gladioli |
CBS 210.79 |
HG778987 |
HG779123 |
- |
Madrid et al. (2014) |
|
||
Cu. graminicola |
BRIP 23186T |
JN192376 |
JN600964 |
JN601008 |
Manamgoda et al. (2012b) |
|
||
Cu. gudauskasii |
DAOM 165085 |
AF071338 |
- |
- |
Berbee et al. (1999) |
|
||
Cu. harveyi |
BRIP 57412IsoT |
KJ415546 |
KJ415400 |
KJ415446 |
Tan et al. (2014) |
|
||
Cu. hawaiiensis |
BRIP 11987IsoLT |
KJ415547 |
KJ415399 |
KJ415445 |
Tan et al. (2014) |
|
||
Cu. heteropogonicola |
BRIP 14579IsoT |
KJ415548 |
KJ415398 |
KJ415444 |
Tan et al. (2014) |
|
||
Cu. heteropogonis |
CBS 284.91T |
JN192379 |
JN600969 |
JN601013 |
Manamgoda et al. (2012b) |
|
||
Cu. hominis |
CBS 136985T |
HG779011 |
HG779106 |
- |
Madrid et al. (2014) |
|
||
Cu. homomorpha |
CBS 156.60T |
JN192380 |
JN600970 |
JN601014 |
Manamgoda et al. (2014) |
|
||
Cu. inaequalis |
CBS 102.42T |
KJ922375 |
KM061787 |
KM196574 |
Manamgoda et al. (2014) |
|
||
Cu. intermedia |
CBS 334.64 |
HG778991 |
HG779155 |
HG779169 |
Madrid et al. (2014) |
|
||
Cu. ischaemi |
CBS 630.82T |
JX256428 |
JX276440 |
- |
Manamgoda et al. (2012b) |
|
||
Cu. kusanoi |
CBS 137.29 |
JN192381 |
- |
JN601016 |
Manamgoda et al. (2015) |
|
||
Cu. lunata |
CBS 730.96NT |
JX256429 |
JX276441 |
JX266596 |
Manamgoda et al. (2012b) |
|
||
Cu. malina |
CBS 131274T |
JF812154 |
KP153179 |
KR493095 |
Tomaso-Peterson et al. (2016) |
|
||
Cu. miyakei |
CBS 197.29SynT |
KJ909770 |
KM083611 |
KM196568 |
Manamgoda et al. (2014) |
|
||
Cu. muehlenbeckiae |
CBS 144.63T |
HG779002 |
HG779108 |
- |
Madrid et al. (2014) |
|
||
Cu. neergaardii |
BRIP 12919IsoT |
KJ415550 |
KJ415397 |
KJ415443 |
Tan et al. (2014) |
|
||
Cu. neoindica |
BRIP 17439 |
AF081449 |
AF081406 |
- |
Berbee et al. (1999) |
|
||
Cu. nicotiae |
CBS 655.74IsoT = BRIP 11983 |
KJ415551 |
KJ415396 |
KJ415442 |
Tan et al. (2014) |
|
||
Cu. nodulosa |
CBS 160.58 |
JN601033 |
JN600975 |
JN601019 |
Manamgoda et al. (2015) |
|
||
Cu. oryzae |
CBS 169.53IsoT |
KP400650 |
KP645344 |
KM196590 |
Manamgoda et al. (2015) |
|
||
Cu. ovariicola |
CBS 470.90T |
JN192384 |
JN600976 |
JN601020 |
Manamgoda et al. (2012b) |
|
||
Cu. papendorfii |
CBS 308.67T |
KJ909774 |
KM083617 |
KM196594 |
Manamgoda et al. (2014) |
|
||
Cu. pallescens |
CBS 156.35T |
KJ922380 |
KM083606 |
KM196570 |
Manamgoda et al. (2015) |
|
||
Cu. perotidis |
CBS 350.90T |
JN192385 |
KJ415394 |
JN601021 |
Manamgoda et al. (2015) |
|
||
Cu. pisi |
CBS 190.48T |
KY905678 |
KY905690 |
KY905697 |
Marin-Felix et al. (2017) |
|
||
Cu. portulacae |
CBS 239.48IsoT = |
KJ909775 |
KM083616 |
- |
Manamgoda et al. (2014) |
|
||
|
BRIP 14541IsoT |
KJ415553 |
KJ415393 |
KJ415440 |
Tan et al. (2014) |
|
||
Cu. prasadii |
CBS 143.64T |
KJ922373 |
KM061785 |
KM230408 |
Manamgoda et al. (2014) |
|
||
Cu. protuberata |
CBS 376.65IsoT |
KJ922376 |
KM083605 |
KM196576 |
Manamgoda et al. (2014) |
|
||
Cu. pseudolunata |
UTHSC 09-2092T |
HE861842 |
HF565459 |
- |
da Cunha et al. (2013) |
|
||
Cu. pseudorobusta |
UTHSC 08-3458 |
HE861838 |
HF565476 |
- |
da Cunha et al. (2013) |
|
||
Cu. ravenelii |
BRIP 13165T |
JN192386 |
JN600978 |
JN601024 |
Manamgoda et al. (2012b) |
|
||
Cu. richardiae |
BRIP 4371IsoLT |
KJ415555 |
KJ415391 |
KJ415438 |
Tan et al. (2014) |
|
||
Cu. robusta |
CBS 624.68IsoT |
KJ909783 |
KM083613 |
KM196577 |
Manamgoda et al. (2014) |
|
||
Cu. ryleyi |
BRIP 12554T |
KJ415556 |
KJ415390 |
KJ415437 |
Tan et al. (2014) |
|
||
Cu. senegalensis |
CBS 149.71 |
HG779001 |
HG779128 |
- |
Madrid et al. (2014) |
|
||
Cu. sesuvi |
Bp-Zj 01 |
EF175940 |
- |
- |
Zhang & Li (2009) |
|
||
Cu. soli |
CBS 222.96T |
KY905679 |
KY905691 |
KY905698 |
Marin-Felix et al. (2017) |
|
||
Cu. sorghina |
BRIP 15900IsoT |
KJ415558 |
KJ415388 |
KJ415435 |
Tan et al. (2014) |
|
||
Cu. spicifera |
CBS 274.52 |
JN192387 |
JN600979 |
JN601023 |
Manamgoda et al. (2012b) |
|
||
Cu. subpapendorfii |
CBS 656.74T |
KJ909777 |
KM061791 |
KM196585 |
Manamgoda et al. (2015) |
|
||
Cu. trifolii |
CBS 173.55 |
HG779023 |
HG779124 |
- |
Madrid et al. (2014) |
|
||
Cu. tripogonis |
BRIP 12375T |
JN192388 |
JN600980 |
JN601025 |
Manamgoda et al. (2011) |
|
||
Cu. tropicalis |
BRIP 14834IsoT |
KJ415559 |
KJ415387 |
KJ415434 |
Tan et al. (2014) |
|
||
Cu. tsudae |
ATCC 44764PT |
KC424596 |
KC747745 |
KC503940 |
Deng et al. (2014) |
|
||
Cu. tuberculata |
CBS 146.63IsoT |
JX256433 |
JX276445 |
JX266599 |
Manamgoda et al. (2012b) |
|
||
Cu. uncinata |
CBS 221.52T |
HG779024 |
HG779134 |
- |
Madrid et al. (2014) |
|
||
Cu. verruciformis |
CBS 537.75 |
HG779026 |
HG779133 |
HG779211 |
Madrid et al. (2014) |
|
||
Cu. verruculosa |
CBS 150.63 |
KP400652 |
KP645346 |
KP735695 |
Manamgoda et al. (2015) |
|
||
1ATCC: American Type Culture Collection, Virginia, USA; BRIP: Queensland Plant Pathology Herbarium, Brisbane, Australia; Bp-Zj: Isolate housed in Biotechnology Institute, Zhejiang University, Hangzhou, China; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; DAOM: Plant Research Institute, Department of Agriculture (Mycology), Ottawa, Canada; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Ria, Thailand, UTHSC: Fungus Testing Laboratory, Department of Pathology at the University of Texas Health Science Center, San Antonio, Texas, USA. ET and IsoT, IsoLT, PT, SynT, T indicate ex-epitype, ex-isotype, ex-isolectotype, ex-paratype, ex-syntype and ex-type strains, respectively.
2ITS: internal transcribed spacers and intervening 5.8S nrDNA; gapdh: partial glyceraldehyde-3-phosphate dehydrogenase gene; tef1: partial translation elongation factor 1-alpha gene.
- Berbee ML, Pirseyedi M, Hubbard S (1999). Cochliobolus phylogenetics and the origin of known, highly virulent pathogens, inferred from ITS and glyceraldehyde-3-phosphate dehydrogenase gene sequences. Mycologia 91: 964–977.
- da Cunha KC, Sutton DA, Fothergill AW, et al. (2013). In vitro antifungal susceptibility and molecular identity of 99 clinical isolates of the opportunistic fungal genus Curvularia. Diagnostic Microbiology and Infectious Disease 76: 168–174.
- Deng H, Tan YP, Shivas RG, et al. (2014). Curvularia tsudae comb. nov. et nom. nov., formerly Pseudocochliobolus australiensis, and a revised synonymy for Curvularia australiensis. Mycoscience 56: 24–28.
- Khemmuk W, Shivas RG, Henry RJ, et al. (2016). Fungi associated with foliar diseases of wild and cultivated rice (Oryza spp.) in northern Queensland. Australasian Plant Pathology 45: 297–308.
- Madrid H, da Cunha KC, Gene J, et al. (2014). Novel Curvularia species from clinical specimens. Persoonia 33: 48–60.
- Manamgoda DS, Cai L, Bahkali AH, et al. (2011). Cochliobolus: an overview and current status of species. Fungal Diversity 51: 3–42.
- Manamgoda DS, Cai L, Hyde KD (2012a). Two new species of Curvularia from nothern Thailand. Sydowia 64: 255–266.
- Manamgoda DS, Cai L, Hyde KD (2012b). A taxonomic evaluation of the holomorphic species complex: Cochliobolus, Bipolaris and Curvularia through multilocus phylogeny. Fungal Diversity 56: 131–144.
- Manamgoda DS, Rossman AY, Castlebury LA, et al. (2014). The genus Bipolaris. Studies in Mycology 79: 221–288.
- Manamgoda DS, Rossman AY, Castlebury LA, et al. (2015). A taxonomic and phylogenetic re-appraisal of the genus Curvularia (Pleosporaceae): human and plant pathogens. Phytotaxa 212: 175–198.
- Manamgoda DS, Udayanga D, Cai L, et al. (2013). Endophytic Colletotrichum from tropical grasses with a new species C. endophytica. Fungal Diversity 61: 107–115.
- Marin-Felix Y, Groenewald JZ, Cai, L, et al. (2017). Genera of phytopathogenic fungi: GOPHY 1. Studies in Mycology xxxx.
- Tan YP, Madrid H, Crous PW, et al. (2014). Johnalcornia gen. et. comb. nov., and nine new combinations in Curvularia based on molecular phylogenetic analysis. Australasian Plant Pathology 43: 589–603.
- Zhang JZ, Li MJ (2009). A new species of Bipolaris from the halophyte Sesuvium portulacastrum in Guangdong Province, China. Mycotaxon 109: 289–300.