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Calonectria Species and Their Cylindrocladium Anamorphs: Species with Clavate Vesicles

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Calonectria Species and Their Cylindrocladium Anamorphs: Species with Clavate Vesicles STUDIES IN MYCOLOGY 55: 213–226. 2006. Calonectria species and their Cylindrocladium anamorphs: species with clavate vesicles Pedro W. Crous1*, Johannes Z. Groenewald1, Jean-Michel Risède2, Philippe Simoneau3 and Kevin D. Hyde4 1Centraalbureau voor Schimmelcultures, Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; 2CIRAD-FLHOR, Station de Neufchâteau, 97130 Capesterre Belle Eau, Guadeloupe, French West Indies; 3UMR PaVé N°77- Faculté des Sciences, Université d’Angers 2, Bd Lavoisier 49045 Angers cedex, France; 4Centre for Research in Fungal Diversity, Department of Ecology & Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong, SAR China *Correspondence: Pedro W. Crous, [email protected] Abstract: The present study compares all known species of Cylindrocladium that have clavate vesicles. Several isolates were obtained from baited soils collected in various parts of the world, while others were associated with leaf litter or symptomatic plant hosts. Isolates were compared based on morphology, as well as DNA sequence data from their β-tubulin and histone gene H3 regions. Cylindrocladium australiense and Cy. ecuadoriae, are described as new species, a decision based on morphology and molecular data. A group of isolates associated with toppling disease of banana in the West Indies is identified as Cy. flexuosum. An epitype is designated for Cy. ilicicola, and a new name, Curvicladiella, proposed to replace the anamorphic genus Curvicladium, which is a homonym. Taxonomic novelties: Cylindrocladium australiense Crous & K.D. Hyde sp. nov., Cylindrocladium ecuadoriae Crous & M.J. Wingf. sp. nov., Curvicladiella Decock & Crous nom. nov., Curvicladiella cignea (Decock & Crous) Decock & Crous comb. nov. Key words: Ascomycetes, Calonectria, Cylindrocladium, Hypocreales, leaf spots, soil fungi, systematics. INTRODUCTION MATERIALS AND METHODS Members of the genus Calonectria De Not. (Ca.) Isolates (Nectriaceae, Hypocreales, Ascomycetes) and Isolates were obtained from plant hosts, or baited their Cylindrocladium Morgan (Cy.) anamorphs are from soil as explained in Crous (2002). Cultural commonly associated with a wide range of plant disease characteristics and morphology were determined on symptoms (Crous 2002). The current paper represents plates containing 2 % malt extract agar (MEA) (20 g/ the second in a series assessing the taxonomy of L), and carnation leaf agar (CLA) [1 % water agar (10 species of Cylindrocladium, by integrating morphology g/L) with autoclaved carnation leaves placed onto the with DNA sequence data and sexual compatibility medium] in the other (Gams et al. 1998). Plates were studies (Crous et al. 2004). incubated for 7 d at 25 °C under continuous near-UV Cylindrocladium species with clavate vesicles are light, to promote sporulation. well-known pathogens from a wide range of hosts in most subtropical to tropical countries (Crous & Wingfield DNA phylogeny 1992, Crous et al. 1995, 1997, 1999, 2000, Kang et al. Genomic DNA was isolated from fungal mycelia 2001, Crous 2002). In the current study, we obtained collected from the plates using the isolation protocol numerous isolates of Cylindrocladium from baited soils collected in tropical areas. Further Cylindrocladium of Lee & Taylor (1990). Two loci were amplified and isolates were obtained from a biotic complex sequenced as explained in Crous et al. (2004), namely, including root rot fungi and plant-parasitic nematodes part of the β-tubulin gene, amplified with primers T1 associated with toppling disease of banana (Risède (O’Donnell & Cigelnik 1997) and CYLTUB1R (Crous et & Simoneau 2001). Previous studies have shown al. 2004); and part of the histone H3 gene using primers that isolates resembling Cy. gracile (Bugn.) Boesew. CYLH3F and CYLH3R (Crous et al. 2004). were pathogenic to banana, and associated with stem The sequences generated in this study were added lesions, root breakage and toppling disease (Risède & to other sequences obtained from GenBank (http://www. Simoneau 2001, 2004). The aim of the present study ncbi.nlm.nih.gov) and TreeBASE (http://www.treebase. was to analyze all available Cylindrocladium strains with org) and the alignment was assembled using Sequence clavate vesicles using morphology and DNA sequence Alignment Editor v. 2.0a11 (Rambaut 2002) with manual analysis of their β-tubulin and histone H3 gene regions adjustments for improvement made visually where in order to resolve the status of Cylindrocladium species necessary. Sequences for Cylindrocladiella peruviana with clavate vesicles. A further aim was to identify the (Bat., J.L. Bezerra & M.M.P. Herrera) Boesew. and Cylindrocladium sp. associated with toppling disease Cylindrocladiella lageniformis Crous, M.J. Wingf. & of banana. Alfenas were added to the alignments as outgroups. 213 CROUS ET AL. Cylindrocladiella peruviana AY725653 Cylindrocladiella lageniformis AY725652 89 Cy. penicilloides CBS 174.55 100 CPC 765 CPC 763 Cy. curvisporum 98 FTCC 1003 CBS 114551 Cy. hurae 98 CBS 114182 100 79 FTCC 1002 100 CBS 114529 99 UFV 215 Cy. rumohrae 54 CBS 111431 CBS 114544 100 CBS 112133 CBS 114767 Cy. angustatum 91 CBS 114766 Ca. leguminum CBS 728.68 CBS 115674 100 CBS 111141 Cy. graciloideum CBS 111040 CBS 114175 100 CBS 111399 67 CBS 111705 Cy. theae 66 CBS 114684 ATCC 48895 Cy. avesiculatum ATCC 38226 100 CBS 114454 CBS 111284 Cy. pseudogracile AR2677 CBS 111383 88 CBS 111425 CBS 111406 Cy. ecuadoriae 80 CBS 111412 CBS 111393 64 CBS 111394 CBS 114557 CBS 114666 100 CPC 11347 CPC 11352 CPC 11350 Cy. flexuosum CBS 112676 63 CBS 112675 CPC 11349 CPC 11351 CPC 11348 CBS 111382 62 CBS 112694 CBS 111475 64 CBS 110676 62 CBS 111869 63 CBS 112673 CBS 114167 CBS 111478 91 CBS 111458 CBS 111465 CBS 115769 Cy. gracile CBS 111456 CBS 111468 CBS 115680 63 CBS 111129 CBS 111474 Cy. clavatum CBS 111776 CBS 111476 CBS 111390 CBS 114171 Cy. hawksworthii CBS 111870 Cy. gordoniae CBS 112142 CBS 111793 CPC 12114 74 CPC 12072 CPC 11619 99 CPC 11618 CPC 11286 CPC 11284 CBS 111867 Cy. pteridis CBS 111871 CBS 111778 60 CPC 2869 CPC 2190 CPC 11285 CPC 11435 86 CPC 11569 CPC 2322 93 CBS 114572 CBS 114571 Cy. madagascariense 97 CBS 114539 100 CBS 114880 Cy. macroconidiale CPC 413 CBS 114660 99 CBS 114704 CBS 114695 96 CBS 112948 68 CBS 111367 CBS 112951 CBS 111205 Cy. colhounii CBS 293.79 CPC 705 CPC 681 68 CPC 1237 CBS 114036 64 CBS 112739 Cy. australiense CBS 112954 Cylindrocladium sp. CBS 112957 100 CBS 112682 Cy. multiseptatum 99 CPC 1602 CBS 114812 100 CPC 10898 Cy. acicola 77 CBS 114813 CBS 113925 CBS 112955 80 83 CBS 112634 CBS 112151 71 97 CBS 112155 Fig. 1. One of 657 most parsimonious trees CBS 112149 CBS 112146 obtained from a heuristic search with 100 CBS 112144 Ca. reteaudii CBS 112153 random taxon additions of the β-tubulin sequence 73 CBS 112147 alignment. The scale bar shows 10 changes and CBS 582.50 CPC 3217 bootstrap support values from 1000 replicates CBS 113582 10 changes CBS 113583 are shown at the nodes. Thickened lines indicate CBS 112143 branches present in the strict consensus tree. The tree was rooted to two Cylindrocladiella species. 214 CALONECTRIA SPECIES AND THEIR CYLINDROCLADIUM ANAMORPHS Cylindrocladiella peruviana AY725700 Cylindrocladiella lageniformis AY725699 Cy. avesiculatum ATCC 38226 92 CBS 114529 100 UFV 215 Cy. rumohrae CBS 111431 CBS 114551 93 100 CBS 114182 FTCC 1002 Cy. hurae 53 FTCC 1003 Ca. leguminum CBS 728.68 65 CBS 114544 100 CBS 112133 CBS 114767 Cy. angustatum CBS 114766 Cy. hawksworthii CBS 111870 CBS 115674 100 CBS 111141 Cy. graciloideum CBS 111040 Cy. gordoniae CBS 112142 88 CBS 111793 CPC 11286 97 CPC 11284 CPC 11285 CPC 2190 54 CPC 11569 CPC 11435 CBS 111778 Cy. pteridis 56 CBS 111867 CBS 111871 CPC 12114 CPC 12072 62 CPC 11619 CPC 11618 CBS 111383 98 CBS 111425 CBS 111393 92 CBS 111394 Cy. ecuadoriae 57 CBS 111412 62 CBS 111406 CBS 114454 100 CBS 111284 Cy. pseudogracile AR2677 CBS 114557 CBS 114666 99 CPC 11347 CPC 11352 CPC 11350 CBS 112676 Cy. flexuosum 54 CBS 112675 CPC 11349 60 CPC 11351 CPC 11348 Cy. clavatum CBS 111776 CBS 111475 CBS 111478 CBS 111458 98 CBS 111465 CBS 114167 CBS 111456 CBS 111468 CBS 115680 CBS 111129 Cy. gracile 53 CBS 111474 CBS 111476 CBS 111390 CBS 114171 CBS 115769 CBS 112673 CBS 111869 CBS 112694 CBS 110676 CBS 111399 95 CBS 111705 CBS 114684 Cy. theae 98 CBS 114175 100 CBS 114880 Cy. macroconidiale CPC 413 100 CPC 765 CPC 763 Cy. curvisporum CBS 114036 72 CPC 705 CPC 1237 CBS 112739 100 CBS 112951 CPC 681 CBS 293.79 Cy. colhounii 52 CBS 112948 CBS 111367 81 CBS 114660 CBS 114704 CBS 114695 100 CBS 114572 Cy. madagascariense CBS 114571 Cy. australiense CBS 112954 Cylindrocladium sp. CBS 112957 Cy. multiseptatum CBS 112682 100 CBS 114812 Cy. acicola 88 CPC 10898 70 CBS 114813 CBS 112955 CBS 112634 85 CBS 112149 94 CBS 112146 CBS 112155 72 CBS 112151 CBS 113925 CPC 3217 Ca. reteaudii CBS 582.50 100 CBS 113582 CBS 113583 CBS 112143 CBS 112147 CBS 112153 10 changes CBS 112144 Fig. 2. One of four most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the histone H3 sequence alignment. The scale bar shows 10 changes and bootstrap support values from 1000 replicates are shown at the nodes. Thickened lines indicate branches present in the strict consensus tree. The tree was rooted to two Cylindrocladiella species. 215 C 216 Table 1. Isolates of Cylindrocladium (Calonectria) species studied. ROUS 1,2 3 3 Species Isolate number β-tubulin Histone H3 Host Country Collector ET Ca. avesiculata (Cy. avesiculatum) ATCC 38226; CPC 2373T AF333392 DQ190620 Ilex vomitoria U.S.A. S.A. Alfieri AL . Ca. clavata (Cy. flexuosum) CBS 112675; SLU2 DQ190547 DQ190621 Musa sp. Saint Lucia J.M. Risède CBS 112676; Gua9 DQ190548 DQ190622 Musa sp. Guadeloupe J.M.
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