StudieS in Mycology 68: 57–78. 2011.
available online at www.studiesinmycology.org
doi:10.3114/sim.2011.68.03
Delimitation of Neonectria and Cylindrocarpon (Nectriaceae, Hypocreales,
Ascomycota) and related genera with Cylindrocarpon-like anamorphs
P. Chaverri1*, C. Salgado1, Y. Hirooka1, 2, A.Y. Rossman2 and G.J. Samuels2
University of Maryland, Department of Plant Sciences and Landscape Architecture, 2112 Plant Sciences Building, College Park, Maryland 20742, USA; 2United States
Department of Agriculture, Agriculture Research Service, Systematic Mycology and Microbiology Laboratory, Rm. 240, B-010A, 10300 Beltsville Avenue, Beltsville, Maryland
20705, USA
1
*Correspondence: Priscila Chaverri, pchaverr@umd.edu
Abstract: Neonectria is a cosmopolitan genus and it is, in part, deined by its link to the anamorph genus Cylindrocarpon. Neonectria has been divided into informal groups on
the basis of combined morphology of anamorph and teleomorph. Previously, Cylindrocarpon was divided into four groups deined by presence or absence of microconidia and
chlamydospores. Molecular phylogenetic analyses have indicated that Neonectria sensu stricto and Cylindrocarpon sensu stricto are phylogenetically congeneric. In addition,
morphological and molecular data accumulated over several years have indicated that Neonectria sensu lato and Cylindrocarpon sensu lato do not form a monophyletic
group and that the respective informal groups may represent distinct genera. In the present work, a multilocus analysis (act, ITS, LSU, rpb1, tef1, tub) was applied to
representatives of the informal groups to determine their level of phylogenetic support as a irst step towards taxonomic revision of Neonectria sensu lato. Results show ive
distinct highly supported clades that correspond to some extent with the informal Neonectria and Cylindrocarpon groups that are here recognised as genera: (1) N. coccineagroup and Cylindrocarpon groups 1 & 4 (Neonectria/Cylindrocarpon sensu stricto); (2) N. rugulosa-group (Rugonectria gen. nov.); (3) N. mammoidea/N. veuillotiana-groups
and Cylindrocarpon group 2 (Thelonectria gen. nov.); (4) N. radicicola-group and Cylindrocarpon group 3 (Ilyonectria gen. nov.); and (5) anamorph genus Campylocarpon.
Characteristics of the anamorphs and teleomorphs correlate with the ive genera, three of which are newly described. New combinations are made for species where their
classiication is conirmed by phylogenetic data.
Key words: Canker-causing fungi, molecular systematics, Nectria-like fungi, phylogeny, polyphasic taxonomy, root-rotting fungi, sequence analysis, systematics, taxonomy.
Taxonomic novelties: Ilyonectria P. Chaverri & C. Salgado, gen. nov.; Ilyonectria coprosmae (Dingley) P. Chaverri & C. Salgado, comb. nov.; Ilyonectria liriodendri (Halleen
et al.) P. Chaverri & C. Salgado, comb. nov.; Ilyonectria macrodydima (Halleen, Schroers & Crous) P. Chaverri & C. Salgado, comb. nov.; Ilyonectria radicicola (Gerlach & L.
Nilsoon) P. Chaverri & C. Salgado, comb. nov.; Rugonectria P. Chaverri & Samuels, gen. nov.; Rugonectria castaneicola (W. Yamam. & Oyasu) Hirooka & P. Chaverri, comb.
nov; Rugonectria neobalansae (Samuels) P. Chaverri & Samuels, comb. nov.; Rugonectria rugulosa (Pat. & Gaill.) Samuels, P. Chaverri & C. Salgado, comb. nov.; Thelonectria
P. Chaverri & C. Salgado, gen. nov.; Thelonectria coronata (Penz. & Sacc.) P. Chaverri & C. Salgado, comb. nov.; Thelonectria discophora (Mont.) P. Chaverri & C. Salgado,
comb. nov.; Thelonectria jungneri (Henn.) P. Chaverri & C. Salgado, comb. nov.; Thelonectria lucida (Höhnel) P. Chaverri & C. Salgado, comb. nov.; Thelonectria olida (Wollenw.)
P. Chaverri & C. Salgado, comb. nov; Thelonectria trachosa (Samuels & Brayford) Samuels, P. Chaverri & C. Salgado, comb. nov.; Thelonectria veuillotiana (Sacc. & Roum.)
P. Chaverri & C. Salgado, comb. nov.; Thelonectria viridispora (Samuels & Brayford) P. Chaverri, C. Salgado, & Samuels, comb. nov.; Thelonectria westlandica (Dingley) P.
Chaverri & C. Salgado, comb. nov.
INTRODUCTION
Species of Neonectria sensu lato and their anamorphs in
Cylindrocarpon are common in tropical and temperate regions.
They are generally found on bark of recently killed woody plants
and sometimes on decaying herbaceous material (Samuels
1988, Samuels & Brayford 1990, Samuels et al. 1990, Samuels &
Brayford 1993, 1994, Rossman et al. 1999, Castlebury et al. 2006).
Some species of this genus are plant pathogens causing cankers,
root rots, and other diseases on hardwood and coniferous trees,
e.g. Abies and Acer cankers caused by Neonectria castaneicola;
beech (Fagus) bark disease caused by N. coccinea, N. ditissima
and N. faginata; black foot disease of grapevines (Vitis) caused
by N. liriodendri; root rots caused by N. radicicola; and cankers
caused by N. rugulosa, among others (Samuels & Brayford 1994,
Hirooka et al. 2005, Kobayashi et al. 2005, Castlebury et al.
2006, Halleen et al. 2006). According to Index Fungorum (www.
indexfungorum.org), 38 species have been placed in Neonectria
and 143 in Cylindrocarpon. These numbers are underestimated
because several species of Nectria-like fungi with Cylindrocarpon
anamorphs have not been transferred to Neonectria (> 20 spp.). To
date, the most comprehensive taxonomic works of Neonectria and
species of Nectria having Cylindrocarpon anamorphs are those
by Booth (1959, 1966 ) and Samuels, Brayford and collaborators
(Samuels 1988, Brayford & Samuels 1993, Samuels & Brayford
1993, 1994, Brayford et al. 2004).
Species of Neonectria sensu lato are characterised by having
perithecia that are subglobose to broadly obpyriform, smooth to
roughened, red, becoming dark red in 3 % potassium hydroxide
(KOH), and with an acute to constricted apex that is sometimes
knobby; the perithecial wall is ca. 50 μm thick and generally
composed of two regions, sometimes with an outer region that forms
textura epidermoidea, that may or may not be covered with another
region of cells; and the ascospores are hyaline, generally bicellular,
rarely multi-cellular, and smooth or inely ornamented (Rossman et
al. 1999). The anamorph of N. ramulariae (type of Neonectria) is
Cylindrocarpon obtusiusculum and, consequently, species with the
Neonectria-like morphology described above and Cylindrocarpon
anamorphs have been classiied as Neonectria. However, species
that have been placed in Neonectria and species of Nectria having
a Cylindrocarpon anamorph vary greatly in the morphology of
their perithecia, some having perithecial walls < 50 μm or > 50 μm
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57
chaverri et al.
thick, others warted, and others with various degrees of ascospore
ornamentation. Some species of Neonectria are morphologically
similar in perithecial morphology with differences seen only in the
anamorph (Samuels et al. 2006b).
The morphological variation in Neonectria resulted in the
subdivision of species into ive informal groups, mostly based
on perithecial characteristics: (1) N. coccinea/galligena-group
(Neonectria sensu stricto) (Booth 1959); (2) N. mammoidea-group
(N. mammoidea = N. discophora (Booth 1959); (3) N. rugulosagroup (Samuels & Brayford 1994); (4) N. radicicola-group (Booth
1959); and (5) N. veuillotiana-group (Brayford & Samuels 1993).
Species in the N. coccinea/galligena-group are characterised by
having few to numerous perithecia clustered on wood; perithecial
walls are ca. 50 μm thick, composed of relatively thick-walled, small
cells; and ascospores are generally smooth (Booth 1959). Species
in the N. mammoidea-group were originally deined as having a
distinctive perithecial wall that comprises a layer of hyphae that
have thickened walls and are typically arranged radially, giving
the appearance of a palisade (Booth 1959, Brayford et al. 2004).
This characteristic generally results in smooth, shiny perithecia. In
addition to the perithecial anatomy, the N. mammoidea-group has
spinulose, often yellow-brown ascospores, and a non-microconidial
anamorph. The N. rugulosa-group includes species with warted
perithecia, a perithecial wall > 50 μm thick, composed of large,
thick-walled cells, and striate ascospores (Samuels & Brayford
1994). The N. radicicola-group includes species that have smooth
to slightly warted, usually solitary perithecia, the outer region
of the perithecial wall composed of large, thin-walled cells, and
smooth ascospores (Samuels & Brayford 1990). Species in the N.
veuillotiana-group have perithecia with a lattened or knobby apex,
perithecial walls composed of thick-walled cells, and tuberculate
ascospores (Brayford & Samuels 1993). Mantiri et al. (2001)
revised the informal groupings of Neonectria based on phylogenetic
analyses of DNA sequence data. Group or clade I was Neonectria
sensu stricto or the N. coccinea/galligena-group; clade II included
the N. mammoidea-, N. rugulosa-, and N. veuillotiana-groups; and
clade III was the N. radicicola-group.
Booth (1966) subdivided Cylindrocarpon into four groups based
on the presence or absence of microconidia and chlamydospores.
The irst three Cylindrocarpon groups in Booth (1966) correlate
with the three groups/clades in Mantiri et al. 2001 (Castlebury et
al. 2006). Anamorphs in the N. coccinea/galligena-group (clade
I in Mantiri et al. 2001) belong to Cylindrocarpon group 1, which
have micro- and macroconidia but lack chlamydospores, except N.
ramulariae/C. obtusiusculum, which has chlamydospores and lacks
microconidia. Cylindrocarpon obtusiusculum was originally placed
in Cylindrocarpon group 4 by Booth (1966). The type species of
Cylindrocarpon, C. cylindroides, belongs in Cylindrocarpon group
1 (Booth 1966, Mantiri et al. 2001, Brayford et al. 2004, Halleen et
al. 2004, Castlebury et al. 2006). Anamorphs in the N. mammoidea/
veuillotiana-group (clade II in Mantiri et al. 2001) belong to
Cylindrocarpon group 2 and are characterised by the lack of
microconidia and chlamydospores. Anamorphs in Cylindrocarpon
group 3 belong to the N. radicicola-group (clade III in Mantiri et al.
2001) and are characterised by the presence of microconidia and
chlamydospores.
The anamorphic genus Campylocarpon was described by
Halleen et al. (2004) for species resembling Cylindrocarpon with
3–5-septate, curved macroconidia and lacking microconidia.
Halleen et al. (2004) segregated Campylocarpon from
Cylindrocarpon based on molecular phylogenetic data that placed
it more closely to N. mammoidea-group than to Cylindrocarpon
58
sensu stricto (N. coccinea-group). Halleen et al. (2004) noted the
similarity of Campylocarpon to the Cylindrocarpon anamorphs of
species in the N. mammoidea-group.
Even though morphological and phylogenetic studies suggest
that Neonectria/Cylindrocarpon represents more than one genus
(Samuels & Brayford 1994, Mantiri et al. 2001, Brayford et al.
2004, Halleen et al. 2004, Hirooka et al. 2005, Castlebury et al.
2006, Halleen et al. 2006), formal taxonomic segregation of these
groups has not been proposed. The objectives of the present
study are to: (1) deine Neonectria sensu stricto; (2) determine if
Neonectria/Cylindrocarpon should be divided into multiple genera
using phylogenetic analyses of multiple loci; and (3) recognise
these species in monophyletic genera as a irst step toward their
taxonomic revision.
MATERIALS AND METHODS
Morphological characterisation
Specimens were obtained from U.S. National Fungus Collections
(BPI), Steere Herbarium, New York Botanical Garden (NY), and
Manaaki Whenua Landcare Research, New Zealand (PDD),
and collected in fresh conidition from the ield. Some cultures
were obtained from the Centraalbureau voor Schimmelcultures
(CBS), Utrecht, Netherlands. For morphological characterisation
of the teleomorph, the macromorphology of the perithecia was
observed and described using the following characters: distribution
of perithecia on the host; perithecial shape, colour, and reaction
to 3 % w/v potassium hydroxide (KOH) and 100 % lactic acid;
perithecial wall structure; and colour and appearance of the
perithecial apex. Colour standards are from Kornerup & Wanscher
(1978). To observe internal and microscopic characteristics, the
perithecia were rehydrated briely in KOH, then supported by
Tissue-Tek O.C.T. Compound 4583 (Miles Inc., Elkhart, Indiana,
USA), and sectioned at a thickness of ca. 15 μm with a freezing
microtome. Characteristics of asci and ascospores were observed
by rehydrating the perithecia in 3 % KOH, removing part of the
centrum with a ine glass needle, and placing it on a glass slide.
Microscopic observations were made using an Olympus BX51
microscope and DP71 digital camera. Cultures were obtained by
isolating asci containing ascospores on cornmeal-dextrose agar
(CMD; Difco™ cornmeal agar + 2 % w/v dextrose supplemented with
antibiotics 0.2 % each neomycin and streptomycin). Morphological
observations of the colonies and anamorph in culture were based
on isolates grown on Difco™ potato-dextrose agar (PDA) and
SNA (low nutrient agar, Nirenberg 1976) for 3 wk in an incubator
at 25 ºC with alternating 12 h/12 h luorescent light/darkness.
Measurements of continuous characters such as length and width
for both anamorph and teleomorph were made using the beta 4.0.2
version of Scion Image software (Scion Corporation, Frederick,
Maryland, USA). Continuous measurements are based on 10–30
measured units and are reported as the extremes (maximum and
minimum) in brackets separated by the mean plus and minus one
standard deviation.
DNA extraction, polymerase chain reaction (PCR),
and sequencing
Strains listed in Table 1 were grown in Petri dishes (6 cm diam)
containing Difco™ potato-dextrose broth. Plates were incubated
NeoNectria/cyliNdrocarpoN and related genera
at 25 ºC for ca. 1 wk. DNA was extracted from the mycelial mat
harvested from the surface of the broth. The PowerPlant™ DNA
Isolation Kit (MO BIO Laboratories, Inc., Carlsbad, California, USA)
was used to extract DNA from the samples. Other sequences used
in the analyses were obtained from GenBank (Table 1).
DNA sequences of partial large subunit (LSU, ca. 900 bp) and
complete internal transcribed spacers 1 and 2 (ITS, ca. 600 bp),
including 5.8S of the nuclear ribosomal DNA; partial β-tubulin (tub,
ca. 500 bp); α-actin (act, ca. 600 bp); RNA polymerase II subunit
1 (rpb1, ca. 700 bp); and translation elongation factor 1α (tef1,
ca. 700 bp) were used in the phylogenetic analyses (Table 2).
The primers used and PCR protocols are listed in Table 2. Each
25 µL PCR reaction consisted of 12.5 µL Promega GreenTaq™
Master Mix 2× (Promega Corporation, Madison, Wisconsin, USA),
1.25 µL 10 mM forward primer, 1.25 µL 10 mM reverse primer,
1 µL of the DNA template, 1 µL of dimethyl sulfoxide (DMSO),
and 8 µL of sterile RNAase-free water. PCR reactions were run
in an Eppendorf Mastercycler ep using the parameters detailed
in Table 2. PCR products were cleaned with ExoSAP-IT® (USB
Corporation, Cleveland, Ohio, USA). Clean PCR products were
sequenced at the DNA Sequencing Facility (Center for Agricultural
Biotechnology, University of Maryland, College Park, Maryland,
USA). Sequences were assembled and edited with Sequencher
v. 4.9 (Gene Codes, Madison, Wisconsin, USA). Sequences were
deposited in GenBank as listed in Table 1.
Phylogenetic analyses
Sixty-nine strains and their corresponding DNA sequences were
analysed. Not all strains had all six loci sequenced and some
sequences were obtained from GenBank; see Table 1. Seven
species in the Bionectriaceae were selected as the outgroup:
Emericellopsis glabra, Hydropisphaera fungicola, Lasionectria
mantuana, Mycoarachis inversa, Nectriopsis exigua, Selinia
pulchra, and Verrucostoma freycinetiae. The included sequences
were aligned with MAFFT v. 5 (Katoh et al. 2005) using the E-INS-i
strategy. The alignment was improved by hand with Seaview v. 2.4
(Galtier et al. 1996) and MESQUITE v. 2.5 (Maddison & Maddison
2009). Gaps (insertions/deletions) were treated as missing
data. Maximum Likelihood (ML) and Bayesian (BI) analyses
were performed with all sequences, irst with each gene/locus
separately, and then with the combined data sets. A reciprocal
70 % BP threshold (Mason-Gamer & Kellogg 1996, Reeb et al.
2004) was used to determine if partitions could be combined into a
single phylogeny.
JMODELTEST (Rannala & Yang 1996, Posada & Buckley
2004, Posada 2008) was used to select the models of nucleotide
substitution for the ML and BI analyses. The number of substitution
schemes was set to 11, base frequencies +F, rate variation +I and
+G, and the base tree for likelihood calculations was set to “ML
optimised.” Once the likelihood scores were calculated, the models
were selected according to the Akaike Information Criterion (AIC).
After jMODELTEST was run, the parameters indicated in Table 2
were used for the ML and BI analyses.
GARLI v. 0.96 (Zwickl 2006) was used for the ML and bootstrap
analyses through the Grid computing (Cummings & Huskamp
2005) and The Lattice Project (Bazinet & Cummings 2008), which
includes clusters and desktops in one encompassing system
(Myers et al. 2008). In GARLI, the starting tree was obtained by
stepwise-addition and the number of runs or search replicates was
set to 50. Bootstrap (BP) analyses were replicated 2000 times. BI
www.studiesinmycology.org
analysis was done with MrBayes v. 3.1.2 (Rannala & Yang 1996,
Mau et al. 1999, Huelsenbeck et al. 2001, Huelsenbeck et al. 2002).
In MrBayes, data were partitioned by locus and the parameters of
the nucleotide substitution models for each partition were set as
described in Table 2. Two independent analyses of two parallel
runs and four chains were carried out for 10 000 000 generations
using MrBayes. Analyses were initiated from a random tree and
trees were sampled every 100th generation. Convergence of the log
likelihoods was analysed with TRACER v. 1.4.1 (beast.bio.ed.ac.uk/
Tracer). The irst 20 % of the resulting trees was eliminated (= “burn
in”). A consensus tree (“sumt” option) and posterior probabilities
(PP) were calculated in MrBayes. Phycas v. 1.1.2 (www.phycas.
org) was used as another tree searching method and also to
resolve possible polytomies (“Star Tree Paradox” problem), if any,
as proposed by Lewis et al. (2005). Phycas uses reversible-jump
MCMC to allow unresolved trees, i.e. with polytomies or very short
and poorly supported branches, and fully resolved tree topologies
to be sampled during a Bayesian analysis. Unresolved trees
generally occur when the time between speciation events is so
short or the substitution rate so low that no substitutions occurred
along a particular internal edge in the true tree. The number of
cycles in Phycas was set to 100 000, sampling every 100 cycles,
and with a starting tree obtained randomly.
RESULTS
Molecular phylogenetic analyses
Multiple sequence alignment resulted in 4184 included base pairs,
1 359 (33 %) phylogenetically informative and 2 500 invariable
sites; 325 sites presented unique non-informative polymorphic sites
(Table 2). Ambiguously aligned regions were excluded from the
analyses, especially in ITS, tef1, and tub loci, which possess highly
variable regions, i.e. introns (Table 2). Phylogenetic analyses of six
loci show high bootstrap (BP) and MrBayes posterior probabilities
(PP) for most nodes in the combined cladogram, except for a few of
the deeper nodes (Fig. 1). BI PPs were either 100 % (high support)
or 50 % (low support). The negative log likelihoods (–Ln) for the
ML, BI, and Phycas trees were 44603.27, 44959.23, and 44957.36,
respectively. The reversible-jump MCMC run in Phycas resulted
in a few improved posterior probabilities for some polytomies or
poorly supported nodes in the ML or BI trees (Fig. 1). The reciprocal
70 % BP threshold used to determine topological conlicts between
partitions resulted in complete congruence, that is, the topologies
of each gene genealogy did not contradict each other (results not
shown). This can be evidenced in the high BP and PP support
found in most nodes (Fig. 1).
Species with Cylindrocarpon-like anamorphs are contained in
two paraphyletic clades (Fig. 1): Clade A with the N. rugulosa-group,
N. mammoidea/veuillotiana-groups, and Campylocarpon (72 % BP,
100 % PP) and Clade B with the N. coccinea- and N. radicicolagroups (97 % BP, 100 % PP). These clades correspond generally
to those reported by Mantiri et al. (2001). Figure 1 also shows
that some of the groups deined by Booth (1959) and Samuels &
Brayford (1994), i.e. N. mammoidea-, N. rugulosa-, N. coccinea-,
and N. radicicola-groups, are supported by high or moderately
high BP and PP values. Campylocarpon, an anamorph genus with
morphology similar to Cylindrocarpon especially to those anamorphs
in the N. mammoidea-group (Halleen et al. 2004), clusters with the N.
mammoidea/veuillotiana-group supported by BI PP (100 %).
59
chaverri et al.
Table 1. Isolates used in the phylogenetic analyses with their corresponding GenBank accession numbers.
Species (sexual/asexual state)**
Campylocarpon fasciculare
Campylocarpon pseudofasciculare
Cosmospora coccinea / ‘Verticillium’
olivaceaum
Cosmospora vilior / Acremonium
berkeleyanum
Cosmospora viliuscula
Cosmospora sp.
Cyanonectria cyanostoma / Fusarium sp.
Cylindrocarpon destructans var. crassum (I)
Cyl. destructans var. crassum (I)
Cyl. olidum (T)
Emericellopsis glabra
Gibberella fujikuroi / Fusarium moniliforme
Gibberella fujikuroi / Fusarium moniliforme
Haematonectria haematococca / Fusarium
solani
Haematonectria illudens / Fusarium illudens
Haematonectria sp.
Hydropisphaera fungicola
Lasionectria mantuana
Leuconectria clusiae / Gliocephalotrichum
bulbilium
Mycoarachis inversa
Nectria antarctica
Nectria aquifolii
Nectria aurigera
Nectria austroamericana / Gyrostroma
austroamericanum
Nectria balansae
Nectria balsamea
Nectria berolinensis / “Tubercularia”
berolinensis
Nectria cinnabarina (dematiosa) /
Tubercularia vulgaris
Nectria coryli
Nectria cucurbitula / Zythiostroma pinastri
Nectria lamyi
Isolate
CBS 112613
CBS 112679
A.R. 2741 (= CBS 114050)
Isolate
ITS
tef1
tub
HM484515
act
rpb1
HM484537
A.R. 4215 (= CBS 126111)
HM484854
HM484869
HM484846
HM484875
HM484838
HM484872
G.J.S. 96-6 (= CBS 455.96)
G.J.S. 93-15
G.J.S. 98-127 (= CBS 101734)
CBS 537.92
CBS 605.92
CBS 215.67
A.R. 3614 (= CBS 125295)
FM 94
PMBMDF092
NRRL 22277
HM484855
HM484856
FJ474076*
GQ506003* HM484851
GQ506006* HM484849
FJ474081* HM484611
HM484876
HM484878
GQ505966*
GQ505968*
GQ505961*
GQ506032*
GQ506035*
GQ506017*
HM484879
HM352884
GQ505969*
HM364334
GQ506023*
NRRL 22090
G.J.S. 93-47 (= CBS 125113)
A.R. 4170 (= CBS 122304)
A.R. 4029 (= CBS 114291)
ATCC 22228
AF178393*
HM484862
HM484863
HM484858
HM484880
HM484877
HM484839
GQ505970*
HM484873
GQ506025*
A.R. 2745 (= ATCC 22107)
A.R. 2767 (= CBS 115033)
A.R. 4108 (= CBS 125147)
A.R. 3717 (= CBS 109874)
A.R. 2808 (= CBS 126114)
HM484861
HM484556
HM484538
HM484551
HM484555
HM364331
HM364332
AY489667*
EF607079*
EF607065*
EF607078*
HM484860
FJ755697*
FJ798606*
AF178401*
HM484589
HM352881
HM352882
GQ505967*
HM364317
GQ505993* HM484843
AF178370*
AF178362*
HM484870 HM484850
GQ505995* HM484845
HM484844
AY489732*
AY489664*
GQ505991*
HM484560
HM484565
HM484573
GQ505988
HM484840
HM484516
HM484522
HM484521
HM484520
HM484882
HM484601
HM484590
HM484600
HM484597
G.J.S. 86-117 (= CBS 125119) HM484857
A.R. 4478 (= CBS 125166)
HM484540
A.R. 2776 (= CBS 126112)
HM484543
HM484868
HM484567
HM484568
HM484848
HM484528
HM484517
CBS 278.48
HM484682
HM484729
Y.H. 0815 (= A.R. 4561)
CBS 259.58
A.R. 2779 (= CBS 115034)
HM484539
HM484541
HM484544
HM484566
GQ505998
HM484569
A.R. 4391 (= CBS 121121)
CBS 551.84
A.R. 2786 (= CBS 125131)
CBS 462.83
A.R. 4280 (= CBS 126113)
G.J.S. 98-32 (= CBS 126110)
TPPH 1
G.J.S. 85-39 (= CBS 119606)
A.R. 4505 (= CBS 125173)
A.R. 4324 (= CBS 125153)
A.R. 4499 (= CBS 125172)
Neo. ditissima / Cyl. heteronemum
CBS 100316
Neo. fuckeliana / Cyl. cylindroides var. tenue A.R. 3103 (= CBS 125133)
A.R. 4109 (= CBS 119723)
A.R. 4110 (= CBS 119200)
A.R. 4480 (= 126652)
G.J.S. 02-67 (= CBS 125109)
Neo. jungneri / Cyl. victoriae (T)
C.T.R. 71-244
Neo. liriodendri / Cyl. liriodendri (I)
CBS 112602
Neo. macrodidyma / Cyl. macrodydimum (I) CBS 112615
Neo. neobalansae / Cyl. sp. (R)
G.J.S. 85-219 (= CBS 125120)
Neo. neomacrospora / Cyl. cylindroides var. CBS 198.62
cylindroides
CBS 324.61
HM484547
HM484554
HM484545
HM484542
HM484546
HM484865
AB233175*
HM364301
Neo. radicicola / Cyl. destructans (I)
Neo. ramulariae / Cyl. obtusiusculum
HM364290
HM364297
HM364303
Nectria miltina
Nectria pseudotrichia / Tubercularia lateritia
Nectria pyrrhochlora
Nectria sinopica / Zythiostroma mougeotii
Nectria zanthoxyli
Nectriopsis exigua / Verticillium rexianum
Neo. castaneicola / Cyl. castaneicola (R)
Neo. coprosmae / Cyl. coprosmae (I)
Neo. coronata / Cyl. coronatum (T)
Neo. discophora / Cyl. ianothele (T)
60
LSU
HM364313
HM364314
AY489734*
A.R. 2553 (= ATCC 208837)
ATCC 16237
CBS 151.29
HM364294
HM364296
HM364298
HM364291
HM364292
HM364293
HM364295
HM364300
HM364299
HM364302
GQ505972*
HM484501
GQ506021*
HM484575
HM484511
HM484586
HM484874
HM484591
HM484594
HM484508
HM484510
HM484871
HM484580
HM484583
HM484647
HM484800
HM484615
HM484760
HM484536
HM484530
HM484518
HM484596
HM484592
HM484593
HM484509
GQ505974
HM484507
GQ506028
HM484582
HM484532
HM484570 HM484519
GQ506001 HM484531
HM484571 HM484523
GQ505986* HM484852
HM484609
HM484602
HM484598
HM484595
HM484599
HM484883
HM484514
GQ505976
HM484587
GQ506030
GQ505973
HM484513
GQ505979*
GQ506031
HM484585
GQ506014*
HM352862
HM352860
HM352878
HM352875
HM352877
HM352880
HM352872
HM352873
HM352874
HM352876
HM352886
HM352885
HM364328
HM364326
HM364327
HM364330
HM352883
HM364333
HM364307
HM364309
HM364311
HM446654
HM364305
HM364306
HM364308
HM364320
HM364319
HM364323
HM364315
HM364322
HM364316
HM364348
HM364345
HM364347
HM364350
HM364342
HM364343
HM364344
HM364346
HM364354
HM364353
HM352864
HM352857
HM352858
HM352859
HM352861
HM352867
HM352866
HM352853
HM364351
HM352869
HM352865
HM364318
HM364352
HM352854
HM364304
HM364310
HM364324
HM364341
HM364349
HM364340
HM352856
HM352863
HM352855
HM364336
HM364335
HM352871
HM352879
HM364325
HM364329
NeoNectria/cyliNdrocarpoN and related genera
Table 1. (Continued).
Species (sexual/asexual state)**
Isolate
Isolate
ITS
LSU
Neo. rugulosa / Cyl. rugulosum (R)
Neo. trachosa / Cyl. sp. (T)
Neo. veuillotiana / Cyl. candidulum (T)
Neo. westlandica / Cyl. sp. (T)
Neocosmospora vasinfecta / Acremoniumlike
TPPH 32
AB233176*
CBS 112467
G.J.S. 90-48 (= CBS 125118)
G.J.S. 83-156 (= CBS 112464)
A.R. 3587
HM484864
Ophionectria trichospora / Antipodium
spectabile
G.J.S. 01-206
tef1
HM364312
HM364321
HM484867
CBS 109876
tub
act
rpb1
HM352888
HM352887
HM364339
HM364338
HM364337
AB237526*
HM364356
HM364357
HM364355
HM484842
HM352870
HM352868
HM484881
HM484847
HM484886
AF543790*
AY489669*
Pseudonectria rousseliana / Volutella buxi
ATCC-MYA 627
U17416*
Rubrinectria olivacea / Nalanthamala sp.
CBS 102268
AY554219*
AY554244*
AY554238*
Selinia pulchra /
A.R. 2812
HM484859
GQ505992* HM484841
HM484884
Verrucostoma freycinetiae / Acremonium-like MAFF240100/h523
HM484866
GQ506013* HM484853
HM484885
Viridispora diparietispora / Penicillifer
CBS 114049
AY489735*
furcatus
*Sequences obtained from GenBank.
** Letters in parenthesis represent their classiication in the newly segregated genera. I: Ilyonectria; R: Rugonectria; T: Thelonectria.
AY489670*
GQ505982*
GQ505984*
GQ506022*
GQ506018*
AY489668*
Table 2. Genes/loci used in the phylogenetic analyses. Information on the primers, included bases pairs, PCR protocols, and models of
nucleotide substitution are indicated.
Locus
Primers used
(reference)
PCR protocol: Nucleotide Included sites
Annealing
substitution (# of excluded
temp. & cycles models
sites)
Phylogenetically
informative sites
(%)
Uninformative
Invariable
polymorphic sites sites
ITS
ITS5, ITS4
53 °C, 1 min, 35´
GTR+G
670 (136)
230 (34 %)
95
345
53 °C, 1 min, 35´
TIM+I+G
915 (0)
142 (16 %)
44
729
tef1-728, tef1-986
66 °C, 55 s, 9´
GTR+I+G
707 (524)
200 (20 %)
39
468
(Carbone & Kohn 1999)
56 °C, 55 s, 35´
Btub-T1, Btub-T2
55 °C, 30 s, 35´
HKY+I+G
535 (127)
260 (26 %)
49
226
Tact1, Tact2
65 °C, 30 s, 15´
GTR+I+G
635 (0)
149 (15 %)
37
4498
(Samuels et al. 2006)
48 °C, 30 s, 30´
crpb1a, rpb1c
50 °C, 2 min, 40´
GTR+I+G
722 (52)
378 (52 %)
61
283
4184
1359 (33 %)
325
2500
(White et al. 1990)
LSU
LR5, LROR
(Vilgalys n.d.)
Tef1
Tub
(O’Donnell & Cigelnik 1997)
Act
Rpb1
(Castlebury et al. 2004)
Total
The type species of Neonectria, N. ramulariae, and
Cylindrocarpon, C. cylindroides, fall in the N. coccinea-group, i.e.
Neonectria/Cylindrocarpon sensu stricto (94 % BP, 100 % PP)
(Fig. 1), part of Clade B. This group also includes N. ditissima and
N. fuckeliana. Morphological characteristics of the Neonectria/
Cylindrocarpon sensu stricto clade include perithecia aggregated
in an erumpent stroma, perithecial walls generally composed of two
regions, somewhat ornamented ascospores, and macroconidia that
are generally > 3-septate, cylindrical, and straight (Table 3). This
clade is sister to the N. radicicola-group (Clade B). The monophyly
of the N. radicicola-group is supported by 98 % BP and 100 % PP
(Fig. 1). Characteristics of the teleomorph and anamorph clearly
separate the N. radicicola-group from Neonectria/Cylindrocarpon
sensu stricto (Table 3). Perithecia in the N. radicicola-group are
supericial on the substrate and have a distinctive perithecial wall
structure, smooth ascospores, and macroconidia that are straight,
< 3-septate, with a prominent basal hilum.
www.studiesinmycology.org
The N. rugulosa group is sister to the N. mammoidea/
veuillotiana-group and Campylocarpon (Clade A). The N. rugulosagroup is monophyletic (100 % BP, 100 % PP). It contains species with
warted perithecia and a perithecial wall structure generally different
that the N. radicicola-group and Neonectria/Cylindrocarpon sensu
stricto, striate ascospores, microconidia, and no chlamydospores
(Table 3). The clade that includes the N. mammoidea/veuillotianagroup is also supported by high BP and PP values (70 % BP and
100 % PP). Species in this clade have a perithecial wall comprised
of thick-walled cells, a knobby or prominent apex, spinulose
or tuberculate ascospores, and generally no microconidia or
chlamydospores. Campylocarpon sequences form a distinct clade
sister to the N. mammoidea-group and is supported by 100 % BP
and 100 % PP. No teleomorph is known for Campylocarpon.
61
Character
Campylocarpon
(Clade A)
Rugonectria (Clade A)
Thelonectria
(Clade A)
Ilyonectria (Clade B)
Neonectria
(Clade B)
Teleomorph groups (Booth 1959, Brayford &
Samuels 1993, Samuels & Brayford 1994)
–
N. rugulosa-group
N. mammoidea /veuillotiana-groups
N. radicicola-group
N. coccinea-group
Anamorph groups (Booth 1966)
–
Group 2
Group 3
Groups 1 & 4
Arrangement of perithecia on substrate
Teleomorph unknown
Perithecia, formed on, or sometimes
partially immersed within a stroma
Perithecia solitary or in groups, supericial,
sometimes seated on an immersed
inconspicuous stroma
Generally solitary and loosely
attached to substrate
Perithecia clustered on wood, generally
seated on an erumpent stroma
Perithecial apex
–
Non-papillate
Most species with a prominent, areolate
(darkened) papilla, if not, then at least with a
darkly pigmented apex
Broadly conical papilla
Blunt or acute apex, rarely papillate
Perithecial wall
–
Warted, 50–150 μm thick; outer region,
including warts, of thick-walled (3–4 μm),
globose, 10–20 μm diam; perithecial wall
merging with surrounding stroma
Smooth or sometimes warted, 20–50 (–100)
μm thick; outer region of intertwined hyphae
or cells lacking a deinite outline, i.e. textura
epidermoidea
Generally smooth to slightly
roughened, 35–50 μm thick; outer
region of thin-walled, globose,
large cells
Generally smooth and shiny, sometimes
scurfy, 35–50 μm thick; outer region of
small, angular to globose, thick-walled
cells (textura epidermoidea in one
species)
Ascospores
–
1-septate, striate
Generally 1-septate, smooth, rarely spinulose 1-septate, smooth
or striate
Macroconidia shape
Fusiform, curved, often
broadest at upper third, with
rounded apical cells and
lattened or rounded basal
cells, inconspicuous hilum
Fusiform, curved, tapering towards ends
(almost Fusarium-like), inconspicuous
hilum
Fusiform, curved, often broadest at upper
third, with rounded apical cells and lattened
or rounded basal cells, inconspicuous hilum
Cylindrical, straight, rounded ends, Cylindrical, generally straight, sometimes
prominent basal hilum
slightly curved toward ends, with
rounded ends (except in one species, N.
fuckeliana, which has fusiform straight
conidia with pointed ends); inconspicuous
hilum
Macroconidia septation
(1–) 3–5 (–6)-septate,
average 4 septa
(3–) 5–7 (–9)-septate
(3–) 5–7 (–9)-septate, average 5
1–3-septate, rarely > 3-septate
3–7 (–9)-septate, average 5-septate
Macroconidia size
(24–) 35–60 (–62) × 6.5–9
μm
(35–) 48–85 × 5–10 μm
(35–) 40–90 (–110) × 4–8 (–11) μm
25–50 (–55) × 5–7.5 μm
35–65 (–110) × 4–7 (–8) μm
Microconidia shape
Absent
Ovoid to cylindrical, hilum inconspicuous
Microconidia rare (seen only on natural
substrate)
Ellipsoidal, prominent basal hilum
Ellipsoidal to oblong, inconspicuous hilum
Microconidia size
Absent
(3–) 5–15 (–20) × 2–5 μm
–
3–15 × 2.5–5 (–6) μm
(2–) 6–10 (–15) × (1–) 2–5 (–6) μm
Chlamydospores
Uncommon
Absent
Uncommon (except in T. olida = C. olidum)
Abundant, generally intercalary,
single or in chains, becoming
brownish
Present in some species
Substrate
Pathogenic on roots and
stems of grapevines
On bark of recently killed, dying or
diseased trees, often causing cankers
On bark of recently killed, dying or diseased
trees, often causing small cankers,
sometimes on rotting roots
Generally a root pathogen.
Anamorph common in the soil.
Perithecia found mostly on
decaying herbaceous material,
sometimes branches or roots.
Generally on bark, sometimes causing
cankers
Geographic distribution
South Africa, Uruguay
Widespread
Widespread, but more common in tropical
regions
Widespread
Mostly in temperate regions
1-septate, smooth or inely ornamented
chaverri et al.
62
Table 3. Comparison of major diagnostic morphological characteristics between the newly segregated genera.
NeoNectria/cyliNdrocarpoN and related genera
Fig. 1. Multilocus phylogenetic tree (Bayesian Inference) with the best log likelihood (-44959.23). Support values indicated at nodes. Bayesian posterior probabilities ≥ 90 %,
Maximum Likelihood bootstrap ≥ 70 % and Phycas posterior probabilities ≥ 90 % indicated by ***. If less than those values, then indicated by -. Cylindrocarpon-like anamorphs
are in two paraphyletic clades: A and B.
www.studiesinmycology.org
63
chaverri et al.
Morphological characterisation
Presence or absence of a stroma
In many cases perithecia are solitary, either seated directly on the
substratum in the N. radicicola-group or on a minute basal stroma
in the N. mammoidea/veuillotiana-group. In other cases, such as
in N. discophora and N. lucida, perithecia are seated amidst erect
hyphae that arise from a basal, almost inconspicuous stroma. A
characteristic of N. coccinea, N. fuckeliana and other species of
Neonectria sensu stricto and N. rugulosa-group is that they form
in great numbers on a rather extensive, subcortical, basal stroma.
An extensive stroma may also form in N. jungneri, but is not as
conspicuous as in Neonectria sensu stricto and the N. rugulosagroup.
Perithecial wall
The perithecial wall of species of Neonectria sensu lato comprises
at least two regions. The inner region is very thin, consisting of
only a few layers of thin-walled, tangentially lattened cells lining
the locule where the spores are formed. The outer regions vary.
Essentially four distinctive types of outer perithecial walls are found
among the groups studied here. In the N. radiciola-group the outer
region of the perithecial wall is formed of one or two layers of large,
round, thin-walled cells. This anatomy can be discerned even in
whole mounts of perithecia. In species that have this anatomy,
the surface of the perithecial wall, when seen in face view, is of
large, round cells, mirroring what is seen in sections. The second
perithecial wall anatomy consists of an outer region that is a
palisade of short hyphae that are perpendicular to the locule, e.g.
in the N. mammoidea-group. When seen in face view cells at the
surface of the perithecium are small, < 5 μm diam. In some species,
such as N. trachosa and N. westlandica, a supericial layer of large,
angular cells that form warts obscure the palisade. When there is a
palisade but no outer layer of large cells, the perithecial surface may
be smooth and shiny. This wall anatomy typiies some members
of the N. mammoidea-group, e.g. N. discophora, N. lucida, N.
westlandica; and N. fuckeliana in Neonectria sensu stricto. The
wall of species such as N. coronata or N. jungneri, both in the N.
mammoidea/veuillotiana-group, which lacks any apparent cellular
structure, is formed of intertwined hyphae having a seemingly
random arrangement rather than a palisadal arrangement. The
third perithecial wall type is characterised by the formation of thickwalled, round cells in the outer region that can be seen in section
and in face view. This wall type characterises species of Neonectria
sensu stricto. The fourth type of perithecial wall is that of the N.
rugulosa-group. The perithecial wall is thick, 50–150 μm, with the
outer region formed of several layers of cells, including warts, with
small globose cells that are very thick-walled and merge with the
surrounding stroma.
Ascospores
Although some species of Neonectria have been reported to have
multiseptate ascospores (Rossman 1983, Samuels & Brayford
1993), the ascospores of the species included in the present
study are bicellular. There is a tendency towards having spinulose
ornamentation but there are exceptions. In species such as N.
veuillotiana the ascospores may be nearly tuberculate. In N. jungneri
the spores are coarsely striate. In N. coronata the spinules may be
arranged in lines giving the appearance of striations. Ascospores
of most species are hyaline, but, in N. discophora, N. lucida, and
N. westlandica, the spores become pale yellow-brown. A species
64
not included in the present study, Nectria viridispora, probably in
the N. mammoidea-group, has green ascospores. Ascospores of
species in Neonectria sensu stricto and the N. radicicola-group are
smooth. Species in the N. rugulosa-group have striate ascospores,
sometimes inconspicuous; cotton blue may be needed to observe
these striations.
Paraphyses
The Nectria-type centrum (Luttrell 1951) is characterised by the
formation of "apical paraphyses," ilaments that originate in a
meristem situated at the top of the locule. Typically these ilaments
have dissolved by the time the ascospores form but often chains
of saccate cells may persist among maturing asci. Most species
of Neonectria sensu stricto have ilaments that appear to be free
at the apex and thus resemble paraphyses. These paraphyses
are septate and constricted at each septum. The paraphyses are
abundant especially in N. fuckeliana.
Conidiophores and phialides
Most conidiophores, especially those that give rise to the
macroconidia, are formed laterally from hyphae; they are irregularly
branched or form fascicles. In the case of Neonectria sensu
stricto and the N. rugulosa-group, the macroconidia are produced
from irregularly branched conidiophores or fascicles, and the
microconidia from simple, generally unbranched, conidiophores.
In the case of the N. radicicola-group, macro- and microconidia
apparently originate from the same type of conidiophore. These are
simple, unbranched or sparsely branched, irregularly or verticillately
branched, or rarely densely branched. The N. mammoidea/
veuillotiana-group and Campylocarpon produce only macroconidia
that originate from irregularly branched conidiophores or fascicles.
The morphology of the phialides is highly conserved. Phialides
are generally long and cylindrical or somewhat lask-shaped, but
mostly long.
Macro- and microconidia
Although the average size of the macroconidia varies among the groups,
there is signiicant overlap. Campylocarpon, the N. radicicola-group,
and Neonectria sensu stricto have macroconidia 25–65 × 4–9 μm,
smaller than those of the N. mammoidea/veuillotiana- and N. rugulosagroups that are 40–90 × 4–10 μm. With respect to shape, species in
Clade A (Fig. 1) have curved macroconidia and species in Clade B have
straight macroconidia. Within Clade A, macroconidia of the N. rugulosagroup can be easily distinguished from those in Campylocarpon and
the N. mammoidea/veuillotiana-group. Species in N. rugulosa-group
have curved, fusoid macroconidia with tapering ends that are almost
Fusarium-like. Campylocarpon and the N. mammoidea/veuillotianagroup also have curved macroconidia but with rounded ends. Even
though the macroconidia of Campylocarpon and the N. mammoidea/
veuilotiana-group are similar, they can be distinguished on the basis
of septation. Campylocarpon has 3–5-septate macroconidia while
the N. mammoidea/veuilotiana-group has 5–7-septate macroconidia.
Regarding septation of macroconidia, most species have on average
ive septa, with exceptions. On average species of Campylocarpon
have four septa, the N. radicicola-group have up to three septa with
exceptions, and N. jungneri (N. mammoidea-group) has generally
> 5 septa. Microconidial morphology is highly conserved. They are
generally ellipsoidal, 0–1-septate and measure 3–15 × 2–5 μm. Only
the N. radicicola-group has microconidia with a prominent hilum or
abscission scar. No microconidia are formed in the N. mammoidea/
veuillotina-group and Campylocarpon. Some species in Neonectria
NeoNectria/cyliNdrocarpoN and related genera
sensu stricto may produce microconidia, but not as abundantly as in
the N. radicicola- and N. rugulosa-groups. The only exception is N.
fuckeliana in which microconidia are abundant and macroconidia are
infrequently seen.
Chlamydospores
Chlamydospores are formed in the N. radicicola-group, in a
few species in Neonectria sensu stricto, and are rarely formed
in Campylocarpon. Species in the N. rugulosa-group rarely
produce swollen and slightly pigmented hyphae that resemble
chlamydospores. Most species in the N. mammoidea/veuillotianagroup do not produce chlamydospores, except in Cylindrocarpon
olidum. The chlamydospores of the N. radicicola-group are
generally intercalary, single or in chains, and yellow-brown. When
produced in Campylocarpon, they are mostly terminal, single, or in
chains of 2–3, and also yellow-brown.
Ecology
Species of Neonectria sensu lato and Cylindrocarpon sensu lato
are either saprobes or plant pathogens. The only two known
Campylocarpon species cause black foot disease of grapevines.
Species in the N. mammoidea/veuillotiana-group are only known
as saprobes growing on bark of recently killed woody trees. The
only exception is Cylindrocarpon olidum, which has been reported
as a root pathogen. Members of the N. rugulosa-group and
Neonectria sensu stricto also grow on bark of recently killed trees
and many species, e.g. N. castaneicola, N. ditissima, N. faginata,
N. rugulosa among others, can cause cankers. In contrast, species
in the N. radicicola-group are generally found in the soil and cause
many root diseases. Based on the present study, the species that
are commonly found in the soil causing root rots are the ones that
produce chlamydospores. On the other hand, the species that
grow on bark do not produce chlamydospores. Members of the N.
rugulosa- and N. radicicola-groups are widespread, N. mammoidea/
veuillotiana-group are mostly tropical and subtropical, Neonectria
sensu stricto occur in temperate regions, and Campylocarpon is
known only from South Africa and Uruguay (Abreo et al. 2010).
DISCUSSION
Genus concept
Several morphological characteristics of the teleomorphs and
anamorphs have been used in deining informal groups in Nectria
sensu lato. In the case of Neonectria and Cylindrocarpon groups
(e.g. Booth 1959, 1966), they have been distinguished by the
anatomy of the outer regions of the perithecial wall and presence
or absence of microconidia and chlamydospores. Results from this
study conirm previous suggestions that Neonectria/Cylindrocarpon
is paraphyletic, comprising ive independent lineages that may be
interpreted as distinct genera. These segregate genera usually
cannot be distinguished based on a single morphological or
ecological character. However, the lineages or segregate genera
correlate strongly with a combination of ecology and morphological
characters of the perithecia and anamorphs (Table 3). Thus, the
following genera are recognised: (1) Neonectria/Cylindrocarpon
sensu stricto (N. coccinea-group); (2) N. rugulosa-group, hereafter
Rugonectria gen. nov.; (3) N. mammoidea/veuillotiana-group,
hereafter Thelonectria gen. nov.; (4) N. radicicola-group, hereafter
www.studiesinmycology.org
Ilyonectria gen. nov.; and (5) Campylocarpon. The Neonectria and
Cylindrocarpon groups deined by Booth (1959, 1966) based on
morphological characters generally agree with the clades observed
in the multilocus phylogeny (Fig. 1).
Based on the morphological similarity between Campylocarpon
and Thelonectria, it could be argued that these two are congeneric.
However, phylogenetic analyses do not support the monophyly
of these two genera (short branch length, and low BP and PP
supports, Fig. 1). Therefore, Campylocarpon and Thelonectria are
recognised as separate. Several morphological and ecological
traits aid in distinguishing these two genera (Table 3).
Although Clades A and B (Fig. 1) could be recognised as
two genera, the multiple morphological and ecological traits of
each of the ive segregate genera are distinctive enough to justify
their taxonomic subdivision. There are other similar cases in the
Ascomycota. For example, although genera with fast-growing
Fusarium anamorphs form a monophyletic group, they are still
recognised as separate genera and have morphologically different
teleomorphs (e.g. Albonectria, Cyanonectria, Gibberella, and
Haematonectria) (O’Donnell 1996, Samuels et al. 2009, Luo &
Zhuang 2010a). Another example is Calonectria/Cylindrocladium
and related genera. Glionectria/Gliocladiopsis, Nectricladiella/
Cylindrocladiella, and Xenocalonectria/Xenocylindrocladium all
have similar anamorph and teleomorph morphology and were
previously classiied in Calonectria. Later they were segregated
from Calonectria/Cylindrocladium based mostly on anamorph
characteristics even though they form a monophyletic group
(Rossman 1983, 1993, Schoch et al. 2000, Crous 2002, Samuels et
al. 2009, Luo & Zhuang 2010a). A third example is Botryosphaeria
sensu lato. Many recognised monophyletic anamorphic genera, e.g.
Fusicoccum, Lasiodiplodia, and Neofusicoccum among others, are
associated with Botryosphaeria teleomorphs, yet, Botryosphaeria
s. l. forms a monophyletic group (Crous et al. 2006).
Results from the present study show that Neonectria fuckeliana
clusters with Neonectria/Cylindrocarpon sensu stricto, and T.
jungneri with Thelonectria. The branch lengths (substitutions/site)
that separate these species from Neonectria/Cylindrocarpon sensu
stricto and Thelonectria, respectively, are similar to the branch
lengths between Thelonectria and Rugonectria (Fig. 1). This could
be interpreted as evidence that N. fuckeliana or T. jungneri should
be recognised as distinct genera. However, these species are not
separated due to the lack of additional morphologically similar
species and to avoid monotypic genera with further splitting of
genera. It is possible that the addition of morphologically similar
species will support the establishment of new genera.
Neonectria/Cylindrocarpon sensu stricto
Neonectria/Cylindrocarpon sensu stricto is characterised by having
few to numerous perithecia clustered on wood and seated on an
erumpent stroma; perithecial walls are generally composed of
two regions with the outer region comprising small, thick-walled
cells; generally septate paraphyses; smooth or inely ornamented
ascospores; generally straight, typically 5-septate macroconidia
with rounded ends; either microconidia or chlamydospores
formed, generally not both; and, if microconidia are present, they
are produced from simple, generally unbranched, conidiophores
and lack a prominent abscission scar. Anamorphs of Neonectria/
Cylindrocarpon belong in Booth’s groups 1 and 4 (Booth 1966).
Neonectria/Cylindrocarpon sensu stricto species are mostly
found in temperate regions on woody substrata, e.g. bark, often
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chaverri et al.
causing cankers, and rarely found in soil. This genus includes
species such as N. coccinea/C. candidum, N. ditissima/C.
heteronemum, N. faginata/C. faginatum, N. fuckeliana/C.
cylindroides var. tenue, N. hederae/C. hederae, N. major/
Cylindrocarpon sp., N. neomacrospora/C. cylindroides, N.
punicea/C. album, and N. ramulariae/C. obtusiusculum (Castlebury
et al. 2006). The monophyly of the N. coccinea-group was shown
in Castlebury et al. (2006). Although some authors suggested that
N. fuckeliana belongs in the N. mammoidea-group based on the
morphology of the perithecia (Booth 1959, Brayford et al. 2004),
this study supports more recent accounts that place this species
close to Neonectria sensu stricto (Halleen et al. 2004, Castlebury
et al. 2006, Luo & Zhuang 2010b).
The teleomorph of the type species of Neonectria, N.
ramulariae, apparently has not been collected again since it was
described by Wollenweber (1917) (Rossman et al. 1999). Rossman
et al. (1999) examined the type specimen and noted that it had only
immature perithecia along with its anamorph, C. obtusiusculum
(= C. magnusianum Wollenw. 1928 non Wollenw. 1926). Domsch
et al. (1980) followed Wollenweber (1928) in recognising N.
ramulariae to be the teleomorph of C. obtusiusculum (then known
as C. magnusianum), based on the anamorph present in the
type specimen of N. ramulariae. Although Rossman et al. (1999)
designated an iconotype for N. ramulariae, new collections of
the anamorph and teleomorph are needed to better describe N.
ramulariae/C. obtusiusculum. The morphology of C. obtusiusculum
is similar to the anamorphs in Ilyonectria. However, the lignicolous
habit, straight macroconidia, absence of microconidia, absence
of a prominent basal abscission scar or hilum, and molecular
phylogenetic analyses place C. obtusiusculum in Neonectria/
Cylindrocarpon sensu stricto.
The segregate genera: Campylocarpon, Ilyonectria,
Rugonectria, and Thelonectria
A sister clade to Neonectria/Cylindrocarpon, Ilyonectria (N.
radicicola-group), is described here based on Ilyonectria radicicola
comb. nov. (anamorph C. destructans). Anamorphs in Ilyonectria
belong in Booth’s group 3 (Booth 1966). Contrary to Neonectria/
Cylindrocarpon, Ilyonectria and its anamorphs are common
in the soil and rhizosphere or as agents causing root rots.
Chlamydospores are generally present in species of Ilyonectria,
possibly as an adaptation for survival in soil. Chlamydospores
are generally absent in species that are associated with bark or
cankers, e.g. Neonectria, Rugonectria and Thelonectria. Perithecia
in Ilyonectria are not as commonly encountered as the anamorphs,
and, if found, they are mostly on herbaceous substrata. The
species of this genus are cosmopolitan and are found on a wide
range of hosts.
Neonectria-like species included here in Ilyonectria are:
I. coprosmae/C. coprosmae, I. liriodendri/C. liriodendri, I.
macrodydima/C. macrodydimum, and I. radicicola/C. destructans
(Samuels & Brayford 1990, Seifert et al. 2003, Halleen et al.
2004, 2006). The monophyly of species in Ilyonectria, viz. the N.
radicicola-group, has also been shown in previous studies (Seifert
et al. 2003, Halleen et al. 2004, 2006). These studies suggest that
C. destructans is a species complex. Thus, deining C. destructans
sensu stricto through the examination of many cultures derived
from ascospores as well as cultures isolated directly from diverse
substrata is a necessary future endeavour. Many other species
have been described that may it in Ilyonectria.
66
Rugonectria gen. nov. (N. rugulosa-group) is described
here based on Rugonectria rugulosa comb. nov. (anamorph C.
rugulosum). Members of the genus occur on recently killed or dying
woody substrata, mostly bark, and are sometimes found causing
cankers. Some species of Neonectria now included in Rugonectria
are: R. castaneicola/C. castaneicola, R. neobalansae, and R.
rugulosa/C. rugulosum. Another species that may it in Rugonectria
is Nectria pulcherrima (Samuels & Brayford 1994). This species
has multiseptate, curved macroconidia with tapering ends,
microconidia, and warted perithecia that are caespitose, somewhat
immersed in an erumpent stroma, all characteristics of Rugonectria.
This species is morphologically similar to R. neobalansae. A new
combination has not been made due to the lack of DNA data to
conirm its phylogenetic placement.
The new genus Thelonectria is established here to accommodate
species in the N. mammoidea- and N. veuillotiana-groups. Species of
Thelonectria are mostly tropical and subtropical, and are found on bark
of recently killed or dying trees, often causing small cankers, rarely
in soil except in one species, C. olidum. Some species included in
this genus are: T. coronata/C. coronatum, T. discophora/C. ianothele,
T. jungneri/C. victoriae, T. lucida/C. lucidum, T. olida, T. trachosa, T.
veuillotiana/C. candidulum, T. viridispora, and T. westlandica (Mantiri
et al. 2001, Brayford et al. 2004). Anamorphs in Thelonectria belong
in Booth’s group 2 (Booth 1966).
Although Thelonectria can generally be recognised by
perithecia with prominent or darkened papilla, macroconidia that
are curved with rounded ends, > 3-septate (average 5-septate),
and absence of microconidia, some species deviate from this
trend. For example, 3-septate macroconidia have been reported
for T. lucida and T. trachosa (Booth 1966, Brayford et al. 2004).
Thelonectria trachosa mostly forms 3-septate macroconidia, but >
3-septate macroconidia can be found in the same culture (Brayford
et al. 2004). Brayford et al. (2004) reported that the majority of the
T. lucida cultures formed > 3-septate macroconidia. Brayford et
al. (2004) also suggested that T. lucida might comprise a species
complex, thus, further taxonomic studies are needed to explain the
morphological variation within this species. Thelonectria lucida and
T. trachosa can be easily classiied in Thelonectria based on the
anatomy of the perithecia and curved macroconidia with rounded
ends and absence of microconidia and chlamydospores. A similar
case is T. olida, which produces 3–5-septate macroconidia and
chlamydospores although Booth (1966) reports many > 3-septate
macroconidia. This species is classiied in Thelonectria based
on the curved macroconidia with rounded ends and absence
of microconidia. However, T. olida is dificult to distinguish from
Campylocarpon based on morphology and ecology.
Conidia in Campylocarpon are similar to those in Thelonectria,
as also reported by Halleen et al. (2004). The only morphological
difference is the average number of septa in the macroconidia: four in
Campylocarpon and ive in Thelonectria. Despite the morphological
similarity of the conidia, phylogenetic analysis distinguishes the
two genera. Campylocarpon species were collected from diseased
roots and stems of grapevines in South Africa. This is in contrast
to most species of Thelonectria, which are found on above ground
parts of woody plants. Thelonectria olida, associated with roots, is
the exception.
Previous molecular phylogenetic studies (Mantiri et al. 2001,
Brayford et al. 2004) did not show that the N. rugulosa-group was
distinct from N. mammoidea-group, as suggested by Samuels &
Brayford (1994). This was probably due to the few phylogenetically
informative loci and few taxa that were used in those studies.
The monophyly of the N. rugulosa-group (= Rugonectria) and its
NeoNectria/cyliNdrocarpoN and related genera
close relationship to the N. mammoidea/veuillotiana-group (=
Thelonectria) are shown here (Fig. 1). Rugonectria is distinguished
from Thelonectria by perithecial anatomy, presence of microconidia
in Rugonectria, and morphology of the macroconidia (Table 3).
As has been the case with several groups of fungi (Chaverri et
al. 2003, Frisvad & Samson 2004, Schmidt et al. 2004, Samuels et
al. 2006a, Chaverri et al. 2008, Degenkolb et al. 2008, Andersen
et al. 2009), a multiphasic approach, i.e. using a combination
of independently derived characters such as morphological,
ecological, and molecular phylogenetic, is necessary to identify
monophyletic groups with Neonectria/Cylindrocarpon-like
morphology. For example, the presence of microconidia alone is
not useful to identify groups with Cylindrocarpon-like morphology,
because microconidia are always present in Ilyonectria and
Rugonectria, sometimes present in Neonectria, and absent in
Thelonectria, and their morphology is highly conserved. However,
if characters are combined such as the presence of 3-septate,
straight macroconidia with a prominent abscission scar, presence
of chlamydospores, and perithecia with a particular wall anatomy,
they can be used to classify a particular specimen as Ilyonectria.
Thus, in this study our genus concept is based on a multilocus
phylogenetic analyses correlated with a combination of multiple
morphological and ecological characters. Each of the proposed
genera is further described in the Taxonomy section.
Species of Neonectria/Cylindrocarpon of uncertain
classiication
In this study we present a general overview of genera with
Neonectria/Cylindrocarpon-like morphology. There are still species
classiied in Neonectria and Cylindrocarpon that have teleomorph
and anamorph morphology different than those presented here
and also quite distinct from Neonectria/Cylindrocarpon sensu
stricto. Additional specimens, cultures, and DNA sequences are
needed to infer their phylogenetic position within the Nectriaceae.
For example, Neonectria macroconidialis has morphological
characteristics of both Neonectria sensu stricto and Ilyonectria. This
species is not formally included in Ilyonectria because phylogenetic
studies including this species in the ITS tree (Seifert et al. 2003,
Halleen et al. 2004) show low bootstrap support for the clade with
N. macroconidialis and other species in the N. radicicola-group.
In contrast, the β-tubulin tree places N. macroconidialis basal and
outside the N. radicicola complex. Therefore, the phylogenetic
position of this species is uncertain. The straight macroconidia,
prominent basal hilum, and anatomy of the perithecia suggest that
N. macroconidialis belongs in Ilyonectria. However, the > 4-septate
macroconidia, a characteristic of Neonectria sensu stricto, would
be an exception if this species were included in Ilyonectria. This
species and others previously placed in the N. radicicola-group
(Samuels & Brayford 1990) are morphologically atypical of this
group, speciically N. austroradicicola/C. austrodestructans and N.
radicicola variant ex Gahnia.
Brayford & Samuels (1993) described three species of Nectria
with Cylindrocarpon-like anamorphs and mentioned that they could
not be classiied in any of the then recognised groups of Nectria.
Nectria neblinensis and N. verrucospora are distinct because
they have macroconidia that are torpedo-like, viz. straight, wider
near the middle or towards the base, and tapering and truncated
at the ends. The perithecial wall anatomy somewhat resembles
Rugonectria, but the ascospores in these two species are warted
and not striate as in Rugonectria. Other species that have been
www.studiesinmycology.org
placed in Cylindrocarpon that have torpedo-like macroconidia are
C. fusiforme, C. supersimplex, and N. laetidiscoides; however,
these are straight in the middle and the terminal cells taper almost
to a point (Matsushima 1975, Samuels & Brayford 1993).
Several species previously classiied in Neonectria/
Cylindrocarpon are distinct from those treated here because they
have phragmosporous ascospores, e.g. N. fusispora, N. laetidisca,
N. laetidiscoides, N. phaeodisca, N. philodendri, N. septospora and
N. vermispora among others (Rossman 1983, Samuels & Brayford
1993). Most of the above appear to belong in Thelonectria, or
at least they are closely related, except N. laetidiscoides and N.
septospora, which have distinct macroconidia.
Another species with uncertain afinity is N. cinnamomea. The
perithecia do not change colour in 3 % KOH, a typical characteristic
of members of the Nectriaceae (Brayford & Samuels 1993). In
addition, the perithecial wall is completely different from the genera
treated in this study or any other genus in Nectriaceae, and the
ascospores have a conspicuous wrinkled sheath. The macroconidia
are also distinct; they are curved, fusiform, and 3-septate.
Luo & Zhuang (2010b) described Neonectria shennongjiana
based mostly on the distinctive macroconidia that are cylindricalclavate to clove-shaped. The phylogenetic analysis in Luo & Zhuang
(2010b) shows that N. shennongjiana may be closely related to
Neonectria sensu stricto. Their parsimony cladogram reveals that
N. shennongjiana clusters within Neonectria sensu stricto (BP 72
% if N. fuckeliana is included). However, in their phylogenetic tree
based on parsimony analysis of two loci (ITS nrDNA and tub),
the position of N. shennongjiana is not clear. The bootstrap value
supporting the clade of N. shennongjiana and C. obtusisporum
is low (62 %). Additional phylogenetic and taxonomic studies are
needed to conirm if N. shennongjiana and other species with oddshaped macroconidia belong in Neonectria s.str. Another species
with clove-shaped macroconidia is described in the literature, i.e.
Nectria lugdunensis (Webster 1959), the teleomorph of Heliscus
lugdunensis.
TAXONOMY
Many of the species of Neonectria sensu lato, including those
considered here, are known in both their teleomorph and anamorph
states. Although Article 59 of the International Code of Botanical
Nomenclature (ICBN) allows the use of two scientiic names for
some groups of pleomorphic fungi including ascomycetes, a trend
exists toward the use of just one scientiic name for each species
regardless of the state manifested (Rossman & Samuels 2005,
Rossman 2009). Additionally, generic names of asexual fungi are
now being used in a narrower, phylogenetic sense rather than as
broad form-genera that encompass unrelated fungi. For example,
the genus Verticillium sensu lato, which traditionally included many
species with verticillate branching, has been segregated into
distinct phylogenetic genera in spite of morphological similarities.
Recently, Verticillium sensu stricto was conserved with a different
type so that it represents the plant pathogenic species such as V.
alboatrum and V. dahliae (Zare et al. 2004). Moreover, other genera
separated from Verticillium sensu stricto are now recognised based
on distinctive morphological and ecological characteristics, e.g.
Lecanicillium and Pochonia (Gams & Van Zaayen 1982, Zare et al.
2000, Gams & Zare 2001, Zare & Gams 2001a, b, Zare et al. 2001).
The anamorphs of Neonectria sensu lato have been classiied
in the genus Cylindrocarpon. Just as Neonectria is now conceived
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chaverri et al.
in a narrow sense, the genus Cylindrocarpon is herein deined
phylogenetically and restricted to only anamorphs of Neonectria
sensu stricto. Thus, the anamorph name in Cylindrocarpon is listed
for only those species that belong in Neonectria sensu stricto.
However, for species in genera segregated from Neonectria sensu
lato with an anamorph name in Cylindrocarpon, the scientiic
name of the anamorph is listed in quotes, e.g. "Cylindrocarpon"
destructans, or as Cylindrocarpon-like, if no epithet exists, to
indicate that it does not belong in Cylindrocarpon sensu stricto.
In this paper, some species described in Cylindrocarpon
have no known teleomorph, but, phylogenetically, they fall into a
recognised genus (e.g. "C." olidum = Thelonectria olida comb. nov).
As permitted by the ICBN this scientiic name is recombined in the
new genus. Recent examples in the literature include Lombard et
al. (2009), in which species are described in Calonectria despite
the lack of known teleomorphs. Although it would be possible and
correct according to ICBN Art. 59 to place these taxa into newly
described or existing anamorph genera, this has not been done
to avoid separating anamorph names from holomorph genera,
which is redundant, confusing, and unnecessary. If and when a
teleomorph were discovered for this species and a new name were
proposed for it, at present, priority would be given to that teleomorph
name rather than the anamorph name. Alternatively, the anamorph
name could be epitypiied with an element that represents the
teleomorph in accordance with ICBN Art. 59.7. Given the confusion
that has arisen because of the dual nomenclature associated with
pleomorphic fungi and the usefulness of molecular systematics
in determining the accurate taxonomic placement of asexually
reproducing fungi, it would seem expedient to move toward the use
of only one scientiic name for all fungi.
KEY TO SEGREGATE GENERA OF NEONECTRIA/CYLINDROCARPON
1. Perithecia generally on herbaceous material, rarely on bark or woody parts; perithecia supericial, loosely attached to substratum;
perithecial wall of two regions, outer region of thin-walled (ca. 1 μm), globose, large cells; ascospores smooth; anamorph in soil, generally
associated with diseased roots; microconidia generally with a prominent abscission scar; chlamydospores present; macroconidia straight,
generally < 3-septate, generally with a prominent abscission scar ............................................................................................ Ilyonectria
1. Perithecia and macroconidia not as above ............................................................................................................................................... 2
2. Perithecia smooth to slightly roughened, generally red, with a prominent papilla or non-papillate; ascospores generally smooth or slightly
ornamented; microconidia present or absent; chlamydospores present or absent; macroconidia curved or almost straight, with rounded
ends, generally 3–5-septate; on bark or roots ........................................................................................................................................... 3
2. Perithecia conspicuously warted, orange-red, generally aggregated, with an inconspicuous papilla, perithecial wall 50–150 μm thick;
ascospores striate; microconidia present; chlamydospores absent; macroconidia fusiform with tapering ends; generally on bark of
recently killed trees or causing small cankers ......................................................................................................................... Rugonectria
3. Perithecia clustered on wood, generally seated on an erumpent stroma, generally smooth and shiny, sometimes scurfy with a blunt
or acute apex, rarely papillate; perithecial walls of 2–3 regions, outer region of small, angular to globose, thick-walled cells, rarely of
textura epidermoidea; many species with septate paraphyses; ascospores ellipsoidal, smooth or inely ornamented; either microconidia
or chlamydospores present; macroconidia generally straight or slightly curved toward ends, rarely clove-shaped, with rounded ends,
rarely tapering, 5–7-septate; chlamydospores rare; on bark of recently killed trees or forming cankers .................................. Neonectria
3. Perithecia mostly aggregated, generally smooth and shiny, with a prominent papilla; ascospores generally ornamented; microconidia and
chlamydospores absent; macroconidia curved, often broadest at upper third, with rounded apical cells and lattened or rounded basal
cells, 3–7-septate; on bark of recently killed trees, on small cankers, or diseased roots .......................................................................... 4
4. Teleomorph unknown; macroconidia on average 4-septate; on diseased roots and stems of grapevines; generally pathogenic;
macroconidia generally 3–5-septate (average 4); known from South Africa and Uruguay ................................................ Campylocarpon
4. Teleomorph common, on bark of recently killed trees or causing small cankers; perithecia supericial, most species with a prominent,
darkened papilla, if not, then at least with a darkly pigmented apex; perithecial walls of 2–3 regions; outer region of intertwined hyphae
or cells lacking a deinite outline i.e. textura epidermoidea, with thickened and pigmented walls; ascospores mostly ornamented, becoming
brownish at maturity; anamorphs rarely encountered apart from their teleomorph; macroconidia (4–)5–7(–9)-septate (average 5) (except
T. olida; see section on Description of Genera) ....................................................................................................................... Thelonectria
DESCRIPTION OF GENERA
In this paper ive genera are described that have neonectria- and
Cylindrocarpon-like morphology: Campylocarpon (teleomorph
unknown); Ilyonectria gen. nov. (anam. Cylindrocarpon-like);
Neonectria sensu stricto (anam. Cylindrocarpon sensu stricto);
Rugonectria gen. nov. (anam. Cylindrocarpon-like); and
Thelonectria (anam. Cylindrocarpon-like). New combinations are
made only for those species that are conirmed to belong to the
new genera based on molecular phylogenetic data presented here
or in previous studies (Seifert et al. 2003, Brayford et al. 2004,
Halleen et al. 2004, 2006, Castlebury et al. 2006).
68
CAMPYLOCARPON Halleen, Schroers & Crous, Stud.
Mycol. 50: 449. 2004. Fig. 2.
Type: Campylocarpon fasciculare Schroers, Halleen & Crous,
Stud. Mycol. 50: 449. 2004.
Teleomorph: Unknown.
Anamorph: Cylindrocarpon-like; microconidia not observed;
chlamydospores rarely observed; conidiophores arising laterally from
hyphae, irregularly branched conidiophores or forming fascicles;
phialides cylindrical, (13–)15–20(–25) × (2–)3.5–4 μm; macroconidia
NeoNectria/cyliNdrocarpoN and related genera
Fig. 2.A–D. Campylocarpon. A–C. C. fasciculare conidiophores and macroconidia (CBS 112613). D. C. pseudofasciculare chlamydospores (CBS 112679). Bars: 10 μm.
curved, often broadest at upper third, with rounded apical cells and
lattened or rounded basal cells, (1–)3–5(–6)-septate (average 4),
with inconspicuous hilum, (24–)35–60(–62) × 6.5–9 μm.
Distribution: South Africa.
Habitat: On roots and stems of grapevines; generally pathogenic.
ILYONECTRIA P. Chaverri & C. Salgado, gen. nov.
MycoBank MB518558. Fig. 3.
Distribution: Known from South Africa and Uruguay (Abreo et al.
2010).
Campylocarpon fasciculare Schroers, Halleen & Crous,
Stud. Mycol. 50: 449. 2004.
Teleomorph: Unknown.
Habitat: On diseased roots, rootstock and stems of grapevines.
Distribution: South Africa.
Description and illustrations: Halleen et al. (2004).
Campylocarpon pseudofasciculare Halleen, Schroers &
Crous, Stud. Mycol. 50: 451. 2004.
Teleomorph: Unknown.
Habitat: On asymptomatic grapevine roots.
www.studiesinmycology.org
Description and illustrations: Halleen et al. (2004).
Type: Ilyonectria radicicola (Gerlach & L. Nilsson) Chaverri & C.
Salgado.
Etymology: “ilyo” = Greek for “mud” or “dirt”. The name is given
because most species are found as soil inhabitants.
Ascomata supericialia, globosa vel sublobosa, verrucata vel squamosa, rubra,
KOH+ phaeorubra, papilla conica vel subconica. Ascosporae ellipsoidea,
1-septatae, hyalinae, glabra. Anamorphosis cylindrocarpon-similis. Microconidia
et chlamydosporae abundans. Phialide cylindrici. Macroconidia cylindrici, recte,
hyaline, 1–3-septatae, hilum conspicue. Microconidia ellipsoidea vel oblonga,
hyaline, 0–1-septatae, hilum conspicue. Typus: Ilyonectria radicicola.
Teleomorph: Perithecia supericial, loosely attached to substrate,
red, KOH+, globose to subglobose, 175–350 μm diam, with a
broadly conical papilla, scaly or slightly warted; perithecial wall of
two regions, 35–50 μm thick: outer region 25–30 μm thick, of thinwalled, ca. 1 μm, globose, large cells; inner region of compressed,
lattened cells. Ascospores ellipsoidal, 1-septate, smooth, hyaline.
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chaverri et al.
Fig. 3. Ilyonectria. A, B. I. radicicola perithecia (A.R. 2553). C, D. Crushed perithecium of I. radicicola showing perithecium wall surface (A.R. 2553). E, F. Longitudinal section
of perithecium (TFM FPH-7807) of I. radicicola. G. Asci and ascospores of I. radicicola (A.R. 2553). H–J. Conidiophores and conidia of I. macrodydima (CBS 112615). K.
Conidiophores and conidia of I. radicicola (C.T.R. 71-76). L. Chlamydospores of I. radicicola (A.R. 2553). Bars: A, B = 500 μm; C, E, F = 100 μm; D, G, J, L = 10 μm; H, I = 20
μm; K = 50 μm.
Anamorph: Cylindrocarpon-like; microconidia and chlamydospores
abundant; macro- and microconidia apparently originating from
same conidiophores. Conidiophores 40–160 um long, generally
simple, unbranched or sparsely branched, irregularly or verticillately
branched, rarely densely branched. Phialides cylindrical, 15–40
(–50) × 1.5–3 μm. Macroconidia straight, hyaline, 1–3-septate,
rarely > 3-septate, 25–50(–55) × 5–7.5 μm, generally with a
prominent basal or lateral abscission scar or hilum. Microconidia
ellipsoidal to ovoid, hyaline, 0–1-septate, with a lateral or basal
hilum, 3–15 × 2.5–5(–6) μm. Chlamydospores abundant, generally
intercalary, globose, single or in chains, becoming brownish.
70
Habitat: On roots, soil, woody and herbaceous plants, often
pathogenic.
Notes: One potential existing generic name for this group is
Coleomyces Moreau & M. Moreau that Booth (1966) listed as
a synonym of Cylindrocarpon. The illustration in the original
description of Coleomyces, based on C. rufus (Moreau & Moreau
1937), suggests that it belongs in the N. radicicola-group. However,
in the original description the authors refer to this name as “ad
interim.” Ad interim means it is a provisional name and, according to
the ICBN (Art. 34.1, Ex. 6), it is not validly published. The authors of
NeoNectria/cyliNdrocarpoN and related genera
the present study were not able to ind a later publication validating
this name. Therefore, Coleomyces cannot be used for species in
the N. radicicola-group.
Distribution: France, Portugal, New Zealand, South Africa, USA.
Ilyonectria coprosmae (Dingley) P. Chaverri & C. Salgado,
comb. nov. MycoBank MB518559.
Ilyonectria macrodidyma (Halleen, Schroers & Crous) P.
Chaverri & C. Salgado, comb. nov. MycoBank MB518562.
Basionym: Nectria coprosmae Dingley, Trans. Roy. Soc. New
Zealand 79: 200. 1951.
≡ Nectria radicicola var. coprosmae (Dingley) Samuels & Brayford,
Mycol. Res. 94: 438. 1990.
≡ Neonectria coprosmae (Dingley) Seifert, Phytopathology 93: 1541.
2003.
Anamorph: "Cylindrocarpon" coprosmae C. Booth, Mycol. Pap.
104: 16. 1966.
Basionym: Cylindrocarpon destructans var. coprosmae (C. Booth)
Brayford & Samuels, Mycol. Res. 94: 438. 1990.
Habitat: On various decaying woody and herbaceous plants.
Distribution: New Zealand.
Descriptions and illustrations: Booth (1966) and Samuels &
Brayford (1990).
Notes: Brayford & Samuels (1990) accepted this species as a
variety of Cylindrocarpon destructans. However, Seifert et al.
(2003) recognised it as a separate species. To better elucidate
the taxonomic and phylogenetic relationship of I. coprosmae/’C.’
coprosmae to I. radicicola/’C.’ destructans sensu stricto, further
detailed taxonomic studies are needed.
Ilyonectria radicicola (Gerlach & L. Nilsson) P. Chaverri &
C. Salgado, comb. nov. MycoBank MB518560.
Basionym: Nectria radicicola Gerlach & L. Nilsson, Phytopath. Z.
48: 225. 1963.
≡ Neonectria radicicola (Gerlach & L. Nilsson) Mantiri & Samuels, Canad.
J. Bot. 79: 339. 2001.
Anamorph: "Cylindrocarpon" destructans (Zinssm.) Scholten var.
destructans, Netherl. J. Plant Path. 70 suppl. (2): 9. 1964.
Basionym: Ramularia destructans Zinssm., Phytopathology 8: 570.
1918.
Description and illustrations: Halleen et al. (2006).
Basionym: Neonectria macrodidyma Halleen, Schroers & Crous,
Stud. Mycol. 50: 446. 2004.
Anamorph: "Cylindrocarpon" macrodidymum Schroers, Halleen &
Crous, Stud. Mycol. 50: 447. 2004.
Habitat: On diseased roots and rootstocks.
Distribution: Australia, Canada, New Zealand, South Africa.
Description and illustrations: Halleen et al. (2004).
NEONECTRIA Wollenw., Ann. Mycol. 15: 52. 1917. Fig. 4.
Type: Neonectria ramulariae Wollenw.
= Chitinonectria Morelet, Bull. Soc. Sci. Nat. Archéol. Toulon Var 178: 6. 1969.
Type: Ch. coccinea (Pers. : Fr.) Morelet (≡ Sphaeria coccinea Pers. : Fr.,
≡ Neonectria coccinea (Pers. : Fr.) Rossman & Samuels).
Anamorph: Cylindrocarpon Wollenw., Phytopathology 3: 225.
1913. Type species Cylindrocarpon cylindroides Wollenw.
[= Fusidium Link : Fr., Syst. Mycol. 1: x1. 1821 : 3(2): 480. 1832 nomen
rejiciendum]
Teleomorph: Perithecia clustered on wood, generally seated on an
erumpent stroma, red, KOH+ dark red, yellow in lactic acid, generally
smooth and shiny, sometimes scurfy, subglobose to broadly
obpyriform, 200–400 μm diam, generally not collapsing when dry,
with a blunt or acute apex, rarely papillate. Perithecial walls of 2–3
regions, generally 35–50 μm thick: outer region of small, angular
to globose, thick-walled cells, rarely of textura epidermoidea; inner
region of lattened thin-walled cells. Paraphyses when present,
septate, slightly constricted at each septum. Ascospores ellipsoidal,
smooth or inely ornamented, 1-septate, hyaline, sometimes
becoming pale brown at maturity.
Descriptions and illustrations: Booth (1966, 1967), Samuels &
Brayford (1990).
Anamorph: Either microconidia or chlamydospores present.
Macroconidia produced from irregularly branched conidiophores
or fascicles. Phialides cylindrical, typically 10–20(–30) μm long.
Macroconidia hyaline, smooth, generally straight, sometimes
slightly curved toward ends, with rounded ends except in one
species, N. fuckeliana, which has fusiform conidia with pointed
ends, 3–7(–9)-septate, mostly 5-septate, lacking a prominent scar
or basal hilum, 35–65(–110) × 4–7(–8) μm. Microconidia produced
from simple, generally unbranched conidiophores, short or long;
microconidia hyaline, smooth, ellipsoidal to oblong, 0–1-septate,
mostly unicellular, (2–)6–10(–15) × (1–)2–5(–6) μm. When present,
chlamydospores globose to subglobose, hyaline.
Ilyonectria liriodendri (Halleen et al.) P. Chaverri & C.
Salgado, comb. nov. MycoBank MB518561.
Habitat: Generally on bark, sometimes causing cankers. Mostly in
temperate regions.
Habitat: On diseased roots and rootstocks.
Representative species: N. coccinea/C. candidum, N. ditissima/C.
heteronemum, N. faginata/C. faginatum, N. fuckeliana/C.
cylindroides var. tenue, N. hederae/C. hederae, N. major/
Cylindrocarpon sp., N. neomacrospora/C. cylindroides, N.
punicea/C. album, and N. ramulariae/C. obtusiusculum.
= Cylindrocarpon radicicola Wollenw., Fus. Autogr. Delin. 2: 651. 1924.
[= Ramularia macrospora Wollenw. Phytopathology 3: 222. 1913 non Fresen.,
Beitr. Mykol. 3: 88. 1863. hom. illeg.]
[= Fusarium polymorphum Marchal, Bull. Soc. Roy. Bot. Belgique 34: 145-148.
1895 non Matruchot, Rech. Dével. Mucéd. 84: 1892. hom. illeg.]
Habitat: On soil, roots, wood, and herbaceous debris.
Distribution: Cosmopolitan.
Basionym: Neonectria liriodendri Halleen, Rego & Crous, Stud.
Mycol. 55: 232. 2006.
Anamorph: "Cylindrocarpon" liriodendri J.D. MacDon. & E.E. Butler,
Pl. Dis. 65: 156. 1981.
www.studiesinmycology.org
71
chaverri et al.
Fig. 4. Neonectria. A, B. N. ditissima perithecia (A.R. 3690 = BPI 870951). C, D. N. fuckeliana perithecia (A.R. 3103 = BPI 842140). E. Top view of surface of N. fuckeliana
perithecium (A.R. 3103 = BPI 842140). F–H. Longitudinal section of N. ditissima perithecia (A.R. 3690 = BPI 870951). I. Asci and ascospores of N. ditissima (A.R. 3703 = BPI
871120). J. Paraphyses of N. ditissima (A.R. = BPI 871120). K. Asci and ascospores of N. ditissima (A.R. 3703 = BPI 871120). L, M. Asci and ascospores of N. fuckeliana (A.R.
3103 = BPI 842140). N–R. Conidiophores and macroconidia of N. ditissima (A.R. 3692 = CBS 119521 = BPI 871119). S–U. Conidiophores and microconidia of N. fuckeliana
(G.J.S. 02-67 = CBS 125109 = BPI 842434). Bars: A, C = 1 mm; B, D = 500 μm; E, I–U = 10 μm; F, G = 100 μm; H = 50 μm.
72
NeoNectria/cyliNdrocarpoN and related genera
Notes: Three names exist that could be considered synonyms of
Cylindrocarpon, i.e. Allantospora, Cylindrodendrum, and Heliscus
(Booth, 1966). The protologue and illustrations of Allantospora
suggest that it is probably not congeneric with Cylindrocarpon
(Wakker 1895). Wakker (1895) illustrated this genus based
on the type species, A. radicicola, as having Verticillium-like
conidiophores and small, allantoid conidia. Therefore, this
synonymy is doubtful. Cylindrodendrum could also be considered
a synonym of Cylindrocarpon, based on the cylindrical conidia
although many other genera in the Hypocreales have cylindrical
conidia. The original description and illustration of this genus based
on the type species, Cylindrodendrum album, shows that most
characteristics are quite distinct from Cylindrocarpon (Bonorden
1851). In Cylindrodendrum, the conidiophore branches have
sterile, terminal elongations that are generally hooked, phialides
with a swollen base and narrow neck, and relatively small conidia
that are not septate. Regarding Heliscus as a possible synonym of
Cylindrocarpon, Luo & Zhuang (2010b) placed the morphologically
similar species, Neonectria shennongjiana, close to Neonectria/
Cylindrocarpon sensu stricto. However, more studies are needed
to conirm if Heliscus is congeneric with Cylindrocarpon. If it is
congeneric, then, Heliscus is an older name (1880) and thus would
have priority over Cylindrocarpon. Due to the extensive use of the
name Cylindrocarpon, its economic importance, and some doubts
about the phylogenetic placement of Heliscus, the authors of the
present study would argue for conservation of Cylindrocarpon over
Heliscus.
RUGONECTRIA P. Chaverri & Samuels, gen. nov.
MycoBank MB518563. Fig. 5.
Etymology: “rugo” = Latin for “wrinkled”. The perithecial wall surface
for species of this genus is warted or rugose.
Type: Rugonectria rugulosa (Pat. & Gaill.) Chaverri, C. Salgado &
Samuels.
Ascomata supericialia vel gregaria in stromatae, ascomata globosa vel sublobosa,
verrucata vel tuberculata, rubra, KOH+ phaeorubra, non papillata. Ascosporae
ellipsoidea vel oblongata, 1-septatae, hyalinae vel pallide brunneae, striatae.
Anamorphosis Cylindrocarpon-similis. Phialide cylindrici. Macroconidia fusiformes,
hyaline, (3–)5–7(–9)-septatae, hilum inconspicue. Microconidia ellipsoidea vel
cylindrici, hyaline, 0–1-septatae, hilum inconspicue. Chlamydosporae absens.
Typus: R. rugulosa.
Teleomorph: Perithecia solitary or in groups, formed on or sometimes
partially immersed within a stroma. Perithecia globose to subglobose,
warted, non-papillate, orange to red, dark red in KOH+, yellow in lactic
acid. Perithecial wall 50–150 μm thick, generally of two indistinct
regions: outer region including warts with cells circular, 10–20 μm
diam, cell walls 3–4 μm thick, merging with surrounding stroma; inner
region with cells becoming progressively lattened, thinner, and less
pigmented toward locule. Ascospores ellipsoidal to oblong, striate,
hyaline, or sometimes yellowish, bicellular.
Anamorph: Cylindrocarpon-like; microconidia present; chlamydospores lacking. Macroconidia arising laterally from hyphae,
irregularly branched conidiophores or in fascicles, generally with a
short base. Phialides from macroconidiophores cylindrical, 15–25
× 3–5 μm. Macroconidia curved, fusiform, tapering towards ends,
(3–)5–7(–9)-septate, with inconspicuous hilum, (35–)48–85 × 5–10
www.studiesinmycology.org
μm. Microconidia produced from simple monophialidic or sparsely
branched conidiophores, scattered, ca. 20–100 μm long. Phialides
from microconidiophores cylindrical, 20–40 × 3–4 μm. Microconidia
ovoid to cylindrical, with rounded ends, generally blunt, 0–1-septate,
hyaline, (3–)5–15(–20) × 2–5 μm, lacking a prominent basal hilum.
Habitat: On bark of recently killed, dying or diseased trees, often
causing cankers.
Rugonectria castaneicola (W. Yamam. & Oyasu) Hirooka &
P. Chaverri, comb. nov. MycoBank MB518564.
Basionym: Nectria castaneicola W. Yamam. & Oyasu, Sci. Rep.
Hyogo Univ. Agric. Biol. 3: 17. 1957.
≡ Neonectria castaneicola (W. Yamam. & Oyasu) Tak. Kobay. & Hirooka,
J. Gen. Plant Pathol. 71: 126. 2005.
Anamorph: "Cylindrocarpon" castaneicola Tak. Kobay. & Hirooka,
J. Gen. Plant Pathol. 71: 126. 2005.
Habitat: On bark of conifers, generally causing cankers.
Distribution: Japan.
Description and illustrations: Kobayashi et al. (2005).
Rugonectria neobalansae (Samuels) P. Chaverri &
Samuels, comb. nov. MycoBank MB518565.
Basionym: Nectria neobalansae Samuels, Mem. N. Y. Bot. Gard.
59: 60. 1990.
Anamorph: Cylindrocarpon-like.
Habitat: On bark of living and recently killed trees.
Distribution: Indonesia, known only from the type locality.
Description and illustrations: Samuels et al. (1990).
Notes: Rugonectria neobalansae is distinct in being almost
completely immersed in an orange-red stroma and having, large
striate ascospores.
Rugonectria rugulosa (Pat. & Gaill.) Samuels, P. Chaverri &
C. Salgado, comb. nov. MycoBank MB518566.
Basionym: Nectria rugulosa Pat. & Gaill., Bull. Soc. Mycol. France
4: 115. 1888 [1889].
≡ Neonectria rugulosa (Pat. & Gaill.) Mantiri & Samuels, Canad. J. Bot.
79: 339. 2001.
= Nectria congoensis Sydow in Hennings in Wildeman Mycetes. Ann. Mus. Congo.
Bot. V. Études Syst. Geog. Bot. Flore du Bas- et du Moyen Congo 14. 1909.
Anamorph: "Cylindrocarpon" rugulosum Brayford & Samuels,
Sydowia 46: 146. 1994.
Habitat: On bark of living and recently killed trees, sometimes
causing cankers.
Distribution: Pantropical.
Descriptions and illustrations: Samuels et al. (1990), Samuels &
Brayford (1994).
73
chaverri et al.
Fig. 5. Rugonectria. A. Perithecia of R. neobalansae (G.J.S. 85-219, NY). B. Perithecia of R. rugulosa (G.J.S. 90-238 = BPI 1107399). C. Top view of surface of R. rugulosa
perithecium (G.J.S. 90-238 = BPI 1107399). D, E. Longitudinal section of R. neobalansae perithecium (G.J.S. 85-219, NY). F–H. Longitudinal section of R. rugulosa (G.J.S.
90-238 = BPI 1107399). I. Ascospores of R. neobalansae (G.J.S. 85-219, NY). J. Asci and ascospores of R. rugulosa (G.J.S. 90-238 = BPI 1107399). K, L. Conidiophores and
macroconidia of R. castaneicola (MAFF 237284). M. Conidiophores and macroconidia of R. rugulosa (G.J.S. 09-1337). N. Macroconidia of R. rugulosa (MAFF 241491). O.
Microconidia of R. castaneicola (MAFF 237284). P, Q. Conidiophores and microconidia of R. rugulosa (09-1337). Bars: A, B = 1 mm; C, I–Q = 10 μm; D–H = 100 μm.
74
NeoNectria/cyliNdrocarpoN and related genera
Fig. 6. Thelonectria. A. T. veuillotiana perithecia (A.R. 4505 = BPI 878946). B. T. discophora perithecia (A.R. 4499 = BPI 878945). C. T. jungneri perithecia (C.T.R. 71-244,
NY). D. T. lucida perithecia (C.T.R. 72-180, NY). E. T. veuillotiana perithecia (G.J.S. 90-48 = BPI 1107127). F. T. westlandica perithecia (G.J.S. 83-156, PDD). G. Top view of
surface of T. veuillotiana perithecium (A.R. 4505 = BPI 878946). H. Longitudinal section of T. discophora perithecium (A.R. 4499 = BPI 878945). I, J. Longitudinal section of T.
veuillotiana perithecium (G.J.S. 90-48 = BPI 1107127). K. Asci and ascospores of T. lucida (C.T.R. 72-180, NY). L, M. Conidiophores and conidia of T. veuillotiana on natural
substrate (G.J.S. 90-48 = BPI 1107127). N. Conidiophores and macroconidia of T. discophora (A.R. 4499 = BPI 878945). O. Conidiophores and macroconidia of T. olida (CBS
215.67). P. Conidiophores and macroconidia of T. veuillotiana (G.J.S. 90-48 = BPI 1107127). Q. Conidia of T. trachosa (CBS 112467). R. Macroconidia of T. westlandica (G.J.S.
83-156, PDD). S. Reverse colony of T. discophora on PDA (A.R. 4499 = BPI 878945). T. Reverse colony of T. veuillotiana on PDA (G.J.S. 90-48 = BPI 1107127). Bars: A–F =
500 μm; G, K–R = 10 μm; H, J = 50 μm; I = 100 μm.
www.studiesinmycology.org
75
chaverri et al.
THELONECTRIA P. Chaverri & C. Salgado, gen. nov.
MycoBank MB518567. Fig. 6.
Etymology: “thelo” – Greek for “nipple”. Many species in this
genus have a raised, papilla that is sometimes darkened, and thus
resembles a nipple.
Type species: Thelonectria discophora (Mont.) P. Chaverri & C.
Salgado (new combination made below).
Ascomata supericialia vel gregaria, ascomata globosa vel sublobosa, glabra,
rubra, KOH+ phaeorubra, atropapillata. Ascosporae ellipsoidea vel oblongata,
1-septatae, hyalinae, glabra. Anamorphosis Cylindrocarpon-similis. Phialide
cylindrici. Macroconidia fusiformes, curva, saepe triente apicali latiore, cellulis
apicalibus rotundatis et cellulis basalibus rotundatis vel complanatis, hyaline,
(3–)5–7(–9)-septatae, hilum inconspicue. Microconidia absens. Chlamydosporae
absens. Typus: T. discophora.
Teleomorph: Perithecia supericial, sometimes seated on an
immersed inconspicuous stroma, smooth or sometimes warted,
sometimes shiny, globose, subglobose, or pyriform to elongated,
300–600 μm diam, most species with a prominent, areolate
(darkened) papilla, if not, then at least with a darkly pigmented
apex; perithecial walls of 2 or 3 regions, 20–50(–100) μm thick:
outer region of intertwined hyphae or cells lacking a deinite outline
i.e. textura epidermoidea, with thickened, pigmented walls; inner
region of thin-walled, non-pigmented, lattened cells. Ascospores
mostly smooth, rarely spinulose or striate, hyaline, becoming
brownish at maturity, generally 1-septate.
Anamorph: Cylindrocarpon-like; microconidia rare, sometimes seen
on natural substrata; chlamydospores rare, abundant in one species;
conidiophores arising laterally from hyphae, irregularly branched
conidiophores or forming fascicles; phialides cylindrical, 10–25 × 3–6
μm; macroconidia curved, often broadest at upper third, with rounded
apical cells and lattened or rounded basal cells, (3–)5–7(–9)-septate,
with inconspicuous hilum, (35–)40–90(–110) × 4–8(–11) μm.
Habitat: On bark of recently killed, dying or diseased trees, often
causing small cankers, sometimes on rotting roots.
Thelonectria coronata (Penz. & Sacc.) P. Chaverri & C.
Salgado, comb. nov. MycoBank MB518568.
Basionym: Nectria coronata Penz. & Sacc., Malpighia 11: 510.
1897.
≡ Neonectria coronata (Penz. & Sacc.) Mantiri & Samuels, Canad. J. Bot.
79: 339. 2001.
Anamorph: "Cylindrocarpon" coronatum Brayford & Samuels,
Sydowia 46: 91. 1993.
Habitat: On bark, often associated with small cankers.
= Nectria tasmanica Berk. in Hooker, Flora Tasmaniae 2: 279. 1860.
= Nectria mammoidea W. Phillips & Plowr. Grevillea 3: 126. 1875.
≡ Creonectria mammoidea (W. Phillips & Plowr.) Seaver, Mycologia 1:
188. 1909 (as Creonectria mammoides).
= Nectria nelumbicola Henn., Verh. Bot. Vereins. Prov. Brandenburg 40: 151. 1898.
= Nectria umbilicata Henn., Hedwigia 41: 3. 1902.
= Nectria mammoidea var. rugulosa Weese, Akad. Wiss. Wien Math.-Naturw.
Kl., Abt. 1, 125: 552. 1916.
= Nectria mammoidea var. minor Reinking, Zentralbl. Bakteriol., Abt. 2, 94:
135. 1936.
= Creonectria discostiolata Chardón, Bol. Soc. Venez. Ci. Nat. 5: 341. 1939.
= Nectria pinea Dingley, Trans. Roy. Soc. New Zealand 79: 198. 1951.
Anamorph: "Cylindrocarpon" ianothele var. majus Wollenw., Z. Parasitenk.
(Berlin) 1: 161. 1928.
= Cylindrocarpon ianthothele var. minus Reinking, Zentralbl. Bakteriol., Abt.
2, 94: 135. 1936.
= Cylindrocarpon ianthothele var. rugulosum C. Booth, Mycol. Pap. 104: 25. 1966.
= Cylindrocarpon pineum C. Booth, Mycol. Pap. 104: 26. 1966.
Habitat: On bark and twigs of recently killed trees, rarely on palm
trunks.
Distribution: Cosmopolitan.
Description and illustrations: Brayford et al. (2004).
Thelonectria jungneri (Henn.) P. Chaverri & C. Salgado,
comb. nov. MycoBank MB518570.
Basionym: Nectria jungneri Henn., Bot. Jahrb. Syst. 22: 75. 1897.
= Nectria eustoma Penz. & Sacc., Malpighia 11: 509. 1898 [1897]
= Nectria leucoloma Starbäck, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 25:
28. 1899.
= Nectria cinereopapillata Henn. & Nyman in Warburg, Monsunia 1: 161. 1900
[1899]
= Nectria striatospora Zimm., Centralbl. Bakteriol. II, 7: 105. 1901.
= Nectria azureostiolata Doi, Mem. Nat. Sci. Mus. Tokyo 10: 23. 1977.
Anamorph: "Cylindrocarpon" victoriae Wollenw., Z. Parasitenk.
(Berlin) 1: 161. 1928.
Habitat: On bark of recently killed or dying trees.
Distribution: Pantropical.
Description and illustrations: Samuels et al. (1990).
Thelonectria lucida (Höhn.) P. Chaverri & C. Salgado,
comb. nov. MycoBank MB518571.
Basionym: Nectria lucida Höhn., Akad. Wiss. Wien math. Naturw.
Kl., Abt. 1, 118: 289. 1909.
≡ Neonectria lucida (Höhn.) Samuels & Brayford, Mycologia 96: 590.
2004.
Anamorph: "Cylindrocarpon" lucidum Booth, Mycol. Pap. 104: 21. 1966.
Habitat: On bark of recently killed or dying trees, rarely on vines.
Distribution: Probably pantropical.
Distribution: Asia, New Zealand, South America, North America,
probably cosmopolitan.
Descriptions and illustrations: Brayford & Samuels (1993); Samuels
& Brayford (1994)
Description and illustrations: Brayford et al. (2004).
Thelonectria discophora (Mont.) P. Chaverri & C. Salgado,
comb. nov. MycoBank MB518569.
Basionym: Sphaeria discophora Mont., Ann. Sci. Nat. Bot. II 3: 353.
1835.
≡ Neonectria discophora (Mont.) var. discophora Mantiri & Samuels,
Canad. J. Bot. 79: 339. 2001.
76
Thelonectria olida (Wollenw.) P. Chaverri & C. Salgado,
comb. nov. MycoBank MB518572.
Basionym: Ramularia olida Wollenw., Phytopathology 3: 223. 1913.
≡ Cylindrocarpon olidum var. olidum (Wollenw.) Wollenw., Fus. Autogr.
Del., ed. 1: 471. 1916.
= Cylindrocarpon curvatum Hochapfel in Wollenw., Z. Parasitenk. 3: 495.
1931.
NeoNectria/cyliNdrocarpoN and related genera
Teleomorph: Unknown.
Habitat: On rotting roots of various plants.
Distribution: Probably widespread.
Thelonectria westlandica (Dingley) P. Chaverri & C.
Salgado, comb. nov. MycoBank MB518576.
Basionym: Nectria westlandica Dingley, Trans. Roy. Soc. New
Zealand 79: 201. 1951.
≡ Neonectria westlandica (Dingley) Samuels & Brayford, Mycologia 96:
595. 2004.
Descriptions and illustrations: Booth (1966), Brayford (1987).
Anamorph: Cylindrocarpon-like.
Notes: This species is somewhat different from the rest of
Thelonectria in having shorter macroconidia, fewer septa, and
abundant chlamydospores. However, it also has similarities with
Thelonectria. Thelonectria olida has short conidiophores, lacks
microconidia, and has curved macroconidia with rounded ends.
Molecular phylogenetic data presented here also places this
species in Thelonectria.
Habitat: On bark of dicotyledonous trees, sometimes gymnosperms.
Thelonectria trachosa (Samuels & Brayford) Samuels, P.
Chaverri & C. Salgado, comb. nov. MycoBank MB518573.
Basionym: Neonectria trachosa Samuels & Brayford, Mycologia
96: 592. 2004.
Anamorph: Cylindrocarpon-like
Distribution: New Zealand.
Description and illustrations: Brayford et al. (2004)
ACKNOWLEDGEMENTS
We appreciate the nomenclatural advice given by Drew Minnis (USDA). We also
greatly appreciate the assistance of Adam Bazinet (UMD) in the use of the GRIDcomputing system. This study was funded by a grant from United States National
Science Foundation (PEET program) DEB-0925696: “Monographic Studies in the
Nectriaceae, Hypocreales: Nectria, Cosmospora, and Neonectria” to University of
Maryland (P. Chaverri, G.J. Samuels & A.Y. Rossman).
Habitat: On bark of unknown conifer.
Distribution: Scotland, only known from the type locality.
Description and illustrations: Brayford et al. (2004).
Thelonectria veuillotiana (Sacc. & Roum.) P. Chaverri & C.
Salgado, comb. nov. MycoBank MB518574.
Basionym: Nectria veuillotiana Sacc. & Roum. in Désmazières,
Rev. Mycol. (Toulouse) 2: 189. 1880.
≡ Dialonectria veuillotiana (Sacc. & Roum.) Cooke, Grevillea 12: 110.
1884.
≡ Cucurbitaria veuillotiana (Sacc. & Roum.) Kuntze, Revis. Gen. Pl.
(Leipzig) 3: 462. 1898.
≡ Neonectria veuillotiana (Sacc. & Roum.) Mantiri & Samuels, Canad. J.
Bot. 79: 339. 2001.
= Sphaerostilbe sanguinea Fuckel, Symb. Myc. App. 3: 22. 1877.
Anamorph: "Cylindrocarpon" candidulum (Sacc.) Wollenw., Z.
Parasitenk. 1: 160. 1928.
≡ Atractium candiduli Sacc., Syll. Fung. (Abellini) 2: 512.1883.
Habitat: On bark of recently killed trees, rarely on wood or leaves.
Distribution: Probably widespread.
Description and illustrations: Brayford & Samuels (1993).
Thelonectria viridispora (Samuels & Brayford) P. Chaverri,
C. Salgado, & Samuels, comb. nov. MycoBank MB518575.
Basionym: Neonectria viridispora Samuels & Brayford, Mycologia
96: 592. 2004.
Anamorph: Cylindrocarpon-like.
Habitat: On bark of Ochroma.
Distribution: Ecuador, only known from the type locality.
Description and illustrations: Brayford et al. (2004).
www.studiesinmycology.org
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Priscila Chaverri