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http://dx.doi.org/10.1007/s40858-017-0201-1
AUTHOR'S PROOF
Metadata of the article that will be visualized in OnlineFirst
Article Title Notes on powdery mildews (Erysiphales) in Thailand V.
Golovinomyces
Article Sub-Title
Article Copyright Year Sociedade Brasileira de Fitopatologia 2017
(This will be the copyright line in the final PDF)
Journal Name Tropical Plant Pathology
Corresponding Author
Family Name Meeboon
Particle
Given Name Jamjan
Suffix
Organization Mie University, Department of Bioresources,
Graduate School
Address 1577 Kurima-Machiya, Tsu 514-8507, Japan
e-mail jamjanm@yahoo.com
Author
Family Name Kokaew
Particle
Given Name Jitra
Suffix
Division Department of Agriculture
Organization Ministry of Agriculture and Cooperatives
Address 50 Phaholyothin Rd., Ladyao, Chatuchak,
Bangkok, Thailand
Author
Family Name Takamatsu
Particle
Given Name Susumu
Suffix
Organization Mie University, Department of Bioresources,
Graduate School
Address 1577 Kurima-Machiya, Tsu 514-8507, Japan
Schedule
Received 7 July 2017
Revised
Accepted 24 October 2017
Abstract Records of Golovinomyces species new to Thailand are described on
the hosts Ageratum conyzoides,Bidens pilosa,Dahlia pinnata,D. ×
hortensis,Helianthus annuus,Lactuca indica, Laggera crispata,Sonchus
oleraceus (Asteraceae), Lygisma inflexum (Asclepiadaceae), Myosotis
scopioides (Boraginaceae), Coccinia indica,Coccinia grandis
(Cucurbitaceae), Vigna umbellata (Fabaceae), Torenia fournieri
AUTHOR'S PROOF
(Linderniaceae), Plantago major (Plantaginaceae) and Verbena ×
hybrida (Verbenaceae). The identifications of the particular
Golovinomyces species have been performed by means of
morphological examinations supplemented by molecular sequence
analyses. On the basis of molecular analyses, the powdery mildew on
Ocimum tenuiflorum (Lamiaceae) proved to represent a species of its
own, which is referred to as Golovinomyces ocimi comb. nov. The
application of Oidium ocimi, the basionym of this combination, is
determined by lecto- and epitypification. Lygisma inflexum,Laggera
crispata and Vigna umbellata are new host records for Golovinomyces
worldwide.
Keywords (separated
by '-')
Anamorph·biodiversity - Erysiphaceae - Molecular phylogeny - Tropics
Foot note information Section Editor: Meike Piepenbring
AUTHOR'S PROOF
UNCORRECTEDPROOF
1
2
3ORIGINAL ARTICLE
4Notes on powdery mildews (Erysiphales) in Thailand
5V. Golovinomyces
6Jamjan Meeboon
1
&Jitra Kokaew
2
&Susumu Takamatsu
1
7
8Received: 7 July 2017 /Accepted: 24 October 2017
9#Sociedade Brasileira de Fitopatologia 2017
10 Abstract Records of Golovinomyces species new to Thailand
11 are described on the hosts Ageratum conyzoides,Bidens
12 pilosa,Dahlia pinnata,D. ×hortensis,Helianthus annuus,
13 Lactuca indica, Laggera crispata,Sonchus oleraceus
14 (Asteraceae), Lygisma inflexum (Asclepiadaceae), Myosotis
15 scopioides (Boraginaceae), Coccinia indica,Coccinia grandis
16 (Cucurbitaceae), Vigna umbellata (Fabaceae), Tore n i a
17 fournieri (Linderniaceae), Plantago major (Plantaginaceae)
18 and Ve r b en a ×hybrida (Verbenaceae). The identifications of
19 the particular Golovinomyces species have been performed by
20 means of morphological examinations supplemented by mo-
21 lecular sequence analyses. On the basis of molecular analyses,
22 the powdery mildew on Ocimum tenuiflorum (Lamiaceae)
23 proved to represent a species of its own, which is referred to
24 as Golovinomyces ocimi comb. nov. The application of
25 Oidium ocimi, the basionym of this combination, is deter-
26 mined by lecto- and epitypification. Lygisma inflexum,
27 Laggera crispata and Vigna umbellata are new host records
28 for Golovinomyces worldwide.
29 Keywords Anamorph·biodiversity .Erysiphaceae .
30 Molecular phylogeny .TropicsQ1
31Introduction Q2
32Golovinomyces (U. Braun) Heluta [type species: Erysiphe
33cichoracearum DC. (≡Golovinomyces cichoracearum (DC.)
34Heluta)] is a genus of powdery mildew having polyascal ascoma
35(chasmothecia) with mycelioid appendages and catenescent co-
36nidia without fibrosin bodies (Braun and Takamatsu 2000;
37Matsuda and Takamatsu 2003). This genus is divided into two
38sections, sect. Golovinomyces and sect. Depressi (U. Braun) U.
39Braun. Golovinomyces is a large genus comprising 45 species
40(Braun and Cook 2012) with an almost worldwide distribution.
41The host range of this genus is mostly restricted to herbaceous
42plants, including up to 2283 species from 58 families such as the
43Asteraceae, Boraginaceae, Scrophulariaceae, and Cucurbitaceae
44(Amano 1986). The taxonomic history of the genus was de-
45scribed in Braun and Cook (2012) in detail. Investigations of
46powdery mildews in Southeast Asia in the past 14 years, in
47particular in Thailand, have contributed to the discovery of new
48taxa and new records mainly belonging to the tribe Erysipheae
49(for example, Meeboon et al. 2016; Meeboon and Takamatsu
502016,2017a,b,c,d). In this paper, three hosts of powdery mil-
51dews are reported that are new worldwide, one new combination
52is introduced, and 11 host-fungus combinations of the powdery
53mildew species (Golovinomyces ambrosiae,G. cynoglossi,
54G. orontii,G. sonchicola,andG. sordidus)newforThailand
55are listed. The identifications of the species concerned were ver-
56ified by phylogenetic methods, i.e. molecular sequence analyses.
57Materials and methods
58Morphological examination
59Morphological examinations were carried out as outlined in
60Meeboon and Takamatsu (2015a). All the specimens were
Section Editor: Meike Piepenbring
*Jamjan Meeboon
jamjanm@yahoo.com
1
Mie University, Department of Bioresources, Graduate School, 1577
Kurima-Machiya, Tsu 514-8507, Japan
2
Department of Agriculture, Ministry of Agriculture and
Cooperatives, 50 Phaholyothin Rd., Ladyao, Chatuchak,
Bangkok, Thailand
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61 examined using a light microscope with phase contrast 10×,
62 20× and 40× objectives. Herbarium samples were rehydrated
63 before examination by boiling a small piece of infected leaf
64 with the fungal mycelium downwards in a drop of lactic acid
65 on a slide (Shin and La 1993). After boiling, the rehydrated
66 mycelium was scraped off and studied in lactic acid using a
67 light microscope. Thirty conidia and conidiophores were mea-
68 sured for each specimen. Herbarium specimens were deposit-
69 ed at Mie University Mycological Herbarium (TSU-MUMH),
70 Japan.
71 The nucleotide sequences of the rDNA internal transcribed
72 spacer (ITS) region including 5.8S rDNA were determined in
73 accordance with Meeboon and Takamatsu (2015b).
74 Representative new sequences determined in this study were
75 deposited in DNA Data Base of Japan (DDBJ) under the ac-
76 cession numbers LC306656–LC306669. These sequences
77 were aligned with closely related sequences of the
78 Erysiphaceae using MUSCLE (Edgar 2004)implementedin
79 MEGA version 6 (Tamura et al. 2013). Alignment was further
80 manually refined using MEGA, and deposited in TreeBASE
81 (http://www.treebase.org/) under the accession number
82 S21253. Phylogenetic trees were obtained from the datasets
83 by using the maximum parsimony (MP) method implemented
84 in PAUP* 4.0b10 (Swofford 2002) according to the proce-
85 dures of Meeboon and Takamatsu (2016).
86 Molecular phylogeny
87 Whole cell DNAwas extracted from mycelia using the chelex
88 method (WalshQ3 et al. 1991) as described in Hirata
Q
4=and
89 Takamatsu (1996). The respective primer pairs of PM5/ITS4
90 and ITS5/PM6 (TakamatsuQ5 and Kano 2001) were used to am-
91 plify ITS fragment 1 and fragment 2, respectively. PCR reac-
92 tion was conducted using KOD FX NeoDNA polymerase
93 (Toyobo, Japan) according to the manufacturer's protocol.
94 The PCR product was sent to SolGent Co. Ltd. (Daejeon,
95 South Korea) for sequencing using primer pair of ITS1 and
96 ITS4 (WhiteQ6 et al. 1990).
97 New representative sequences determined in this study
98 were deposited in DNA Data Base of Japan (DDBJ) under
99 the accession numbers LC163909, LC163911, LC163913,
100 LC163917 and LC163922. Sequences generated in this study
101 were aligned with other sequences of Golovinomyces re-
102 trieved from DNA databases (DDBJ, EMBL, NCBI) using
103 MUSCLE (Multiple Sequence Comparison by Log
104 Expectation) (Edgar 2004) implemented in MEGA 6
105 (Tamura et al. 2013). The alignments were deposited in
106 TreeBASE (http://www.treebase.org/) under the accession
107 number S20344.
108 Phylogenetic trees were obtained from the data set using
109 the maximum parsimony (MP) method performed in PAUP*
110 4.0b10 (Swofford 2002) with the heuristic search option using
111 the tree bisection reconstruction (TBR) algorithm. This search
112was repeated 100 times with different random starting points,
113using the stepwise addition option to increase the likelihood of
114finding the most parsimonious tree. All sites were treated as
115unordered and unweighted, with gaps treated as missing data.
116Tree scores, including tree length, consistency index (CI),
117retention index (RI) and rescaled consistency index
118(RC), were calculated. The strength of the internal
119branches of the resulting trees was tested with bootstrap
120(BS) analysis (Felsenstein Q71985) using 1000 replications
121with the stepwise addition option set to simple and a
122maximum tree number of 100.
123Results
124Phylogenetic analyses
125Fourteen ITS sequences of Golovinomyces species deter-
126mined in this study were aligned with other Golovinomyces
127sequences retrieved from DNA databases. Arthrocladiella
128mougeotii (Lév.) Vassilkov (AB329690) was used as outgroup
129taxon. Of the 510 total characters, 376 were constant, 46 were
130variable but parsimony-uninformative, and 88 were parsimo-
131ny-informative. The MP analysis produced about 115K equal-
132ly parsimonious trees with 284 steps. Topologies were almost
133consistent among the trees except for branching orders of the
134terminal branches and branch length. A typical tree is shown
135in Fig. 1.
136Taxono m y
137Golovinomyces ocimi (S. Naray. & K. Ramakr.) Meeboon &
138S. Takam., comb. nov. Fig. 2.
139MycoBank no.: MB821878
140Basionym: Oidium ocimi S. Naray. & K. Ramakr., Madras
141Univ. J. 37–38: 87, 1967.
142Lectotype (designated here, MycoBank, (MBT377568 Q8):
143Fig. 15 in Narayanaswami & Ramakrishnan (Narayanaswami
144and Ramakrishnan 1967: 87). Epitype (designated here,
145MycoBank, MBT377567): on Ocimum tenuiflorum L.
146(Lamiaceae), THAILAND, Chiang Rai, Wiangpapao, 15
147December 2015, J. Meeboon (TNS-F-xxxxx), TSU-
148MUMH6621 (isoepitype).
149Gene sequence (ex epitype): LC306657 (ITS).
150Mycelium amphigenous, effuse or in irregular patches, al-
151most persistent or evanescent. Hyphae hyaline, walls thin,
152smooth or almost so, 4–9μm wide. Hyphal appressoria nip-
153ple-shaped, sometimes poorly developed. Conidiophores ter-
154minal on the surface of mother cells, 70–173 μm long, often
155increasing from base to top, erect, straight or curved at the
156base. Foot cells 27–42 × 7–11 μm, basal septum of the foot
157cell mostly raised, 5–15 μm above the junction with the moth-
158er cell, foot cell followed by 1–3 shorter cells, forming
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Fig. 1 Phylogenetic analysis of
the rDNA ITS1-5.8S-ITS2 for 63
sequences from Golovinomyces
spp. BS (>70%) values by the
maximum parsimony (MP) are
shown on the branches. Newly
determined sequences were
shown as boldface
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159 catenescent conidia. Conidia doliiform, 32–40 × 15–20 μm,
160 germ tubes subapically inserted, non to one-septate, mostly
161 short to moderately long, terminating simply or in a club-
162 shaped appressorium.
163 Further collections examined –on Ocimum tenuiflorum L.
164 (Lamiaceae), THAILAND, Chiang Mai, Su Thep, 29
165 December 2007, J. Meeboon, MUMH1803; Chiang Rai,
166 Wiangpapao, 25 December 2016, J. Meeboon, MUMH6892;
167 Chiang Rai, Mae Suai, 29 December 2016, J. Meeboon & S.
168 Takamatsu, MUMH6903, MUMH6942.
169 Notes –Ocimum tenuiflorum, commonly known as holy
170 basil,is an aromatic plant which is native to the Indian sub-
171 continent and widespread as a cultivated plant throughout the
172 Southeast Asian tropics. It is used in Thai cuisine as Thai holy
173 basil (kaphrao). Golovinomyces biocellatus (Ehrenb.) Heluta
174 was reported to occur on many host species of Lamiaceae,
175 including Ocimum spp., in Europe, Asia, North and South
176 America (Braun and Cook 2012Q9 ). Narayanaswami and
177 Ramakrishnan (1967) described Oidium ocimi on
178 O. tenuiflorum (= O. sanctum) from India. The original de-
179 scription is rather brief and too insufficient to determine the
180taxonomic position of this asexual powdery mildew.
181Therefore, Braun Q10(1987) listed O. ocimi under “Anamorphs
182of uncertain position”. Bappammal et al. (1995) collected
183additional Indian specimens of O. ocimi and provided a
184more detailed description and illustration, suggesting a
185close affinity of this species to Golovinomyces biocellatus.
186This illustration was again published in Hosagoudar and
187Agarwal (2009). Furthermore, Amano (1986) listed
188O. tenuiflorum (= O. sanctum) as host species of Erysiphe
189biocellata from India. Therefore, Braun and Cook (2012)re-
190duced O. ocimi to synonymy with G. biocellatus. Identification
191and application of the name O. ocimi require a clarification of
192this species via lecto- and epitypification. Type material of this
193species could not be traced and is probably not preserved.
194Bappammal et al. (1995) also failed to locate and examine type
195material. Therefore, the original illustration published in
196Narayanaswami and Ramakrishnan (1967), which is part of
197the original material in the sense of the ICN, has to be taken
198into consideration for lectotypification (see Art. 9.2, 9.3). An
199epitype (new material with corresponding sequence data) is
200proposed to allow a phylogenetic analysis of O. ocimi which
Fig. 2 Golovinomyces ocimi on
Ocimum tenuiflorum
(MUMH6621). a. Germinating
conidia. b.Conidia.c.
Conidiophores. d. Appressoria.
Bar = 20 μm
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201 is essential for a taxonomic reassessment of this species.
202 The G. biocellatus complex has recently been divided
203 into four species (Scholler et al. 2016), viz, G. biocellatus,
204 G. monardae,G. neosalviae,andG. salviae. The present col-
205 lections from Thailand on Ocimum tenuiflorum are morpho-
206 logically similar to these Golovinomyces species but differ in
207 having relatively short conidiophores foot cells, 27–42 μm
208 long, vs. 40–100(−140) μm long in G. monardae,55–
209 100(−130) μm long in G. biocellatus,30–100(−120) μmlong
210 in G. salviae,45–75(−115) μm long in G. neosalviae.
211 Furthermore, two sequences retrieved from powdery mildew
212 on O. tenuiflorum formed a distinct monophyletic clade. This
213 clade grouped with the clades of G. biocellatus,G. monardae,
214 G. neosalviae,andG. salviae with 100% BS value. The num-
215 ber of nucleotide differences of Golovinomyces on
216 O. tenuiflorum was 3/486 characters (99.3% similarity, not
217 including gaps) from G. biocellatus on Glechoma hederacea
218 (LC076805), 3/485 characters (99.3 % similarity) from
219 G. monardae on Monarda citriodora (LC076809), 4/485
220 characters (99.1 % similarity) from G. neosalviae on Salvia
221 lavandulifolia (LC076838), and 4/486 characters (99.1 %
222 similarity) from G. salviae on Salvia nemorosa (LC100001).
223 These results suggest that the powdery mildew on
224 O. tenuiflorum represents a species of its own morpholog-
225 ically and phylogenetically different from the allied
226 Golovinomyces species on Glechoma,Mentha,Origanum,
227Rosmarinus,Salvia and Thymus, supported by the
228phylogenetic-taxonomic affinity and position of the host
229plant within the Lamiaceae [subfam. Nepethoideae tribe
230Ocimeae] (Walker et al. 2004). ICN (International Code
231of Nomenclature for algae, fungi, and plants), Art. 59.1, is
232applicable to this fungus. Thus, although its sexual morph is
233still unknown, it is assignable to Golovinomyces, which has
234priority over the anamorph-typified genus Euoidium that is
235now a heterotypic synonym of Golovinomyces (Rossman
236et al. 2016), and the name Oidium ocimi is used for this
237species (see above).
238The Golovinomyces ambrosiae clade (III sensu
239Takama t s u e t a l. 2013)
240This genetic assemblage is heterogeneous and unresolved. It
241comprises at least two morphologically distinguishable taxa
242that have been described as Golovinomyces ambrosiae,con-
243fined to Ambrosia,Helianthus,Iva, and Rudbeckia spp. as
244well as Zinnia angustifolia, all belonging to the composite
245tribe Heliantheae (Braun and Cook 2012). This species is
246characterized by having broadly ellipsoid-ovoid, doliiform to
247limoniform conidia, 25–45 × 15–27 μmwhenfresh,witha
248length/width ratio < 2, 1.3–1.9, mosty 1.4–1.6, and dimorphic
249germ tubes with a high percentage of longitubus pattern within
250the Euoidium type. The other taxon involved in this clade has
Fig. 3 Golovinomyces
ambrosiae on Helianthus annuus
(MUMH6871). a.Conidia.b.
Conidiophores. c. Appressoria.
Bar = 25 μm
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251been referred to as Golovinomyces spadiceus, which differs
252from G. ambrosiae in having narrower conidia, 25–40 × 14–
25320 μm, with a length/width ratio of 1.5–2andaEuoidium type
254of the conidial germination, lacking longitubus pattern germ
255tubes. G. spadiceus occurs on many hosts belonging to tribe
256Heliantheae, including species of the genera Coreopsis,
257Dahlia,Xanthium,andZinnia, but is undoubtedly plurivorous
258with various hosts belonging to genera of other composite
259tribes and possibly even to some non-composite hosts. The
260situation is further complicated by the fact that some of the
261hosts involved in this complex are colonized by several
262Golovinomyces species, e.g., Dahlia spp. and Helianthus
263spp. are also hosts of G. orontii and Zinnia spp. are known
264to be hosts of G. ambrosiae as well as G. spadiceus (Braun
265and Cook 2012). It cannot be excluded that even Helianthus
266spp. might be hosts of G. spadiceus. Lineage III in Takamatsu
267et al. (2013) is probably a complex of several species. ITS
268sequences are not sufficient for a reliable resolution and dif-
269ferentiation on species level. Furthermore, the application of
270the species names involved can only be considered to be ten-
271tative since Erysiphe ambrosiae and E. spadicea were de-
272scribed from North America on Ambrosia sp. and Xanthium
273sp., respectively, but reference sequences and epitypes based
274on North American samples are not yet available. The appli-
275cation of the name G. circumfusus (Schltdl.) U. Braun to a
Fig. 4 Golovinomyces cynoglossi on Myosotis scopioides (MUMH1825).
aConidiophore. bConidia. cGerminating conidium. dAppressoria.
Bar = 20 μm
Fig. 5 Golovinomyces orontii on
Coccinia grandis (MUMH6613).
aConidia. bConidiophores.
Bar = 20 μm
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276 sequence retrieved from a Japanese sample on Eupatorium
277 chinense in lineage III in Takamatsu et al. (2013)isalsoun-
278 clear and unproven since reference sequences obtained on the
279 basis of European collections on Eupatorium cannabinum
280 (type of Erysiphe circumfusa) are also still lacking. Up until
281 a comprehensive genetic re-examination of the whole
282 G. ambrosiae complex based on additional markers and
283 epitypifications of the taxa involved, collections with
284 sequences pertaining to lineage III can currently only
285 tentatively be assigned to G. ambrosiae and G. spadiceus
286 just based on morphology, as for instance done by Dugan
287 (2013) in the case of Golovinomyces on Coreopsis.Inthis
288 sense, Golovinomyces specimens recently collected in
289 Thailand on Dahlia pinnata,D. ×hortensis, and Laggera
290 crispata canbeassignedtoG. spadiceus, and collections on
291 Helianthus annuus to G. ambrosiae.Golovinomyces on
292 Ver b en a ×hybrida can only tentatively be assigned to
293 G. spadiceus (conidiophores and conidia of the sample from
294 Thailand are barely distinguishable from G. verbenae, but
295 comparative sequence data retrieved from North American
296collection on Ve r b en a are not yet available, i.e. the phyloge-
297netic position of the latter species is still unknown). The iden-
298tity of Golovionomyces on Ageratum conyzoides and Bidens
299pilosa in Thailand is unclear, since they are clearly different
300from G. ambrosiae andprobablyalsodistinctfrom
301G. spadiceus, and can currently only be classified as
302Golovinomyces sp. Further research is necessary. For the fun-
303gus on Ageratum conyzoides (Eupatorieae), the name
304Euoidium agerati (J.M. Yen) U. Braun & R.T.A. Cook, de-
305scribed on this host from Taiwan, is available (Braun and
306Cook 2012).
307Golovinomyces ambrosiae (Schwein.) U. Braun & R.T.A.
308Cook, in Cook & Braun, Mycol. Res. 113(5): 628, 2009
309Mycelium amphigenous, mainly epiphyllous, effuse or
310forming patches, thin, white, persistent or almost so; hyphae
311hyaline, walls thin, smooth, 4–8μm wide. Hyphal appressoria
312nipple-shaped, solitary or in opposite pairs. Conidiophores
313arising centrally or towards one end of hyphal mother cells
314and from their upper surface, erect, straight, 177–298 μm
315long. Foot cells cylindrical, straight in the base, 71–129 ×
Fig. 6 Golovinomyces orontii on
Lactuca indica (MUMH6936).
aConidia. bConidiophores.
Bar = 15 μm
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316 11–18 μm, followed by 1–3 shorter cells, forming catenescent
317 conidia. Conidia broadly ellipsoid-ovoid, doliiform to some-
318 what limoniform, 39–60 × 23–27 μm, germ tubes terminal to
319 subterminal, tips not swollen or only slightly swollen (Fig. 3).
320 Material examined –On Helianthus annuus L.
321 (Asteraceae), Chiang Rai, Mae Suai, 30 December 2016, J.
322 Meeboon & S. Takamatsu, MUMH6871; Chiang Mai, Mae
323 Rim, Mon Jam, 19 January 2016, J. Meeboon, MUMH6683.
324 Notes –Golovinomyces ambrosiae and G. orontii have
325 been recorded on Helianthus spp. (Braun and Cook 2012).
326 These two species differ from each other by the shape of foot
327 cells; G. ambrosiae has straight in the base of foot cells,
328 G. orontii has curved foot cells. The morphological
329 characteristics of the current specimen are more similar
330 to G. ambrosiae than to G. orontii due to having straight foot
331 cells and broad conidia, 25–40 × (10–)15–23(–25) μmin
332 G. orontii vs. 25–45 × 15–27 μminG. ambrosiae (Braun
333 and Cook 2012).This is the first report of G. ambrosiae on
334 Helianthus annuus from Thailand.
335 Golovinomyces cynoglossi (Wallr.) Heluta, Ukrayins’k.
336 Bot. Zhurn. 45(5): 62, 1988
337 Mycelium amphigenous, dense, effuse or in patches, eva-
338 nescent to almost persistent. Hyphae hyaline, thin-walled,
339 smooth, 3.5–7μm wide. Hyphal appressoria nipple-shaped.
340 Conidiophores erect, arising from upper surface of hyphal
341 mother cells, 90–160(–200) μm long. Foot cells straight, cy-
342 lindrical, 30–110 × 10–13 μm, followed by 1–2(–3) shorter
343cells, forming catenescent conidia; conidia ellipsoid-ovoid,
34433–45 × 18–25 μm, germ tubes arising from an end, moder-
345ately long, apex often with somewhat swollen appressorium
346as in Euoidium type (Fig. 4).
347Material examined –on Myosotis scopioides L.
348(Boraginaceae), THAILAND, Chiang Mai, Su Thep, 5
349January 2005, J. Meeboon, MUMH1825.
350Notes –Braun and Cook (2012) described G. cynoglossi on
351many host species of various host genera of Boraginaceae includ-
352ing Myosotis throughout Europe, Asia, North and South Africa
353and North America. The asexual morph found on Myosotis
354scopioides agrees completely with G. cynoglossi and is the first
355record of powdery mildew on M. scopioides in Thailand.
356Golovinomyces orontii (Castagne) Heluta, Ukrayins’k.
357Bot. Zhurn. 45(5): 63, 1988
358Mycelium amphigenous, mainly epiphyllous, effuse or in
359patches, evanescent or persistent, white; hyphae slightly flex-
360uous, branched at right angles. Hyphal appressoria nipple-
361shaped, often poorly developed. Conidiophores erect, arising
362laterally or from the upper surface of hyphal mother cells and
363almost centrally or towards one end of the cell. Foot
364cells straight or curved in the basal half, followed by
3652–3 shorter cells, forming catenescent conidia. Conidia
366ellipsoid, doliiform, subcylindrical, germ tubes arising
367from an end, occasionally from a side, straight, bent,
368rarely forked, apically often with a somewhat swollen
369appressorium, Euoidium type.
Fig. 7 Golovinomyces orontii on
Torenia fournieri (MUMH6881).
aConidiophores. bConidia.
cGerminating conidia.
Bar = 15 μm
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370 Golovinomyces orontii ex Coccinia grandis
371 Conidiophores erect, arising laterally or the upper surface of
372 hyphal mother cells, 130–200 μm long. Foot cells curved,
373 rarely straight, 60–87 × 13–16 μm, followed by 1–3shorter
374 cells, forming catenescent conidia. Conidia subcylindrical,
375 36–44 × 18–20 μm(Fig.5).
376 Golovinomyces orontii ex Lactuca indica
377 Conidiophores erect, usually arising laterally from the hyphal
378 mother cell but occasionally from its upper surface, 100–
379 210 μm long. Foot cells straight to curved at the base, 36–
380 61 × 5–10 μm, followed by 1–3 shorter cells, forming
381 catenescent conidia. Conidia doliiform to subcylindrical, 28–
382 32 × 12–14 μm(Fig.6).
383Golovinomyces orontii ex Torenia fournieri
384Conidiophores erect, arising from the upper surface of hyphal
385mother cells, 157–290 μm long. Foot cells straight or curved,
38626–124 × 9.5–13 μm, followed by 1–3 shorter cells, forming
387catenescent conidia. Conidia ellipsoid, doliiform to
388subcylindrical, 30–39 × 15–20 μm(Fig.7).
389Golovinomyces orontii ex Vigna umbellata
390Conidiophores erect, arising from the upper surface of hyphal
391mother cells, 115–235 μm long. Foot cells straight or curved,
39232–57 × 13–15 μm, followed by 1–3 shorter cells, forming
393catenescent conidia. Conidia doliiform to subcylindrical,
39438–50 × 20–23 μm(Fig.8).
395Materials examined –on Coccinia grandis (L.) Voigt
396(Cucurbitaceae), THAILAND, Chiang Rai, 5 January 2016,
Fig. 8 Golovinomyces orontii on
Vigna umbellata (MUMH6893).
aConidia. bConidiophores.
cAppressoria. Bar = 20 μm
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397 J. Meeboon, MUMH6613; Lactuca indica L. (Asteraceae),
398 THAILAND, Chiang Rai, Mae Suai, 29 December 2016, J.
399 Meeboon & S. Takamatsu, MUMH6936; Torenia fournieri
400 Linden ex Fourn. (Linderniaceae), Chiang Rai, Wiangpapao,
401 5 January 2016, J. Meeboon, MUMH6881, Vigna umbellata
402 (Thunb.) Ohwi & H.Ohashi (Fabaceae), Chiang Rai, Mae
403 Suai, 29 December 2016, J. Meeboon, MUMH6893.
404 Notes–Golovinomyces cucurbitacearum (R.Y. Zheng &
405 G.Q. Chen) Vakal. & Kliron. is listed as powdery mildew on
406 Coccinia spp. (Braun and Cook 2012). The asexual morph of
407 the present sample is in good agreement with G. orontii,and
408 theidentityofthiscollectionisconfirmedbymeansof
409 molecular sequence analyses (see Fig. 1). The fungus on
410 Coccinia differs from G. cucurbitacearum in having longer
411 foot cells and broader conidia. Golovinomyces cichoracearum,
412 G. orontii,Leveillula lactucarum,L. lactucae-serriolae,and
413 Podosphaera xanthii have been recorded on Lactuca spp.
414 worldwide (Matsuda and Takamatsu 2003; Braun and Cook
Fig. 9 Golovinomyces spadiceus on Dahlia pinnata (MUMH3708). a
Conidiophores. bConidia. cGerminating conidia. Bar = 15 μm
Fig. 10 Golovinomyces spadiceus on Laggera crispata (MUMH1748).
aConidiophores. bGerm tubes. cGerminating conidia. Bar = 15 μm
Fig. 11 Golovinomyces spadiceus on Verbena hybrida (MUMH6684). a
Conidia. bConidiophores. cAppressoria. Bar = 20 μm
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415 2012; Takamatsu et al. 2013;Parketal.2015; Cho et al. 2016).
416 The asexual morph of the powdery mildew found in Thailand
417 on L. indica having conidiophores arising laterally and from the
418 upper surface of hyphae. However, some previous reports men-
419 tioned that the conidiophores of G. orontii obtained from
420 Lactuca spp. are only arising laterally, thus producing curved
421 foot cells (Braun and Cook 2012;Parketal.2015;Choetal.
422 2016). In addition, the identification as G. orontii has been
423 confirmed by means of molecular sequence analyses (see Fig.
424 1). Vági et al. (2007)reportedGolovinomyces sp. on Tor e n i a
425 fournieri from Hungary. The morphology of the specimen on
426 Torenia fournieri from Thailand is similar to the fungus col-
427 lected in Hungary, and a sequence retrieved from this collection
428 clusters within G. orontii group 3 according to Takamatsu et al.
429 (2013). Vigna umbellata was previously unknown as host of
430 Golovinomyces spp. The sequences of the powdery mildews
431 from C. grandis and V. umbellata belong to the big G. orontii
432 complex and form a clade of its own with 94% BS support
433 (Fig. 1). These hosts are new to Thailand.
434 Golovinomyces spadiceus (Berk. & M.A. Curtis) U. Braun
435 ex Dahlia pinnata and Dahlia × hortensis
436 Mycelium amphigenous, forming white patches, confluent,
437 sometimes covering entire leaves, persistent or almost so.
438Hyphae 4–8μm wide, thin-walled, smooth, hyaline. Hyphal
439appressoria solitary, nipple-shaped, 3–8μmdiam.
440Conidiophores erect, arising from upper surface of hyphal
441mother cell and usually towards one end of it, 125–188 μm
442long. Foot cells cylindrical, 52–98 × 9–13 μm, followed by 1–
4433 shorter cells, forming catenescent conidia. Conidia ellipsoid-
444ovoid, 27–33 × 15–25 μm. Conidial germination of the
445Euoidium type (Fig. 9).
446Materials examined –On Dahlia pinnata Cav.
447(Asteraceae), Chiang Mai, Su Thep, 21 January 2005, J.
448Meeboon, MUMH3708; Dahlia ×hortensis Guillaumin,
449Chiang Rai, Mae Suai, 27 December 2016, J. Meeboon,
450MUMH6874.
451Notes –Braun and Cook (2012) described the morpholog-
452ical characteristics of G. spadiceus, which overlapped with
453our own specimens. The molecular phylogenetic analysis
454based on the ITS rDNA sequence of this specimen showed
455that the present fungus nested in the G. ambrosiae clade (III
456sensu Takamatsu et al. 2013) with 98% BS (Fig. 1). This is the
457first report of G. spadiceus on Dahlia pinnata and Dahlia ×
458hortensis from Thailand.
459Golovinomyces spadiceus (Berk. & M.A. Curtis) U. Braun
460ex Laggera crispata
461Mycelium amphigenous, effuse or in thin, irregular
462patches, white, persistent. Hyphae sparingly branched,
Fig. 12 Golovinomyces
sonchicola on Sonchus oleraceus
(MUMH1772). aConidiophores.
bConidia. cGerminating conidia.
dAppressoria. Bar = 18 μm
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463 straight to moderately sinuous, 4–8μm wide, hyaline, thin-
464 walled, smooth. Hyphal appressoria almost indistinct to nip-
465 ple-shaped, usually solitary, 2–7μm diam. Conidiophores
466 156–187 μm long, arising laterally or from the upper surface
467 of hyphal mother cells, and positioned almost centrally or
468 towards one end of the cells, slightly curved at the base.
469 Foot cells subcylindrical, 61–112 × 10–13.5 μm, followed
470 by 0–3 shorter cells, forming catenescent conidia. Conidia
471 ellipsoid-obovoid, often constricted at the ends, 35–41 × 16–
472 21 μm. Conidial germination of the Euoidium type (Fig. 10).
473 Material examined –On Laggera crispata (Vahl) Hepper
474 & J.R.I. Wood (= L. pterodonta (DC.) Sch.Bip. ex Oliv.)
475 (Asteraceae, Inuleae), Chiang Mai, Mae Rim, 18 January
476 2005, J. Meeboon, MUMH1748.
477 Notes –This is the first report of powdery mildew on
478 L. crispata. The morphological characteristics were typical
479 of the asexual morph of the genus Golovinomyces,Footcells
480 of current material are longer than previous data, 30–80 × 9–
481 15 μm (Braun and Cook 2012), and conidia are longer, 25–40
482 ×14–20 μm (Braun and Cook 2012). Based on these morpho-
483 logical characteristics, we identify the powdery mildew on
484 L. crispata as G. spadiceus.
485 Golovinomyces spadiceus (Berk. & M.A. Curtis) U. Braun
486 ex Verbena × hybrida
487 Mycelium amphigenous, effuse or in thin, irregular patches,
488 white, persistent. Hyphae sparingly branched, straight to mod-
489 erately sinuous, 4–8μm wide, hyaline, thin-walled, smooth.
490 Hyphal appressoria almost indistinct to nipple-shaped, usually
491 solitary, 2–7μm diam. Conidiophores 81–182 μm long, arising
492 from the upper surface of hyphal mother cells, and positioned
493 almost centrally or towards one end of the cells, straight to
494 slightly curved at the base. Foot cells subcylindrical, 32–66 ×
495 10–15 μm, followed by 1–3 shorter cells, forming catenescent
496 conidia. Conidia doliiform-limoniform, 38–46 × 20–26 μm.
497 Conidial germination of the Euoidium type (Fig. 11).
498 Material examined –On Ve r b e n a ×hybrida Groenland &
499 Rümpler (Verbenaceae), Chiang Mai, Mae Rim, 19 January
500 2016, J. Meeboon, MUMH6684.
501 Notes –Golovinomyces verbenae (Schwein.) Heluta and
502 G. orontii have been recorded on Ver b e na spp. (Amano 1986;
503 Braun and Cook 2012). The current specimen was confirmed to
504 be G. spadiceus by the size of foot cells and conidial shape. This
505 is the first report of G. spadiceus on V. ×hybrida from Thailand.
506 Golovinomyces sonchicola U. Braun & R.T.A. Cook, in
507 Cook & Braun, Mycol. Res. 113(5): 629, 2009
508 Mycelium amphigenous, mainly epiphyllous, effuse or
509 forming white patches, thin. Hyphae straight to sinuous, hyaline,
510 thin-walled, smooth or almost so, 4–7μm wide. Hyphal appres-
511 soria nipple-shaped. Conidiophores arising from the hyphal
512 mother cell and towards one end of the cell, often close to a
513 septum, rarely in the middle, 112–160 μm long. Foot cells
514curved, 38–70 × 12–16 μm, slightly constricted at the basal
515septum, followed by 1–3 shorter cells, forming catenescent co-
516nidia. Conidia ellipsoid-obovoid, 34–48 × 18–22 μm, germ tubes
517terminal or almost so, short to moderately long, often with a
518slightly swollen appressorium at the tip, Euoidium type (Fig. 12).
519Material examined –on Sonchus oleraceus L.
520(Asteraceae), THAILAND, Chiang Mai, Su Thep, 5 January
5212005, J. Meeboon, MUMH1772.
522Notes –Shin (2000) described the asexual morphs of pow-
523dery mildews on Sonchus asper,S. brachyotus,and
524S. oleraceus collected in Korea. The ITS sequence of the
525powdery mildew on S. oleraceus was compared with the nu-
526cleotide sequences obtained from DNA databases. This fun-
527gus has the highest sequence similarity with G. sonchicola on
528S. oleaceus collected in Japan (99.8%). The present fungus
529formed a distinct clade with G. sonchicola on S. oleraceus
530(AB077623) collected in Japan with strong bootstrap support
531(98%) (Fig. 1). Based on the morphological and molecular
532characteristics, the powdery mildew on S. oleaceus is
Fig. 13 Golovinomyces sordidus on Plantago major (MUMH6633).
aConidiophore. bConidia. cGerminating conidia. dAppressoria.
Bar = 20 μm
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533 identified as G. sonchicola. This is the first report of
534 G. sonchicola on S. oleraceus from Thailand.
535 Golovinomyces sordidus (L. Junell) Heluta, Ukrayins’k.
536 Bot. Zhurn. 45(5): 63, 1988
537 Mycelium amphigenous, in irregular patches, almost per-
538 sistent or evanescent. Hyphae hyaline, walls thin, smooth or
539 almost so, 5–8μm wide. Hyphal appressoria nipple-shaped or
540 occasionally slightly lobed, sometimes poorly developed.
541 Conidiophores arising more or less laterally from the hyphal
542 mother cell and towards one end of the cell, often close to a
543 septum, 98–240 μm long. Foot cells almost cylindrical,
544 40–100 × 11–14 μm, followed by 3–4 shorter cells,
545 forming catenescent conidia. Conidia ellipsoid-ovoid to
546 subcylindrical, 30–40 × 20–24 μm, germ tubes terminal
547 or almost so, short to moderately long, ending in an
548 unlobed, somewhat swollen appressorium, Euoidium
549 type (Fig. 13).
550Material examined –on Plantago major L.
551(Plantaginaceae), THAILAND, Chiang Mai, Inthanon
552National Park, 11 December 2014, MUMH6633.
553Notes –The asexual morph of G. sordidus is the only
554Euoidium species occurring on Plantago spp. (Braun and
555Cook 2012). Although there are only minor differences in
556the sizes of conidiophores and foot cells, this fungus was
557identified as G. sordidus. This is the first report of
558G. sordidus on P. major from Thailand.
559Golovinomyces sp.ex Ageratum conyzoides
560Mycelium amphigenous, mainly epiphyllous, effuse or
561forming white patches, thin. Hyphae straight to sinuous, hya-
562line, thin-walled, smooth or almost so, 3–7μm wide. Hyphal
563appressoria nipple-shaped. Conidiophores 110–237 μmlong,
564erect, arising from upper surface of hyphal mother cell and
Fig. 14 Golovinomyces sp. on
Ageratum conyzoides
(MUMH6739). aConidia. b
Conidiophores. cAppressoria.
Bar = 20 μm
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565 usually towards one end of it. Foot cells cylindrical, 42–95 ×
566 12–16 μm, followed by 1–3 shorter cells, forming catenescent
567 conidia. Conidia doliiform to subcylindrical 38–53 × 17–20
568 μm. Conidial germination of the Euoidium type (Fig. 14).
569 Material examined –On Ageratum conyzoides L.
570 (Asteraceae, Eupatorieae), THAILAND, Lampang, Wang
571 Nue, 13 January 2016, J. Meeboon, MUMH6739.
572 Notes –Ageratum conyzoides is known as host of
573 G. cichoracearum s. lat. (Amano 1986). The foot cells
574 and conidia of this specimen are longer than in
575 G. cichoracearum s. str. (30–)40–80 μmlongvs.42–
576 95 μm long and 25–42 × 14–23 μmvs.38–53 × 17–20
577 μm, respectively (Braun and Cook 2012). The sequence
578 of the powdery mildew on A. conyzoides clustered with
579 those on Bidens pilosa in the clade of G. ambrosiae (III
580 sensu Takamatsu et al. 2013)with92%BSandthe
581 morphological characteristics are clearly different from
582 G. ambrosiae, foot cells are shorter, 35–80 × 9–15 μm
583 and conidia are broader, 25–45 × 15–27 μm(BraunandCook
584 2012). Identification of Golovinomyces on A. conyzoides
585 using solely based on the ITS rDNA sequences is insufficient.
586 This is the first report of Golovinomyces sp. on A. conyzoides
587 from Thailand.
588 Golovinomyces sp. ex Bidens pilosa
589 Mycelium amphigenous, mainly epiphyllous, effuse or
590 forming white patches, thin. Hyphae straight to sinuous, hya-
591 line, thin-walled, smooth or almost so, 4–6μm wide. Hyphal
592 appressoria nipple-shaped. Conidiophores arising from upper
593surface of hyphal mother cell and usually towards one end of
594it, 95–190 μm long. Foot cells 41–68 × 10–16 μm, slightly
Fig. 15 Golovinomyces sp. on
Bidens pilosa (MUMH6685). a
Conidia. bConidiophores. c
Appressoria. Bar = 20 μm
Fig. 16 Golovinomyces sp. on Lygisma inflexum (MUMH1784). a
Conidiophores. bConidia with germ tubes. Bar = 20 um
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595 constricted at the basal septum, followed by 1–3 shorter cells,
596 forming catenescent conidia. Conidia doliiform 34–50 × 20–
597 25 μm. Conidial germination of the Euoidium type (Fig. 15).
598 Material examined –On Bidens pilosa L. Chiang Rai,
599 Wiangpapao, 3 January 2016, J. Meeboon, MUMH6685
600 Notes –Bidens pilosa is known as host of
601 G. cichoracearum s. lat. (Amano 1986). This is the first report
602 of Golovinomyces sp. on B. pilosa from Thailand.
603 Golovinomyces sp. ex Lygisma inflexum
604 Mycelium amphigenous, forming patches or effuse, often con-
605 fluent, persistent, particularly on the upper leaf surface, often
606 evanescent on the lower surface, white or dingy greyish white.
607 Hyphae straight to sinuous-geniculate, walls thin, smooth or
608 almost so, hyaline, 3–8μm wide. Hyphal appressoria almost
609 indistinct to nipple-shape. Conidiophores arising from upper
610 surface and usually towards one end of hyphal mother cells,
611 186–262 μm long. Foot cells straight to curved, cylindrical,
612 76–82 × 12–16 μm, followed by 1–3 shorter cells, forming
613 catenescent conidia, often in long chains. Conidia ellipsoid-
614 cylindrical, 35–43.5 × 20–21 μm, germ tubes short to moder-
615 ately long, Euoidium type (Fig. 16).
616 Material examined –on Lygisma inflexum (Costantin)
617 Kerr (Asclepiadaceae), THAILAND, Chiang Mai, Su Thep,
618 5 January 2005, J. Meeboon, MUMH1784.
619 Notes –Golovinomyces cichoracearum s. lat. has been
620 recorded on the asclepidaceous hosts Asclepias syriaca L.
621 and A. tuberosa L. and G.orontii on A. incarnata L. and
622 Hoya carnosa (L.) R.Br. (Farr and Rossman 2017), but
623 Lygisma inflexum has not yet been listed as host of any
624 Golovinomyces species (Braun and Cook 2012). This is the
625 first record of powdery mildews on L. inflexum. Although
626 DNA isolation of this specimen failed, based on conidial
627 chains with sinuous edge lines and conidia without fibrosin
628 bodies, this powdery mildew is assigned to the genus
629 Golovinomyces.
630 Acknowledgements This work was financially supported in part by a
631 Grant-in-Aid for Scientific Research (No. 16K07613 and 16F16097)
632 from the Japan Society for the Promotion of Science to ST; and The
633 JSPS postdoctoral fellowship to JM (P16097).
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JrnlID 40858_ArtID 201_Proof# 1 - 28/10/2017
AUTHOR'S PROOF
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AUTHOR QUERIES
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Q1. Please check captured Keywords if correct.
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Q3. Ref. "Walsh et al. 1991" is cited in the body but its bibliographic information is missing. Kindly
provide its bibliographic information in the list.
Q4. Ref. "Hirata and Takamatsu (1996)" is cited in the body but its bibliographic information is
missing. Kindly provide its bibliographic information in the list.
Q5. Ref. "Takamatsu and Kano 2001" is cited in the body but its bibliographic information is missing.
Kindly provide its bibliographic information in the list.
Q6. Ref. "White et al. 1990" is cited in the body but its bibliographic information is missing. Kindly
provide its bibliographic information in the list.
Q7. Ref. "Felsenstein 1985" is cited in the body but its bibliographic information is missing. Kindly
provide its bibliographic information in the list.
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Q9. The citation “Narayanaswamy and Ramakrishnan (1969)”has been changed to “Narayanaswami
and Ramakrishnan (1967)”to match the author name/date in the reference list. Please check if the
change is fine in this occurrence and modify the subsequent occurrences, if necessary.
Q10. Ref. "Braun (1987)" is cited in the body but its bibliographic information is missing. Kindly
provide its bibliographic information in the list.
Q11. References [Hirata 1976, Paul and Thakur 2006, Takamatsu et al. 2010] were provided in the
reference list; however, this was not mentioned or cited in the manuscript. As a rule, all references
given in the list of references should be cited in the main body. Please provide its citation in the
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