Morphology and Phylogeny of Pestalotiopsis (Sporocadaceae, Amphisphaeriales) from Fagaceae Leaves in China

Ning Jiang; Hermann Voglmayr; Han Xue; Chun-Gen Piao; Yong Li

doi:10.1128/spectrum.03272-22

. 2022 Nov 10;10(6):e03272-22. doi: 10.1128/spectrum.03272-22

Morphology and Phylogeny of Pestalotiopsis (Sporocadaceae, Amphisphaeriales) from Fagaceae Leaves in China

Ning Jiang ^a, Hermann Voglmayr ^b, Han Xue ^a, Chun-Gen Piao ^a, Yong Li ^a,^✉

Editor: Florian M Freimoser^c

PMCID: PMC9769744 PMID: 36354327

ABSTRACT

Fagaceae is a family of flowering plants widely distributed in the Northern Hemisphere, including deciduous and evergreen trees and shrubs. Species of Pestalotiopsis are well-known agents of leaf spot diseases, but targeted sampling on Fagaceae is still missing. To determine the diversity of Pestalotiopsis species associated with Fagaceae leaf spot in China, investigations were conducted in the main areas of Fagaceae distribution from 2016 to 2021. Diseased leaf tissues were collected, and fungal isolates were obtained from leaf spots. In the present study, 43 isolates of Pestalotiopsis were studied based on combined morphology and phylogeny. As a result, 10 new species were identified, viz., Pestalotiopsis anhuiensis, P. castanopsidis, P. changjiangensis, P. cyclobalanopsidis, P. foliicola, P. guangxiensis, P. guizhouensis, P. lithocarpi, P. shaanxiensis, and P. silvicola, and six new host records were recognized.

IMPORTANCE Pestalotiopsis is a common fungal genus inhabiting plant tissues as endophytes, pathogens, and saprophytes. Fagaceae is a plant family including many important tree species, such as Castanea mollissima and Quercus spp. In this study, diseased leaves of Fagaceae in China were investigated, and 16 Pestalotiopsis species were identified based on morphology and phylogeny of combined loci of internal transcribed spacers (ITS), the translation elongation factor 1-α (tef1), and the beta-tubulin (tub2) genes. Among these, 10 new species were found, and six new host records were revealed. Our study significantly updates the taxonomy of Pestalotiopsis and enhances our understanding of leaf diseases of Fagaceae hosts.

KEYWORDS: new species, phylogeny, plant disease, taxonomy

INTRODUCTION

Fagaceae is an ecologically important family of flowering plants including eight genera and about 927 species worldwide (1). Most species of Fagaceae are deciduous trees or shrubs in temperate regions, and some species are evergreens distributed in subtropical to tropical regions. As we know, chestnuts (Castanea spp.) provide delicious natural foods, and oaks (Quercus spp.) and beeches (Fagus spp.) are commonly used as timbers. Many fagaceous species in temperate forests are ecologically highly important as main components of various forest types and provide an important food source for wildlife, and some of them are prominent ornamental trees (2).

More than 320 Fagaceae species have been recorded from China, including Chinese chestnut (Castanea mollissima), which provides tree crops, and oriental cork oak (Quercus variabilis), which is widely used as a shade, street, or ornamental tree in China. Leaf spot diseases are common on Fagaceae hosts in China, from which several fungal species were revealed based on morphology and phylogeny in recent years (3). For example, Diaporthe eres and Ophiognomonia castaneae cause Castanea mollissima brown margin leaf blight in Shandong Province of China (4). Botryosphaeria qinlingensis causes oak frogeye leaf spot disease in China (5). Monochaetia castaneae and 25 other species were reported to be associated with C. mollissima leaf diseases in chestnut plantations in China (6).

Pestalotioid fungi are easily characterized by multiseptate and more or less fusiform conidia with appendages at one or both ends, frequently with some melanized cells (7 –9). Fungi currently known as pestalotioid fungi are classified in the family Sporocadaceae, with 30 accepted genera based on multiple-locus phylogeny and conidial characters (8).

The classification of pestalotioid genera is complicated and has undergone substantial rearrangements in the past decades. The genus Pestalotia, originally described by De Notaris (10), contains almost 600 species epithets, the majority of which were over time transferred to separate genera. More than 100 years after its description, in his pioneering revision, Steyaert (11) restricted Pestalotia to the generic type species, Pestalotia pezizoides, and separated Pestalotiopsis and Truncatella based on cell numbers in the conidium: 4 cells in Truncatella, 5 cells in Pestalotiopsis, and 6 cells in Pestalotia. However, Steyaert (11) did not accept Monochaetia as a distinct genus but placed its species into Pestalotiopsis or Truncatella based on cell numbers of the conidium. In contrast, in his revision of Pestalotia, Guba (12) did not accept Pestalotiopsis and Truncatella as distinct genera but re-established Monochaetia for species having a single apical and basal appendage, which he placed in three sections based on the number of conidial cells (4, 5, or 6). Subsequently, a hybrid classification scheme was established recognizing all four genera (Pestalotia, Pestalotiopsis, Truncatella, and Monochaetia), which was primarily based on the cell numbers per conidium in combination with the number of apical appendages (one versus several). Monochaetia was restricted to species with five-celled conidia, while species with four-celled conidia were transferred to Truncatella or Seimatosporium and species with six-celled conidia to Seiridium (13).

Molecular phylogenetic analysis largely confirmed the morphological generic classification of Sutton (13), and it was subsequently further refined. Phylogenetically, Monochaetia sensu stricto, Pestalotiopsis, Seiridium, and Truncatella were shown to form distinct clades in Sporocadaceae (8, 14), while Pestalotia was shown to be synonymous with the older genus Seiridium (15). Based on phylogeny of multiple genes and conidial morphology, Pestalotiopsis sensu lato was further split into Pestalotiopsis sensu stricto, Neopestalotiopsis, and Pseudopestalotiopsis (16). While Neopestalotiopsis can be distinguished from the other genera by a versicolorous medium part of the conidia, viz., a lighter brown second cell and darker brown third and fourth cells, conidia of Pestalotiopsis and Pseudopestalotiopsis are morphologically indistinguishable but differ molecularly, e.g., by the lengths of their internal transcribed spacer (ITS) sequences (489 to 495 bp in Pestalotiopsis versus 536 to 540 bp in Pseudopestalotiopsis) (16). Ecologically, species of Pestalotiopsis sensu lato are common leaf pathogens infecting various hosts worldwide (17 –20).

As Pestalotiopsis species are known as leaf spot pathogens on Fagaceae, typical spotted leaves were collected to obtain fungal isolates, which were subsequently identified based on both morphological and phylogenetic approaches. The aims of the present study were to reveal hidden species diversity of Pestalotiopsis from diseased fagaceous leaves and to evaluate the practicability of host association for species distinction.

RESULTS

The combined sequence data set of ITS, tef1, and tub2 comprised 1,552 characters (524 for the ITS, 529 for tef1, and 499 for tub2) from 162 isolates, including one outgroup taxon, Neopestalotiopsis magna (MFLUCC 12-0652). Of the 1,552 characters included in the phylogenetic analyses, 457 were parsimony informative (84 from the ITS, 200 from tef1, and 173 from tub2). The best maximum-likelihood (ML) tree (lnL = −12,981.92) revealed by RAxML is shown as a phylogram in Fig. 1. The topologies resulting from ML and Bayesian inference (BI) analyses of the concatenated data set were congruent. Isolates from the present study formed 16 individual clades representing 16 species of Pestalotiopsis, including 10 new species (Pestalotiopsis anhuiensis, P. castanopsidis, P. changjiangensis, P. cyclobalanopsidis, P. foliicola, P. guangxiensis, P. guizhouensis, P. lithocarpi, P. shaanxiensis, and P. silvicola) and six known species (P. chamaeropis, P. kenyana, P. lushanensis, P. nanjingensis, P. neolitseae, and P. rhodomyrtus).

FIG 1 — Phylogram of *Pestalotiopsis* resulting from a maximum-likelihood analysis based on a combined matrix of ITS, *tef1*, and *tub2* loci. Numbers above the branches indicate ML bootstrap values (left; values of ≥50% are shown) and Bayesian posterior probabilities (right; values of ≥0.9 are shown). The tree is rooted with *Neopestalotiopsis magna* (MFLUCC 12–0652). Isolates from the present study are marked in blue.

TAXONOMY

Pestalotiopsis anhuiensis Ning Jiang, sp. nov. (Fig. 2). MycoBank number MB843387. Etymology: named after the collection site of the type specimen, Anhui Province. Typus: China, Anhui Province, Hefei City, Shushan District, Dashushan Forest Park, on diseased leaves of Cyclobalanopsis glauca, 2 November 2019, Dan-ran Bian (holotype CAF 800044; ex-holotype culture CFCC 54791).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 50 to 300 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 3.5 to 13.5 by 2 to 3.5 μm, mean ± standard deviation (SD) = 8.2 ± 2.6 by 3.1 ± 1.1 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (18.5)20.5 to 23.5(25) by (6)6.5 to 7.5(8) μm (measurements of conidia are reported as maximum and minimum in parentheses and the range representing the mean ± standard deviation of the number of measurements given in parentheses), mean ± SD = 21.8 ± 1.4 by 7.1 ± 0.4 μm, length/width ratio (L/W) = 2.7 to 3.8; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3.5)4.5 to 5.5(6) μm; median cells 3, trapezoid or subcylindrical, concolorous, pale brown to brown, thick-walled, the first median cell from base (4)5 to 6 μm long, the second cell 4 to 5 μm long, the third cell (3.5)4 to 5(5.5) μm long, together (12)13 to 15.5(17) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 4(4.5) μm long; basal appendage single, unbranched, tubular, centric, straight or slightly bent, (4)4.5 to 6.5(7) μm long, mean ± SD = 5.4 ± 0.9 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or bent, (7.5)12.5 to 17.5(20) μm long, mean ± SD = 14.9 ± 2.6 μm. Sexual morph unknown.

Colonies on malt extract agar (MEA) flat, spreading, with flocculent aerial mycelium and entire edge, white, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on potato dextrose agar (PDA), flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Notes: Pestalotiopsis anhuiensis from Cyclobalanopsis glauca is phylogenetically close to Pestalotiopsis abietis, P. disseminata, and P. guangxiensis (Fig. 1). Morphologically, P. anhuiensis shares similar conidial sizes with P. abietis and P. disseminata (18.5 to 25 by 6 to 8 μm in P. anhuiensis versus 19.9 to 31.2 by 5.8 to 8 μm in P. abietis and 18 to 25 by 6.5 to 8 μm in P. disseminata) (21 –23) and has narrower conidia than P. guangxiensis (6 to 8 μm versus 7.5 to 9.5 μm in P. guangxiensis) (Table 1). However, P. anhuiensis can be distinguished by sequence data (nucleotide differences from P. abietis: in the ITS, 1/506 [0.2%]; in tef1, 4/470 [0.85%]; in tub2, 1/442 (0.23%); nucleotide differences from P. disseminata: in the ITS, 3/506 [0.59%], 1 insertion; in tef1, 8/470 [1.7%]; in tub2, 1 or 2/406 [0.25 to 0.5%]; nucleotide differences from P. guangxiensis: in the ITS, 1/506 [0.2%], 1 insertion; in tef1, 12/470 [2.55%], 1-bp gap; in tub2, 2 or 3/442 [0.45 to 0.68%]).

TABLE 1.

Synopsis of Pestalotiopsis occurring on fagaceous hosts^a

Species	Host(s)	Length of conidia (μm)	Width of conidia (μm)	Length of 3 median cells (μm)	Length of apical appendage (μm)	Length of basal appendage (μm)	Reference
P. anhuiensis	Cyclobalanopsis glauca	18.5–25	6–8	12–17	7.5–20	4–7	This study
P. castanopsidis	Castanopsis hystrix, C. lamontii	23–29	7–11.5	16–20.5	17–24.5	8.5–15	This study
P. chamaeropis	C. fissa, Quercus acutissima, Q. aliena, Q. variabilis	20.5–31.5	6.5–9	13–20	8.5–27.5	2.5–12	This study
P. changjiangensis	Castanopsis hainanensis, C. tonkinensis	19–24	7–8.5	13.5–16.5	1.5–7	absent	This study
P. cyclobalanopsidis	Cyclobalanopsis glauca	18.5–25.5	6–8.5	13–16	5.5–14.5	2–6.5	This study
P. foliicola	Castanopsis faberi	19.5–24	7–9.5	10.5–16	10.5–37	3–5	This study
P. guangxiensis	Quercus griffithii	17.5–21	7.5–9.5	12–14	14–19	3–4.5	This study
P. guizhouensis	Cyclobalanopsis glauca	21–26.5	7–9.5	13–18	7–15	2–8	This study
P. kenyana	Castanea henryi, Ca. mollissima, Castanopsis fissa, C. hystrix, Cyclobalanopsis glauca, Cy. fleuryi, Quercus aliena, Q. aliena var. acutiserrata	20.5–28	6–8	NA	3.5–15	1.5–3.5	6; this study
P. lithocarpi	Lithocarpus chiungchungensis	17–23	5.5–8	12.5–14.5	9–24	2.5–5	This study
P. lushanensis	Quercus serrata	19.5–26.5	7–9.5	13–17.5	10.5–26.5	2.5–0.5	This study
P. monochaeta	Quercus robur	25–42	7–11.5	17–26	40–75	6–14	16
P. nanjingensis	Quercus aliena	17.5–23.5	6.5–9	13–7.5	8.5–20.5	2–6.5	This study
P. neolitseae	Lithocarpus amygdalifolius	19–23.5	5.5–7	13–14.5	9.5–14.5	2–3.5	This study
P. rhodomyrtus	Cyclobalanopsis augustinii, Quercus aliena	20–27	6–8	13.5–18	8–17.5	2–7	This study
P. shaanxiensis	Quercus variabilis	21–25	7–9	13.5–17.5	13–22	1.5–7.5	This study
P. silvicola	Cyclobalanopsis kerrii	20–26	6–8.5	12–16	12–22.5	3.5–8	This study

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NA, not available.

Pestalotiopsis castanopsidis Ning Jiang, sp. nov. (Fig. 3). MycoBank number MB841308. Etymology: named after the host genus, Castanopsis. Typus: China, Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Castanopsis lamontii, 4 December 2019, Shang Sun (holotype CAF 800021; ex-holotype culture CFCC 54430).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 50 to 350 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 5 to 11.5 by 2.5 to 7 μm, mean ± SD = 7.2 ± 2.5 by 4.4 ± 1.4 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (23)24 to 27.5(29) by (7)8 to 11(11.5) μm, mean ± SD = 25.7 ± 1.8 by 9.3 ± 1.4 μm, L/W = 2.3 to 4; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, 4 to 5 μm; median cells, 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (5.5)6 to 7 μm long, the second cell (5.5)6 to 7.5(8) μm long, the third cell 5.5 to 6.5(7) μm long, together (16)17 to 20(20.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 4.5(5) μm long; basal appendage single, unbranched, tubular, centric, straight or slightly bent, (8.5)9.5 to 14(15) μm long, mean ± SD = 11.7 ± 2.4 μm; apical appendages, 3 or 4, unbranched, tubular, knobbed, centric, straight or slightly bent, (17)17.5 to 23(24.5) μm long, mean ± SD = 20.3 ± 2.7 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, off-white to sienna, reaching 60 mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, pale luteous to fawn, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional materials, China, Guangdong Province, Shaoguan City, Lechang City, Dayaoshan Forest Farm, on diseased leaves of Castanopsis hystrix, 4 December 2019, Dan-ran Bian (CFCC 54305 and CFCC 54384).

Notes: Three isolates of Pestalotiopsis castanopsidis from Castanopsis hystrix and C. lamontii clustered into a distinct clade phylogenetically close to P. jesteri (Fig. 1). However, P. castanopsidis differs from P. jesteri by obviously larger conidia (23 to 29 by 7 to 11.5 μm in P. castanopsidis versus 19 to 23 by 5 to 7 μm in P. jesteri) (24). In addition, P. castanopsidis can be distinguished from P. jesteri by sequence data (nucleotide differences: in the ITS, 4/364 [1.1%]; in tub2, 14 or 15/438 [3.2 to 3.42%]).

Pestalotiopsis chamaeropis S. S. Maharachch, K. D. Hyde & P. W. Crous, Studies in Mycology 79:158 (2014).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 150 to 450 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 4.5 to 12 by 2.5 to 4.5 μm, mean ± SD = 6.9 ± 1.5 by 3.6 ± 0.7 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (20.5)23 to 28.5(31.5) by (6.5)7.5 to 8.5(9) μm, mean ± SD = 25.8 ± 2.6 by 8 ± 0.7 μm, L/W = 2.5 to 4.4; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3.5)4.5 to 7(8) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (4)4.5 to 6(7) μm long, the second cell (4.5)5 to 6.5(7) μm long, the third cell (4)5 to 6(6.5) μm long, together (13)15 to 18(20) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (3)3.5 to 5(5.5) μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (2.5)4 to 8.5(12) μm long, mean ± SD = 6.3 ± 2.2 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (8.5)12 to 21.5(27.5) μm long, mean ± SD = 16.8 ± 4.8 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white to buff, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Materials examined, China, Anhui Province, Hefei City, Shushan District, Dashushan Forest Park, on diseased leaves of Quercus aliena, 2 November 2019, Dan-ran Bian (CFCC 55019 and CFCC 55122); Anhui Province, Hefei City, Shushan District, Dashushan Forest Park, on diseased leaves of Quercus acutissima, 2 November 2019, Dan-ran Bian (CFCC 54977); Anhui Province, Hefei City, Shushan District, Dashushan Forest Park, on diseased leaves of Quercus variabilis, 2 November 2019, Dan-ran Bian (CFCC 54776); Guangdong Province, Shaoguan City, Lechang City, Dayaoshan Forest Farm, on diseased leaves of Castanopsis fissa, 4 December 2019, Shang Sun (CFCC 55023); Shaanxi Province, Xian City, Zhouzhi County, Heihe Forest Park, on diseased leaves of Quercus variabilis, 6 September 2019, Yong Li (CFCC 55338); Sichuan Province, Yaan City, Shimian County, on diseased leaves of Quercus acutissima, 10 September 2020, Ning Jiang (CFCC 55124).

Notes: Seven new isolates of Pestalotiopsis chamaeropis were collected from four fagaceous hosts, forming a well-supported clade with the ex-type strain CBS 186.71 (Fig. 1). In addition, samples from the present study agree well with the ex-type strain in conidial dimension and morphological characters (21 to 28 by 6 to 9.5 μm in CBS 186.71) (16). Hence, Castanopsis fissa, Quercus acutissima, Q. aliena, and Q. variabilis become new hosts for Pestalotiopsis chamaeropis, which was originally described from Chamaerops humilis (16).

Pestalotiopsis changjiangensis Ning Jiang, sp. nov. (Fig. 4). MycoBank number MB841309. Etymology: named after the collection site of the type specimen, Changjiang Li Autonomous County. Typus: China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Castanopsis tonkinensis, 16 November 2018, Yong Li (holotype CAF 800024; ex-holotype culture CFCC 54314).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, dark brown, 300 to 850 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 3.5 to 6 by 2 to 5.5 μm, mean ± SD = 4.7 ± 1.1 by 3.4 ± 1 μm. Conidia fusoid, straight, 4-septate, smooth, not constricted or slightly constricted at the septa, (19)20 to 22.5(24) by 7 to 8(8.5) μm, mean ± SD = 21.2 ± 1.4 by 7.7 ± 0.4 μm, L/W = 2.4 to 3.3; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (2)3 to 4 μm; median cells 3, trapezoid or subcylindrical, pale brown to brown, thick-walled, the first median cell from base 4.5 to 5.5 μm long, the second cell (4.5)5 to 5.5 μm long, the third cell (4)4.5 to 6 μm long, together (13.5)14.5 to 16(16.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline or pale brown, (2)2.5 to 4(5) μm long; basal appendage indistinct or absent; apical appendage indistinct, tubular, bent, 1.5 to 4.5(7) μm long, mean ± SD = 3 ± 1.1 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium and undulate edge, pale luteous, reaching a 70-mm diameter after 10 days at 25°C, sterile; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white, reaching a 65-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional materials examined, China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Castanopsis hainanensis, 14 November 2018, Yong Li (CFCC 54433); Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis austrocochinchinensis, 16 November 2018, Yong Li (CFCC 52803).

Notes: Three isolates of Pestalotiopsis collected from Castanopsis hainanensis, C. tonkinensis, and Cyclobalanopsis austrocochinchinensis clustered into a distinct and well-supported clade, which is newly described as P. changjiangensis here (Fig. 1). Its conidia are characterized by the lack of a basal appendage and an indistinct apical appendage, which is unique within the genus Pestalotiopsis.

Pestalotiopsis cyclobalanopsidis Ning Jiang, sp. nov. (Fig. 5). MycoBank number MB841310. Etymology: named after the host genus, Cyclobalanopsis. Typus: China, Guangdong Province, Shaoguan City, Lechang City, Dayaoshan Forest Farm, on diseased leaves of Cyclobalanopsis glauca, 4 December 2019, Shang Sun (holotype CAF 800022; ex-holotype culture CFCC 54328).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, dark brown, 250 to 700 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 6 to 8 by 2 to 4.5 μm, mean ± SD = 7.2 ± 0.9 by 3.6 ± 0.8 μm. Conidia fusoid, curved, 4-septate, smooth, slightly constricted at the septa, (18.5)20 to 24.5(25.5) by (6)6.5 to 8(8.5) μm, mean ± SD = 22.3 ± 1.3 by 7.3 ± 0.5 μm, L/W = 2.6 to 3.6; basal cell obconic with a truncate base, thin-walled, hyaline to pale brown, 4 to 6(6.5) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base 4.5 to 6(6.5) μm long, the second cell (3.5)4 to 5 μm long, the third cell (4)4.5 to 5.5 μm long, together 13 to 15.5(16) μm long; apical cell conic with an acute apex, thin-walled, hyaline to pale brown, 3 to 4 μm long; basal appendage unbranched, tubular, centric, straight, (2)2.5 to 5.5(6.5) μm long, mean ± SD = 3.8 ± 1.5 μm; apical appendages, 2 to 5, unbranched, tubular, bent, (5.5)7 to 13(14.5) μm long, mean ± SD = 10.1 ± 3 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire, hazel to off-white, reaching a 70-mm diameter after 10 days at 25°C, sterile; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white to luteous, reaching a 50-mm diameter after 10 days at 25°C, forming dark brown conidiomata with black conidial masses.

Additional material examined, China, Guangdong Province, Shaoguan City, Lechang City, Dayaoshan Forest Farm, on diseased leaves of Cyclobalanopsis glauca, 4 December 2019, Shang Sun (CFCC 55891).

Notes: Pestalotiopsis cyclobalanopsidis is closely related to P. castanopsidis, P. guizhouensis, and P. jesteri (Fig. 1). However, P. cyclobalanopsidis differs from them by distinctly curved conidia (24).

Pestalotiopsis foliicola Ning Jiang, sp. nov. (Fig. 6). MycoBank number MB843388. Etymology: folium = “leaf” and -cola = “inhabiting”; in reference to substrate origin of the type strain, leaves. Typus: China, Jiangxi Province, Xinyu City, Fenyi County, Dagangshan Nature Reserve, on diseased leaves of Castanopsis faberi, 13 November 2019, Shang Sun (holotype CAF 800045; ex-holotype culture CFCC 54440).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 100 to 450 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, annelidic, 6.5 to 20 by 2.5 to 6.5 μm, mean ± SD = 12.7 ± 4.2 by 3.1 ± 1.2 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (19.5)20 to 23(24) by (7)7.5 to 9(9.5) μm, mean ± SD = 21.5 ± 1.5 by 8.5 ± 0.5 μm, L/W = 2.1 to 2.9; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3)3.5 to 5(6) μm; median cells 3, trapezoid or subcylindrical, concolorous, pale brown to brown, thick-walled, the first median cell from base (4)4.5 to 5.5(6) μm long, the second cell (4)4.5 to 5.5(6) μm long, the third cell (3)4.5 to 5.5(6) μm long, together (10.5)13.5 to 15.5(16) μm long; apical cell conic with an acute apex, thin-walled, hyaline or pale brown, (2.5)3 to 4(4.5) μm long; basal appendage single, unbranched, tubular, centric, straight or slightly bent, (3)3.5 to 4.5(5) μm long, mean ± SD = 3.7 ± 0.7 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or bent, (10.5)14 to 26(37) μm long, mean ± SD = 20 ± 5.8 μm. Sexual morph unknown.

Colonies on MEA flat, with flocculent aerial mycelium and crenate edge, white, reaching a 35-mm diameter after 10 days at 25°C, sterile; on PDA, flat, spreading, with flocculent aerial mycelium and irregular edge, white to isabelline, reaching a 40-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional materials examined, China, Jiangxi Province, Xinyu City, Fenyi County, Dagangshan Nature Reserve, on diseased leaves of Castanopsis faberi, 13 November 2019, Shang Sun (CFCC 57359 and CFCC 57360).

Notes: Three isolates of Pestalotiopsis foliicola from Castanopsis faberi clustered into a distinct clade phylogenetically close to P. pinicola and P. rosea (Fig. 1). However, P. foliicola differs from P. pinicola and P. rosea by wider conidia (7 to 9.5 μm in P. foliicola versus 5 to 7 μm in P. pinicola and 5.7 to 7 μm in P. rosea) (25, 26).

Pestalotiopsis guangxiensis Ning Jiang, sp. nov. (Fig. 7). MycoBank number MB841311. Etymology: named after the collection site of the type specimen, Guangxi Zhuang Autonomous Region. Typus: China, Guangxi Zhuang Autonomous Region, Nanning City, Qingxiushan District, Qingxiushan Park, on diseased leaves of Quercus griffithii, 4 December 2019, Dan-ran Bian (holotype CAF 800023; ex-holotype culture CFCC 54308).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 350 to 800 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 8.5 to 17.5 by 2 to 5 μm, mean ± SD = 11.4 ± 2.7 by 3.1 ± 1.3 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (17.5)18 to 20.5(21) by (7.5)8 to 9(9.5) μm, mean ± SD = 19.2 ± 1.2 by 8.3 ± 0.5 μm, L/W = 1.9 to 2.6; basal cell obconic with a truncate base, thin-walled, hyaline, 2.5 to 4.5 μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base 4 to 4.5 μm long, the second cell 4 to 5(5.5) μm long, the third cell 4 to 5 μm long, together (12)12.5 to 13.5(14) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2)2.5 to 3(3.5) μm long; basal appendage unbranched, tubular, centric, straight, (3)3.5 to 4(4.5) μm long, mean ± SD = 4 ± 0.5 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (14)15 to 18.5(19) μm long, mean ± SD = 16.8 ± 1.9 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming radially folded surface and undulate edge, isabelline, reaching a 55-mm diameter after 10 days at 25°C, producing yellow droplet, sterile; on PDA, flat, spreading, with flocculent aerial mycelium and crenate edge, white to fawn, reaching a 65-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional material examined, China, Guangxi Zhuang Autonomous Region, Nanning City, Qingxiushan District, Qingxiushan Park, on diseased leaves of Quercus griffithii, 4 December 2019, Dan-ran Bian (CFCC 54300).

Notes: Two isolates of P. guangxiensis from Quercus griffithii formed a well-supported clade phylogenetically close to P. disseminata (Fig. 1). P. guangxiensis can be distinguished from P. disseminata by wider conidia (7.5 to 9.5 μm in P. guangxiensis versus 6.5 to 8 μm in P. disseminata) (21, 23). Additionally, P. guangxiensis differs from P. disseminata by sequence data (nucleotide differences: in the ITS, 4/506 [0.8%], 1 insertion; in tef1, 7/471 [1.49%], 6 insertions; in tub2, 1 to 3/406 [0.25 to 0.74%]).

Pestalotiopsis guizhouensis Ning Jiang, sp. nov. (Fig. 8). MycoBank number MB843389. Etymology: named after the collection site of the type specimen, Guizhou Province. Typus: China, Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Cyclobalanopsis glauca, 23 November 2019, Dan-ran Bian (holotype CAF 800046; ex-holotype culture CFCC 54803).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 100 to 400 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, ampulliform to spherical, annelidic, 6 to 17.5 by 3 to 6 μm, mean ± SD = 9.6 ± 4.3 by 4.5 ± 0.9 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (21)22.5 to 25.5(26.5) by (7)7.5 to 8.5(9.5) μm, mean ± SD = 23.8 ± 1.6 by 8.1 ± 0.6 μm, L/W = 2.4 to 3.5; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (2.5)3.5 to 5.5(6.5) μm; median cells 3, trapezoid or subcylindrical, concolorous, pale brown to brown, thick-walled, the first median cell from base (4.5)5 to 7(7.5) μm long, the second cell (3.5)4.5 to 5.5(6) μm long, the third cell (4)4.5 to 5.5(6) μm long, together (13)14.5 to 17.5(18) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 4.5(5) μm long; basal appendage single, unbranched, tubular, centric, straight or slightly bent, (2)3.5 to 6.5(8) μm long, mean ± SD = 4.8 ± 1.5 μm; apical appendages, 3 or 4, unbranched, tubular, attenuate toward the apex, centric, straight or slightly bent, (7)8 to 12(15) μm long, mean ± SD = 10.2 ± 2 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium and entire edge, white, reaching a 35-mm diameter after 10 days at 25°C, sterile; on PDA, flat, spreading, with flocculent aerial mycelium and irregular edge, sienna, reaching a 55-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional material examined, China, Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Cyclobalanopsis glauca, 23 November 2019, Dan-ran Bian (CFCC 57364).

Notes: Two isolates of Pestalotiopsis guizhouensis from Cyclobalanopsis glauca clustered into a distinct clade phylogenetically close to P. cyclobalanopsidis (Fig. 1). However, P. guizhouensis differs from P. cyclobalanopsidis by conidial shape (straight or slightly curved conidia in P. guizhouensis versus distinctly curved conidia in P. cyclobalanopsidis).

Pestalotiopsis kenyana S. S. Maharachch, K. D. Hyde & P. W. Crous, Studies in Mycology 79:158 (2014).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 150 to 650 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 5.5 to 15.5 by 2 to 4.5 μm, mean ± SD = 8.9 ± 1.4 by 3.3 ± 0.9 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (21)24 to 28.5(30) by (6)6.5 to 8(8.5) μm, mean ± SD = 26.1 ± 2.3 by 7.3 ± 0.5 μm, L/W = 2.6 to 4.9; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (4)4.5 to 6.5(8.5) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (4.5)5 to 6(6.5) μm long, the second cell (4.5)5 to 6(8) μm long, the third cell (4.5)5 to 6(6.5) μm long, together (13.5)15 to 17.5(19) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (3)4 to 5.5(6.5) μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (2.5)4 to 6.5(8.5) μm long, mean ± SD = 5.2 ± 1.5 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (4)9 to 15(20) μm long, mean ± SD = 12.2 ± 3.1 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white to pale luteous, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium and entire edge, white to luteous, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Materials examined: China, Henan Province, Xinyang City, Shihe District, Jigong Mountain, on diseased leaves of Cyclobalanopsis fleuryi, 7 August 2019, Yong Li (CFCC 55330); Henan Province, Xinyang City, Shihe District, Jigong Mountain, on diseased leaves of Cyclobalanopsis neglecta, 7 August 2019, Yong Li (CFCC 54732); Henan Province, Xinyang City, Shihe District, Jigong Mountain, on diseased leaves of Castanopsis fissa, 7 August 2019, Yong Li (CFCC 55088); Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Castanopsis hystrix, 4 December 2019, Shang Sun (CFCC 54742); Guangdong Province, Qingyuan City, Yangshan County, Guangdong Nanling Nature Reserve, on diseased leaves of Cyclobalanopsis glauca, 4 December 2019, Shang Sun (CFCC 54805); Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Quercus aliena, 23 November 2019, Dan-ran Bian (CFCC 54962); Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Quercus aliena var. acutiserrata, 23 November 2019, Dan-ran Bian (CFCC 54621 and CFCC 54618).

Notes: Eight new isolates of Pestalotiopsis kenyana were collected from six species and a variety of Fagaceae, which agree well with the ex-type strain CBS 442.67 in conidial dimension and characters (22 to 29 by 7 to 9 μm) (16). Hence, Castanopsis fissa, C. hystrix, Cyclobalanopsis fleuryi, Cy. glauca, Cy. neglecta, Quercus aliena, and Q. aliena var. acutiserrata become new hosts for Pestalotiopsis kenyana, which was originally described from Coffea species (16).

Pestalotiopsis lithocarpi Ning Jiang, sp. nov. (Fig. 9). MycoBank number MB841312. Etymology: named after the host genus, Lithocarpus. Typus: China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus chiungchungensis, 30 March 2019, Yong Li (holotype CAF 800025; ex-holotype culture CFCC 55100).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 450 to 1,100 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 4 to 10 by 3 to 5.5 μm, mean ± SD = 5.9 ± 2.8 by 4.2 ± 0.8 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (17)18.5 to 21.5(23) by (5.5)6 to 7(8) μm, mean ± SD = 20.2 ± 1.4 by 6.6 ± 0.6 μm, L/W = 2.5 to 3.7; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3.5)4 to 5 μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (3.5)4 to 5 μm long, the second cell 4 to 4.5(5) μm long, the third cell 4 to 5 μm long, together 12.5 to 14(14.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 3.5 μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, 2.5 to 4.5(5) μm long, mean ± SD = 3.6 ± 0.8 μm; apical appendages, 2 to 4 (mostly 3), unbranched, tubular, centric, straight or slightly bent, (9)12.5 to 21(24) μm long, mean ± SD = 16.7 ± 4.1 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, umber, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium and feathery edge, white to pale luteous, reaching a 65-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional material examined, China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus chiungchungensis, 30 March 2019, Yong Li (CFCC 55893).

Notes: Two isolates from leaf spots of Lithocarpus chiungchungensis clustered into a well-supported clade newly described here as Pestalotiopsis lithocarpi (Fig. 1). Phylogenetically, P. lithocarpi is close to P. dracontomelonis from diseased leaves of Dracontomelon dao (Anacardiaceae) collected in Thailand. Pestalotiopsis lithocarpi is similar to P. dracontomelonis in conidial size (17 to 23 by 5.5 to 8 μm in Pestalotiopsis lithocarpi versus 18 to 23 by 5.5 to 7.5 μm in P. dracontomelonis) (27). However, they can be distinguished by the length of the three median cells (12.5 to 14.5 μm in P. lithocarpi versus 13 to 17 μm in P. dracontomelonis) (27). In addition, P. lithocarpi can be distinguished from P. dracontomelonis by sequence data (nucleotide differences: in the ITS, 2/505 [0.4%], 1 insertion; in tef1, 5/457 [1.2%], 7 insertions).

Pestalotiopsis lushanensis F. Liu & L. Cai, Scientific Reports 7(no. 866):9 (2017).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 150 to 750 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 5.5 to 26.5 by 2.5 to 4.5 μm, mean ± SD = 13.6 ± 3.1 by 3.6 ± 0.9 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (19.5)21 to 25(26.5) by (7)7.5 to 9(9.5) μm, mean ± SD = 23 ± 1.9 by 8.4 ± 0.6 μm, L/W = 2.3 to 3.5; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3.5)4 to 5.5(6.5) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (4)4.5 to 5.5(6) μm long, the second cell (4)4.5 to 5.5(6) μm long, the third cell (4.5)5 to 6(6.5) μm long, together (13)14 to 16.5(17.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3.5 to 4.5(5.5) μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (5.5)6 to 9(11) μm long, mean ± SD = 7.5 ± 1.4 μm; apical appendages, 3, unbranched, tubular, centric, straight or slightly bent, (10.5)15 to 22.5(26.5) μm long, mean ± SD = 18.6 ± 3.7 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings, radially folded surface and entire edge, white to isabelline, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white to pale luteous, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Materials examined: China, Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Quercus serrata, 23 November 2019, Dan-ran Bian (CFCC 54894).

Notes: A new isolate of Pestalotiopsis lushanensis was collected from Quercus serrata, which agrees well with the ex-type strain LC4344 in conidial dimension and characters (20 to 27 by 7.5 to 10 μm in LC4344) (18). Hence, Quercus serrata becomes a new host for Pestalotiopsis lushanensis, which was originally described from Camellia sp. (18).

Pestalotiopsis nanjingensis Qin Yang & He Li, Journal of Fungi 7(12, no. 1080):21 (2021).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 50 to 500 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 8.5 to 19 by 2 to 4.5 μm, mean ± SD = 13.4 ± 1.7 by 2.9 ± 0.9 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (17.5)20 to 22(23.5) by (6.5)7 to 8.5(9) μm, mean ± SD = 20.8 ± 1 by 7.6 ± 0.7 μm, L/W = 2.2 to 3.5; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (2.5)3 to 4.5(5) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base 4 to 5(5.5) μm long, the second cell (4.5)5 to 5.5(6) μm long, the third cell (4)4.5 to 5.5(7) μm long, together (13)13.5 to 15.5(17.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2)3 to 4(4.5) μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (2)3.5 to 5(6.5) μm long, mean ± SD = 4.3 ± 0.9 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (8.5)11.5 to 16.5(20.5) μm long, mean ± SD = 14 ± 2.7 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium and undulate edge, white to pale luteous, reaching a 60-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white to pale gray, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Material examined, China, Henan Province, Xinyang City, Shihe District, Jigong Mountain, on diseased leaves of Quercus aliena, 7 August 2019, Yong Li (CFCC 53882).

Notes: A new isolate of Pestalotiopsis nanjingensis was collected from Quercus aliena, which has ITS, tef1, and tub2 sequences identical to those of the ex-type strain CSUFTCC16 (28). Hence, Quercus aliena becomes a new host for Pestalotiopsis nanjingensis, which was originally described from Camellia oleifera (28).

Pestalotiopsis neolitseae H. A. Ariyaw & K. D. Hyde, Mycosphere 9(5):1005 (2018).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 250 to 800 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 6 to 10 by 2.5 to 4 μm, mean ± SD = 7.7 ± 1.3 by 3.4 ± 0.6 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (19)20 to 22.5(23.5) by 5.5 to 6.5(7) μm, mean ± SD = 21.2 ± 1.3 by 6.1 ± 0.4 μm, L/W = 2.8 to 4.1; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, 3.5 to 5(6) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base 4.5 to 5 μm long, the second cell 4 to 5(5.5) μm long, the third cell 4.5 to 5 μm long, together 13 to 14.5 μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 4.5 μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (2)2.5 to 3.5 μm long, mean ± SD = 2.9 ± 0.6 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (9.5)11 to 14(14.5) μm long, mean ± SD = 12.6 ± 1.6 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white to pale gray, reaching a 30-mm diameter after 10 days at 25°C, sterile; on PDA, flat, spreading, with flocculent aerial mycelium and entire edge, white, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Material examined, China, Hunan Province, Changsha City, Changsha Forest Botanical Garden, on diseased leaves of Lithocarpus amygdalifolius, 9 November 2020, Cheng-ming Tian and Ning Jiang (CFCC 54590).

Notes: A new isolate of Pestalotiopsis neolitseae was collected from Lithocarpus amygdalifolius, which agrees well with the type specimen in conidial dimension and characters (29). Hence, Lithocarpus becomes a new host genus for Pestalotiopsis neolitseae, which was originally described from Neolitsea villosa (Lauraceae).

Pestalotiopsis rhodomyrtus Yu Song, K. Geng, K. D. Hyde & Yong Wang bis, Phytotaxa 126(1): 27 (2013).

Conidiomata in culture sporodochial, solitary, erumpent, pulvinate, black, 100 to 450 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to subcylindrical, annelidic, 2.5 to 6 by 2.5 to 4 μm, mean ± SD = 4.3 ± 1.1 by 2.9 ± 0.8 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (20)21.5 to 24.5(27) by (6)6.5 to 7.5(8) μm, mean ± SD = 23 ± 1.7 by 7 ± 0.5 μm, L/W = 2.6 to 3.8; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3)3.5 to 5(6) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (4)4.5 to 6(6.5) μm long, the second cell (4.5)5 to 5.5(6) μm long, the third cell (4)4.5 to 5.5(6) μm long, together (13.5)14.5 to 16.5(18) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (2.5)3 to 4.5(5) μm long; basal appendage unbranched, tubular, centric, straight or slightly bent, (2)3 to 5.5(7) μm long, mean ± SD = 4.3 ± 1.3 μm; apical appendages, 2 or 3, unbranched, tubular, centric, straight or slightly bent, (8)9.5 to 14.5(17.5) μm long, mean ± SD = 12 ± 2.4 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white to amber, reaching a 65-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, pale gray, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Materials examined: China, Guizhou Province, Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, on diseased leaves of Cyclobalanopsis augustinii, 23 November 2019, Dan-ran Bian (CFCC 55052); Shaanxi Province, Hanzhong City, Foping County, Dongshan Mountain, on diseased leaves of Quercus aliena, 7 September 2019, Yong Li (CFCC 54733).

Notes: Two new isolates of Pestalotiopsis rhodomyrtus were collected from Cyclobalanopsis augustinii and Quercus aliena, which agree with the type specimen in conidial dimension and characters (30). Hence, Cyclobalanopsis augustinii and Quercus aliena become new hosts for Pestalotiopsis rhodomyrtus, which was originally described from Rhodomyrtus tomentosa (Myrtaceae) (30).

Pestalotiopsis shaanxiensis Ning Jiang, sp. nov. (Fig. 10). MycoBank number MB843390. Etymology: named after the collection site of the type specimen, Shaanxi Province. Typus: China, Shaanxi Province, Hanzhong City, Foping County, Dongshan Mountain, on diseased leaves of Quercus variabilis, 7 September 2019, Yong Li (holotype CAF 800047; ex-holotype culture CFCC 54958).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 100 to 550 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 7 to 15 by 2 to 5.5 μm, mean ± SD = 10.5 ± 3 by 4.3 ± 1.1 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (21)22 to 24.5(25) by (7)7.5 to 8.5(9) μm, mean ± SD = 23.1 ± 1.3 by 8 ± 0.4 μm, L/W = 2.5 to 3.3; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (3.5)4 to 5(5.5) μm; median cells 3, trapezoid or subcylindrical, concolorous, pale brown to brown, thick-walled, the first median cell from base (4)4.5 to 5.5(6) μm long, the second cell (4.5)5 to 5.5(6) μm long, the third cell (4.5)5 to 5.5(6) μm long, together (13.5)14.5 to 16.5(17.5) μm long; apical cell conic with an acute apex, thin-walled, hyaline or pale brown, (3)3.5 to 4.5(5) μm long; basal appendage single, unbranched, tubular, centric, straight or slightly bent, (1.5)2.5 to 6(7.5) μm long, mean ± SD = 4.2 ± 1.6 μm; apical appendages, 3, unbranched, tubular, centric, straight or slightly bent, (13)13.5 to 18(22) μm long, mean ± SD = 15.6 ± 2.3 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and undulate edge, white to pale luteous, reaching a 40-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white, reaching a 70-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional material examined, China, Shaanxi Province, Hanzhong City, Foping County, Dongshan Mountain, on diseased leaves of Quercus variabilis, 7 September 2019, Yong Li (CFCC 57356).

Notes: Two isolates of Pestalotiopsis shaanxiensis from Quercus variabilis formed a distinct clade phylogenetically close to P. biciliata and P. camelliae-oleiferae (Fig. 1). Morphologically, P. shaanxiensis has obviously wider conidia than P. camelliae-oleiferae (7 to 9 μm in P. shaanxiensis versus 5 to 7 μm in P. camelliae-oleiferae) (28). P. shaanxiensis can be distinguished from P. biciliata by one versus two basal appendages in the latter (16).

Pestalotiopsis silvicola Ning Jiang, sp. nov. (Fig. 11). MycoBank number MB843391. Etymology: silva = “forest” and -cola = “inhabiting”; in reference to its woody host. Typus: China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis kerrii, 30 March 2019, Yong Li (holotype CAF 800048; ex-holotype culture CFCC 55296).

Conidiomata in culture sporodochial, aggregated or solitary, erumpent, pulvinate, black, 50 to 450 μm in diameter, exuding black conidial masses. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to spherical, annelidic, 7.5 to 15 by 3 to 6 μm, mean ± SD = 10 ± 2.2 by 4 ± 1 μm. Conidia fusoid, straight or slightly curved, 4-septate, smooth, slightly constricted at the septa, (20)21 to 24(26) by (6)6.5 to 7.5(8.5) μm, mean ± SD = 22.5 ± 1.8 by 7 ± 0.7 μm, L/W = 2.6 to 3.8; basal cell obconic with a truncate base, thin-walled, hyaline or pale brown, (2.5)4 to 5.5(6) μm; median cells 3, trapezoid or subcylindrical, concolorous, brown, thick-walled, the first median cell from base (4)4.5 to 5 μm long, the second cell (3.5)4 to 5(6) μm long, the third cell 4 to 4.5(5.5) μm long, together (12)13 to 15(16) μm long; apical cell conic with an acute apex, thin-walled, hyaline, (3)4 to 5(5.5) μm long; basal appendages, 1 to 2, unbranched, tubular, centric, straight or slightly bent, (3.5)4.5 to 7(8) μm long, mean ± SD = 5.6 ± 1.1 μm; apical appendages, 3, unbranched, tubular, centric, straight or bent, (12)12.5 to 18.5(22.5) μm long, mean ± SD = 15.6 ± 2.9 μm. Sexual morph unknown.

Colonies on MEA flat, spreading, with flocculent aerial mycelium forming concentric rings and entire edge, white, reaching a 60-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses; on PDA, flat, spreading, with flocculent aerial mycelium and undulate edge, white, reaching a 60-mm diameter after 10 days at 25°C, forming black conidiomata with black conidial masses.

Additional materials examined, China, Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Cyclobalanopsis kerrii, 30 March 2019, Yong Li (CFCC 54915 and CFCC 57363).

Notes: Three isolates of Pestalotiopsis silvicola from Cyclobalanopsis kerrii clustered into a distinct clade phylogenetically close to P. aggestorum, P. colombiensis, and P. jinchanghensis (Fig. 1). Morphologically, they share similar conidial characters. However, P. silvicola can be distinguished by sequence data (nucleotide differences from P. aggestorum: in the ITS, 1/505 [0.2%]; in tef1, 1/464 [0.22%]; in tub2, 1/441 [0.23%]; from P. colombiensis: in the ITS, 2/505 [0.4%]; in tef1, 8/464 [1.72%]; in tub2, 2/441 [0.45%]; from P. jinchanghensis: in the ITS, 1/500 [0.2%]; in tef1, 18/464 [3.88%], 4-bp insertions; in tub2, 1/441 [0.23%]).

DISCUSSION

In the present study, 43 isolates from Fagaceae leaf spots in China belonging to 16 Pestalotiopsis species were characterized. However, we investigated only 20 Fagaceae hosts of more than 320 reported species in China, and several provinces were not deeply investigated. Based on morphology and phylogeny, 10 of the 16 species described here have been proven to be new, indicating that many hidden Pestalotiopsis species may remain to be discovered from Fagaceae in the future.

Species of Pestalotiopsis are usually isolated from plant leaves (7, 8, 16, 18, 25). Two plurivorous species, Pestalotiopsis kenyana and P. monochaeta, were previously recorded from Castanea henryi and C. mollissima in China and from Quercus robur in the Netherlands, respectively (6, 16). These two species are not host-specific as they have been reported from various unrelated hosts, such as Camellia sinensis (Theaceae), Coffea sp. (Rubiaceae), and Taxus baccata (Taxaceae) (6, 16, 18). In the present study, 15 additional species are newly recorded from leaves of Fagaceae (Table 1), of which 10 are so far known only from fagaceous hosts.

Among pestalotioid species, appendages vary in number, origin, position, numbers of branches and the branching patterns (8). These characters have been proven appropriate and useful in delineating certain genera (8). For example, Seimatosporium is different from Sporocadus in that it forms appendages (9). However, Pestalotiopsis changjiangensis discovered in the present study is characterized by having only a single short, indistinct apical appendage, which is unique in Pestalotiopsis (Fig. 4).

This study revealed 16 Pestalotiopsis species associated with Fagaceae leaf spot symptoms in China. Further studies are now required, however, to confirm their pathogenicity.

MATERIALS AND METHODS

Sample collection and isolation.

Fresh specimens of diseased fagaceous leaves were collected from Anhui, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hunan, Jiangxi, Shaanxi, and Sichuan Provinces in China from 2016 to 2021. A total of four host genera, 18 species, and one variety were investigated in the present study, viz., Castanopsis faberi, C. fissa, C. hainanensis, C. hystrix, C. lamontii, C. tonkinensis, Cyclobalanopsis augustinii, Cy. austrocochinchinensis, Cy. fleuryi, Cy. glauca, Cy. kerrii, Cy. neglecta, Lithocarpus amygdalifolius, L. chiungchungensis, Quercus aliena, Q. aliena var. acutiserrata, Q. griffithii, Q. serrata, and Q. variabilis. The leaf samples were packed in paper bags and transferred to the laboratory for fungal isolation.

The leaf samples with typical spot symptoms were first surface sterilized for 1 min in 75% ethanol, 3 min in 1.25% sodium hypochlorite, and 1 min in 75% ethanol, then rinsed for 2 min in distilled water, and blotted on dry sterile filter paper. Then, the diseased areas of the leaves were cut into 0.5- by 0.5-cm pieces using an aseptic razor blade, transferred onto the surface of potato dextrose agar (PDA; 200 g potatoes, 20 g dextrose, 20 g agar per L) and malt extract agar (MEA; 30 g malt extract, 5 g mycological peptone, 15 g agar per L) plates, and incubated at 25°C to obtain fungal hyphae. Hyphal tips were then removed to new PDA plates to obtain pure cultures. The cultures were deposited in the China Forestry Culture Collection Center (CFCC; http://cfcc.caf.ac.cn/) and the specimens in the herbarium of the Chinese Academy of Forestry (CAF; http://museum.caf.ac.cn/).

DNA extraction, sequencing, and phylogenetic analyses.

Genomic DNA was extracted from colonies grown on cellophane-covered PDA using a CTAB (cetyltrimethylammonium bromide) method (31). The amount of DNA was estimated by electrophoresis in 1% agarose gels, and the quality was measured using a NanoDrop 2000 instrument (Thermo Scientific, Waltham, MA, USA) following the user manual.

The following primer pairs were used for amplification of the gene regions sequenced in the present study: ITS1/ITS4 for the 5.8S nuclear ribosomal DNA gene with the two flanking internally transcribed spacer (ITS1 and ITS2) regions (32), EF1-728F/EF2 for the translation elongation factor 1-α (tef1) gene (33, 34), and T1/Bt2b and Bt2a/Bt2b for the beta-tubulin (tub2) gene (35, 36). The PCR conditions were set as follows: an initial denaturation step of 5 min at 94°C, followed by 35 cycles of 30 s at 94°C, 50 s at 52°C (ITS) or 54°C (tef1 and tub2), and 1 min at 72°C, and a final elongation step of 7 min at 72°C. PCR amplification products were checked via electrophoresis in 2% agarose gels. DNA sequencing was performed using an ABI Prism 3730XL DNA analyzer with a BigDye Terminator kit v.3.1 (Invitrogen, USA) at the Shanghai Invitrogen Biological Technology Company Limited (Beijing, China).

The quality of the amplified nucleotide sequences was checked and the sequences assembled using SeqMan v.7.1.0. Reference sequences were retrieved from the National Center for Biotechnology Information (NCBI). Sequences were aligned using MAFFT v. 6 (37) and corrected manually using MEGA 6 (38). The phylogenetic analyses of the combined matrices were performed using maximum-likelihood (ML) and Bayesian inference (BI) methods. ML was implemented on the CIPRES Science Gateway portal (https://www.phylo.org) using RAxML-HPC BlackBox 8.2.10 (39), employing a GTRGAMMA substitution model with 1,000 bootstrap replicates, while BI was performed using a Markov chain Monte Carlo (MCMC) algorithm in MrBayes v. 3.0 (40). Two MCMC chains were run, starting from random trees, for 1,000,000 generations, and trees were sampled every 100th generation, resulting in a total of 10,000 trees. The first 25% of trees were discarded as burn-in of each analysis. Branches with significant Bayesian posterior probabilities (BPP) were estimated in the remaining 7,500 trees. Phylogenetic trees were viewed with FigTree v.1.3.1 and graphically processed by Adobe Illustrator CS5.

For closely related species with similar morphology, ITS, tef1, and tub2 sequences of the respective species were compared pairwise. For this, the sequences of species pairs were aligned and the parts containing leading/trailing gaps were removed. Sequence differences of this alignment are recorded in the following way: number of nucleotide substitutions (excluding insertions and gaps)/total number of nucleotide characters, percent of sequence substitutions (in brackets), and the number of base pair insertions and gaps.

Morphology.

The morphological data of the isolates collected in the present study were obtained from sporulating pure cultures grown on PDA or MEA in the dark at 25°C. The conidiomata were observed and photographed using a dissecting microscope (M205 C; Leica, Wetzlar, Germany). Microscope slides of conidiogenous cells and conidia were prepared in tap water, and the slides were examined and photographed with an Axio Imager 2 microscope (Zeiss, Oberkochen, Germany) equipped with an AxioCam 506 color camera or a Nikon Eclipse 80i microscope (Nikon, Tokyo, Japan) equipped with a Nikon digital sight DS-Ri2 camera, using differential interference contrast (DIC) illumination. For measurements, 50 conidia were randomly selected. Measurements of the conidia are reported as maximum and minimum in parentheses and the range representing the mean ± standard deviation of the measurements given in parentheses. Culture characteristics were recorded from 9-cm PDA or MEA plates after 10 days incubation at 25°C in the dark. To enable comparison of species growing on fagaceous hosts, available measurement data and sequence data are summarized in Table 1.

Data availability.

The nucleotide sequence data from the present study were deposited in GenBank, and the accession numbers are listed in Table 2.

TABLE 2.

Isolates and GenBank accession numbers used for phylogenetic analyses in this study^a

Species	Isolate	Host/substrate	Origin	GenBank accession no.
Species	Isolate	Host/substrate	Origin	ITS	tub2	tef1
Neopestalotiopsis magna	MFLUCC 12-0652^b	Pteridium sp.	France	KF582795	KF582793	KF582791
Pestalotiopsis abietis	CFCC 53011^b	Abies fargesii	China	MK397013	MK622280	MK622277
	CFCC 53012	Abies fargesii	China	MK397014	MK622281	MK622278
P. adusta	ICMP 6088^b	Refrigerator door	Fiji	JX399006	JX399037	JX399070
	MFLUCC 10-146	Syzygium sp.	Thailand	JX399007	JX399038	JX399071
P. aggestorum	LC6301^b	Camellia sinensis	China	KX895015	KX895348	KX895234
	LC8186	Camellia sinensis	China	KY464140	KY464160	KY464150
P. anacardiacearum	IFRDCC 2397^b	Mangifera indica	China	KC247154	KC247155	KC247156
*P. anhuiensis*	CFCC 54791 ^b	*Cyclobalanopsis glauca*	China	ON007028	ON005056	ON005045
P. arceuthobii	CBS 434.65^b	Arceuthobium campylopodum	USA	KM199341	KM199427	KM199516
P. arenga	CBS 331.92^b	Arenga undulatifolia	Singapore	KM199340	KM199426	KM199515
P. australasiae	CBS 114126^b	Knightia sp.	New Zealand	KM199297	KM199409	KM199499
	CBS 114141	Protea sp.	New South Wales	KM199298	KM199410	KM199501
P. australis	CBS 111503	Protea neriifolia × susannae	South Africa	KM199331	KM199382	KM199557
	CBS 114193^b	Grevillea sp.	New South Wales	KM199332	KM199383	KM199475
P. biciliata	CBS 124463^b	Platanus × hispanica	Slovakia	KM199308	KM199399	KM199505
	CBS 236.38	Paeonia sp.	Italy	KM199309	KM199401	KM199506
P. brachiata	LC2988^b	Camellia sp.	China	KX894933	KX895265	KX895150
	LC8188	Camellia sp.	China	KY464142	KY464162	KY464152
	LC8189	Camellia sp.	China	KY464143	KY464163	KY464153
P. brassicae	CBS 170.26^b	Brassica napus	New Zealand	KM199379	NA	KM199558
P. camelliae	MFLUCC 12-0277^b	Camellia japonica	China	JX399010	JX399041	JX399074
P. camelliae-oleiferae	CSUFTCC08^b	Camelliae oleiferae	China	OK493593	OK562368	OK507963
	CSUFTCC09	Camelliae oleiferae	China	OK493594	OK562369	OK507964
*P. castanopsidis*	CFCC 54430 ^b	*Castanopsis lamontii*	China	OK339732	OK358508	OK358493
	CFCC 54305	*Castanopsis hystrix*	China	OK339733	OK358509	OK358494
	CFCC 54384	*Castanopsis hystrix*	China	OK339734	OK358510	OK358495
P. chamaeropis	CBS 186.71^b	Chamaerops humilis	Italy	KM199326	KM199391	KM199473
	LC3619	Camellia sp.	China	KX894991	KX895322	KX895208
	CFCC 55124	Quercus acutissima	China	OM746221	OM839894	OM839993
	CFCC 54977	Quercus acutissima	China	OM746223	OM839896	OM839995
	CFCC 55019	Quercus aliena	China	OM746224	OM839897	OM839996
	CFCC 55122	Quercus aliena	China	OM746229	OM839902	OM840001
	CFCC 55023	*Castanopsis fissa*	China	OM746233	OM839906	OM840005
	CFCC 54776	*Quercus variabilis*	China	OM746234	OM839907	OM840006
	CFCC 55338	*Quercus variabilis*	China	OM746235	OM839908	OM840007
*P. changjiangensis*	CFCC 54314 ^b	*Castanopsis tonkinensis*	China	OK339739	OK358515	OK358500
	CFCC 54433	*Castanopsis hainanensis*	China	OK339740	OK358516	OK358501
	CFCC 52803	*Cyclobalanopsis austrocochinchinensis*	China	OK339741	OK358517	OK358502
P. clavata	MFLUCC 12-0268^b	Buxus sp.	China	JX398990	JX399025	JX399056
P. colombiensis	CBS 118553^b	Eucalyptus urograndis	Colombia	KM199307	KM199421	KM199488
*P. cyclobalanopsidis*	CFCC 54328 ^b	*Cyclobalanopsis glauca*	China	OK339735	OK358511	OK358496
	CFCC 55891	*Cyclobalanopsis glauca*	China	OK339736	OK358512	OK358497
P. digitalis	MFLU 14-0208^b	Digitalis purpurea	New Zealand	KP781879	KP781883	NA
P. dilucida	LC3232	Camellia sinensis	China	KX894961	KX895293	KX895178
P. dilucida	LC8184	Camellia sinensis	China	KY464138	KY464158	KY464148
P. diploclisiae	CBS 115449	Psychotria tutcheri	China	KM199314	KM199416	KM199485
	CBS 115587^b	Diploclisia glaucescens	China	KM199320	KM199419	KM199486
P. disseminata	CBS 143904	Persea americana	New Zealand	MH554152	MH554825	MH554587
	MEAN 1165	Pinus pinea	Portugal	MT374687	MT374712	MT374699
P. diversiseta	MFLUCC 12-0287^b	Rhododendron sp.	China	JX399009	JX399040	JX399073
P. doitungensis	MFLUCC 14-0115^b	Dendrobium sp.	Thailand	MK993574	MK975837	MK975832
P. dracaenicola	MFLUCC 18-0913^b	Dracaena sp.	Thailand	MN962731	MN962733	MN962732
P. dracontomelonis	MFLU 14-0207^b	Dracontomelon dao	Thailand	KP781877	NA	KP781880
P. ericacearum	IFRDCC 2439^b	Rhododendron delavayi	China	KC537807	KC537821	KC537814
P. etonensis	BRIP 66615^b	Sporobolus jacquemontii	Australia	MK966339	MK977634	MK977635
P. formosana	NTUCC 17-009^b	Poaceae sp.	China	MH809381	MH809385	MH809389
P. furcata	MFLUCC 12-0054^b	Camellia sinensis	Thailand	JQ683724	JQ683708	JQ683740
	LC6691	Camellia sinensis	China	KX895030	KX895363	KX895248
*P. foliicola*	CFCC 54440 ^b	*Castanopsis faberi*	China	ON007029	ON005057	ON005046
	CFCC 57359	*Castanopsis faberi*	China	ON007030	ON005058	ON005047
	CFCC 57360	*Castanopsis faberi*	China	ON007031	ON005059	ON005048
P. gaultheriae	IFRD 411-014^b	Gaultheria forrestii	China	KC537805	KC537819	KC537812
P. gibbosa	NOF 3175^b	Gaultheria shallon	Canada	LC311589	LC311590	LC311591
P. grevilleae	CBS 114127^b	Grevillea sp.	Australia	KM199300	KM199407	KM199504
*P. guangxiensis*	CFCC 54308 ^b	*Quercus griffithii*	China	OK339737	OK358513	OK358498
	CFCC 54300	*Quercus griffithii*	China	OK339738	OK358514	OK358499
*P. guizhouensis*	CFCC 54803 ^b	*Cyclobalanopsis glauca*	China	ON007035	ON005063	ON005052
	CFCC 57364	*Cyclobalanopsis glauca*	China	ON007036	ON005064	ON005053
P. hawaiiensis	CBS 114491^b	Leucospermum sp.	USA	KM199339	KM199428	KM199514
P. hispanica	CBS 115391^b	Protea sp.	Spain	MH553981	MH554640	MH554399
P. hollandica	CBS 265.33^b	Sciadopitys verticillata	Netherlands	KM199328	KM199388	KM199481
P. humicola	CBS 336.97^b	Soil	Papua New Guinea	KM199317	KM199420	KM199484
P. hunanensis	CSUFTCC15^b	Camellia oleifera	China	OK493599	OK562374	OK507969
	CSUFTCC18	Camellia oleifera	China	OK493600	OK562375	OK507970
P. inflexa	MFLUCC 12-0270^b	Unidentified tree	China	JX399008	JX399039	JX399072
P. intermedia	MFLUCC 12-0259^b	Unidentified tree	China	JX398993	JX399028	JX399059
P. italiana	MFLU 14-0214^b	Cupressus glabra	Italy	KP781878	KP781882	KP781881
P. jesteri	CBS 109350^b	Fragraea bodenii	Papua New Guinea	KM199380	KM199468	KM199554
P. jiangxiensis	LC4399^b	Camellia sp.	China	KX895009	KX895341	KX895227
P. jinchanghensis	LC6636^b	Camellia sinensis	China	KX895028	KX895361	KX895247
	LC8190	Camellia sinensis	China	KY464144	KY464164	KY464154
P. kaki	KNU-PT-1804^b	Diospyros kaki	Korea	LC552953	LC552954	LC553555
P. kandelicola	NCYU 19-0355^b	Kandelia candel	China	MT560723	MT563100	MT563102
P. kenyana	CBS 442.67^b	Coffea sp.	Kenya	KM199302	KM199395	KM199502
	LC6633	Camellia sinensis	China	KX895027	KX895360	KX895246
	CFCC 54962	*Quercus aliena*	China	OM746237	OM839910	OM840009
	CFCC 55330	*Cyclobalanopsis fleuryi*	China	OM746238	OM839911	OM840010
	CFCC 54621	**Quercus aliena var. acutiserrata**	China	OM746239	OM839912	OM840011
	CFCC 54732	*Cyclobalanopsis neglecta*	China	OM746243	OM839916	OM840015
	CFCC 54742	*Castanopsis hystrix*	China	OM746245	OM839918	OM840017
	CFCC 54618	**Quercus aliena var. acutiserrata**	China	OM746248	OM839921	OM840020
	CFCC 54805	*Cyclobalanopsis glauca*	China	OM746253	OM839926	OM840025
	CFCC 55088	*Castanopsis fissa*	China	OM746254	OM839927	OM840026
P. knightiae	CBS 111963	Knightia sp.	New Zealand	KM199311	KM199406	KM199495
	CBS 114138^b	Knightia sp.	New Zealand	KM199310	KM199408	KM199497
P. krabiensis	MFLUCC 16-0260^b	Pandanus sp.	Thailand	MH388360	MH412722	MH388395
P. leucadendri	CBS 121417^b	Leucadendron sp.	South Africa	MH553987	MH554654	MH554412
P. licualicola	HGUP 4057^b	Licuala grandis	China	KC492509	KC481683	KC481684
P. lijiangensis	CFCC 50738^b	Castanopsis carlesii var. spinulosa	China	KU860520	KU844184	KU844185
P. linearis	MFLUCC 12-0271^b	Trachelospermum sp.	China	JX398992	JX399027	JX399058
*P. lithocarpi*	CFCC 55100 ^b	*Lithocarpus chiungchungensis*	China	OK339742	OK358518	OK358503
	CFCC 55893	*Lithocarpus chiungchungensis*	China	OK339743	OK358519	OK358504
P. lushanensis	LC4344^b	Camellia sp.	China	KX895005	KX895337	KX895223
	LC8182	Camellia sp.	China	KY464136	KY464156	KY464146
	LC8183	Camellia sp.	China	KY464137	KY464157	KY464147
	CFCC 54894	*Quercus serrata*	China	OM746282	OM839955	OM840054
P. macadamiae	BRIP 63738b	Macadamia integrifolia	Australia	KX186588	KX186680	KX186621
	BRIP 63739b	Macadamia integrifolia	Australia	KX186587	KX186679	KX186620
	BRIP 637441a	Macadamia integrifolia	Australia	KX186586	KX186678	KX186619
P. malayana	CBS 102220^b	Macaranga triloba	Malaysia	KM199306	KM199411	KM199482
P. monochaeta	CBS 144.97^b	Quercus robur	Netherlands	KM199327	KM199386	KM199479
	CBS 440.83	Taxus baccata	Netherlands	KM199329	KM199387	KM199480
*P. nanjingensis*	CFCC 53882	*Quercus aliena*	China	OM746295	OM839968	OM840067
	CSUFTCC16^b	Camellia oleifera	China	OK493602	OK562377	OK507972
P. nanningensis	CSUFTCC10^b	Camellia oleifera	China	OK493596	OK562371	OK507966
P. neolitseae	NTUCC 17-011^b	Neolitsea villosa	China	MH809383	MH809387	MH809391
	CFCC 54590	*Lithocarpus amygdalifolius*	China	OK339744	OK358520	OK358505
P. novae-hollandiae	CBS 130973^b	Banksia grandis	Australia	KM199337	KM199425	KM199511
P. oryzae	CBS 111522	Telopea sp.	USA	KM199294	KM199394	KM199493
	CBS 171.26	NA	Italy	KM199304	KM199397	KM199494
	CBS 353.69^b	Oryza sativa	Denmark	KM199299	KM199398	KM199496
P. pallidotheae	MAFF 240993^b	Pieris japonica	Japan	AB482220	NA	NA
P. pandanicola	MFLUCC 16-0255^b	Pandanus sp.	Thailand	MH388361	MH412723	MH388396
P. papuana	CBS 331.96^b	Coastal soil	Papua New Guinea	KM199321	KM199413	KM199491
	CBS 887.96	Cocos nucifera	Papua New Guinea	KM199318	KM199415	KM199492
P. parva	CBS 265.37	Delonix regia	NA	KM199312	KM199404	KM199508
	CBS 278.35^b	Delonix regia	NA	KM199313	KM199405	KM199509
P. photiniicola	GZCC 16-0028^b	Photinia serrulata	China	KY092404	KY047663	KY047662
P. pini	MEAN 1092	Pinus pinea	Portugal	MT374680	MT374705	MT374693
P. pinicola	KUMCC 19-0183^b	Pinus armandii	China	MN412636	MN417507	MN417509
P. portugalica	CBS 393.48^b	NA	Portugal	KM199335	KM199422	KM199510
P. rhizophorae	MFLUCC 17-0416^b	Rhizophora mucronata	Thailand	MK764283	MK764349	MK764327
P. rhododendri	IFRDCC 2399^b	Rhododendron sinogrande	China	KC537804	KC537818	KC537811
P. rhodomyrtus	LC4458	Camellia sinensis	China	KX895010	KX895342	KX895228
	HGUP4230^b	Rhodomyrtus tomentosa	China	KF412648	KF412642	KF412645
	CFCC 54733	*Quercus aliena*	China	OM746310	OM839983	OM840082
	CFCC 55052	*Cyclobalanopsis augustinii*	China	OM746311	OM839984	OM840083
P. rosea	MFLUCC 12-0258^b	Pinus sp.	China	JX399005	JX399036	JX399069
P. scoparia	CBS 176.25^b	Chamaecyparis sp.	China	KM199330	KM199393	KM199478
P. sequoiae	MFLUCC 13-0399^b	Sequoia sempervirens	Italy	KX572339	NA	NA
*P. shaanxiensis*	CFCC 54958 ^b	*Quercus variabilis*	China	ON007026	ON005054	ON005043
	CFCC 57356	*Quercus variabilis*	China	ON007027	ON005055	ON005044
*P. silvicola*	CFCC 55296 ^b	*Cyclobalanopsis kerrii*	China	ON007032	ON005060	ON005049
	CFCC 54915	*Cyclobalanopsis kerrii*	China	ON007033	ON005061	ON005050
	CFCC 57363	*Cyclobalanopsis kerrii*	China	ON007034	ON005062	ON005051
P. spathulata	CBS 356.86^b	Gevuina avellana	Chile	KM199338	KM199423	KM199513
P. spathuliappendiculata	CBS 144035^b	Phoenix canariensis	Australia	MH554172	MH554845	MH554607
P. telopeae	CBS 114137	Protea sp.	Australia	KM199301	KM199469	KM199559
	CBS 114161^b	Telopea sp.	Australia	KM199296	KM199403	KM199500
	CBS 113606	Telopea sp.	Australia	KM199295	KM199402	KM199498
P. terricola	CBS 141.69^b	Soil	Pacific Islands	MH554004	MH554680	MH554438
P. thailandica	MFLUCC 17-1616^b	Rhizophora mucronata	Thailand	MK764285	MK764351	MK764329
P. trachycarpicola	OP068^b	Trachycarpus fortunei	China	JQ845947	JQ845945	JQ845946
	IFRDCC 2403	Podocarpus macrophyllus	China	KC537809	KC537823	KC537816
	LC4523	Camellia sinensis	China	KX895011	KX895344	KX895230
P. unicolor	MFLUCC 12-0276^b	Rhododendron sp.	China	JX398999	JX399030	NA
	MFLUCC 12-0275	Unidentified tree	China	JX398998	JX399029	JX399063
P. verruculosa	MFLUCC 12-0274^b	Rhododendron sp.	China	JX398996	NA	JX399061
P. yanglingensis	LC4553^b	Camellia sinensis	China	KX895012	KX895345	KX895231
	LC3412	Camellia sinensis	China	KX894980	KX895312	KX895197
P. yunnanensis	HMAS 96359^b	Podocarpus macrophyllus	China	AY373375	NA	NA

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Isolates and sequences generated during the present study are in bold. NA, not available.

Ex-type strain.

ACKNOWLEDGMENTS

This research was funded by Fundamental Research Funds for the Central Non-profit Research Institution of Chinese Academy of Forestry (grant CAFYBB2018ZB001), and National Microbial Resource Center of the Ministry of Science and Technology of the People’s Republic of China (grant NMRC-2021-7).

Contributor Information

Yong Li, Email: lylx@caf.ac.cn.

Florian M. Freimoser, Agroscope

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The nucleotide sequence data from the present study were deposited in GenBank, and the accession numbers are listed in Table 2.

TABLE 2.

Isolates and GenBank accession numbers used for phylogenetic analyses in this study^a

Species	Isolate	Host/substrate	Origin	GenBank accession no.
Species	Isolate	Host/substrate	Origin	ITS	tub2	tef1
Neopestalotiopsis magna	MFLUCC 12-0652^b	Pteridium sp.	France	KF582795	KF582793	KF582791
Pestalotiopsis abietis	CFCC 53011^b	Abies fargesii	China	MK397013	MK622280	MK622277
	CFCC 53012	Abies fargesii	China	MK397014	MK622281	MK622278
P. adusta	ICMP 6088^b	Refrigerator door	Fiji	JX399006	JX399037	JX399070
	MFLUCC 10-146	Syzygium sp.	Thailand	JX399007	JX399038	JX399071
P. aggestorum	LC6301^b	Camellia sinensis	China	KX895015	KX895348	KX895234
	LC8186	Camellia sinensis	China	KY464140	KY464160	KY464150
P. anacardiacearum	IFRDCC 2397^b	Mangifera indica	China	KC247154	KC247155	KC247156
*P. anhuiensis*	CFCC 54791 ^b	*Cyclobalanopsis glauca*	China	ON007028	ON005056	ON005045
P. arceuthobii	CBS 434.65^b	Arceuthobium campylopodum	USA	KM199341	KM199427	KM199516
P. arenga	CBS 331.92^b	Arenga undulatifolia	Singapore	KM199340	KM199426	KM199515
P. australasiae	CBS 114126^b	Knightia sp.	New Zealand	KM199297	KM199409	KM199499
	CBS 114141	Protea sp.	New South Wales	KM199298	KM199410	KM199501
P. australis	CBS 111503	Protea neriifolia × susannae	South Africa	KM199331	KM199382	KM199557
	CBS 114193^b	Grevillea sp.	New South Wales	KM199332	KM199383	KM199475
P. biciliata	CBS 124463^b	Platanus × hispanica	Slovakia	KM199308	KM199399	KM199505
	CBS 236.38	Paeonia sp.	Italy	KM199309	KM199401	KM199506
P. brachiata	LC2988^b	Camellia sp.	China	KX894933	KX895265	KX895150
	LC8188	Camellia sp.	China	KY464142	KY464162	KY464152
	LC8189	Camellia sp.	China	KY464143	KY464163	KY464153
P. brassicae	CBS 170.26^b	Brassica napus	New Zealand	KM199379	NA	KM199558
P. camelliae	MFLUCC 12-0277^b	Camellia japonica	China	JX399010	JX399041	JX399074
P. camelliae-oleiferae	CSUFTCC08^b	Camelliae oleiferae	China	OK493593	OK562368	OK507963
	CSUFTCC09	Camelliae oleiferae	China	OK493594	OK562369	OK507964
*P. castanopsidis*	CFCC 54430 ^b	*Castanopsis lamontii*	China	OK339732	OK358508	OK358493
	CFCC 54305	*Castanopsis hystrix*	China	OK339733	OK358509	OK358494
	CFCC 54384	*Castanopsis hystrix*	China	OK339734	OK358510	OK358495
P. chamaeropis	CBS 186.71^b	Chamaerops humilis	Italy	KM199326	KM199391	KM199473
	LC3619	Camellia sp.	China	KX894991	KX895322	KX895208
	CFCC 55124	Quercus acutissima	China	OM746221	OM839894	OM839993
	CFCC 54977	Quercus acutissima	China	OM746223	OM839896	OM839995
	CFCC 55019	Quercus aliena	China	OM746224	OM839897	OM839996
	CFCC 55122	Quercus aliena	China	OM746229	OM839902	OM840001
	CFCC 55023	*Castanopsis fissa*	China	OM746233	OM839906	OM840005
	CFCC 54776	*Quercus variabilis*	China	OM746234	OM839907	OM840006
	CFCC 55338	*Quercus variabilis*	China	OM746235	OM839908	OM840007
*P. changjiangensis*	CFCC 54314 ^b	*Castanopsis tonkinensis*	China	OK339739	OK358515	OK358500
	CFCC 54433	*Castanopsis hainanensis*	China	OK339740	OK358516	OK358501
	CFCC 52803	*Cyclobalanopsis austrocochinchinensis*	China	OK339741	OK358517	OK358502
P. clavata	MFLUCC 12-0268^b	Buxus sp.	China	JX398990	JX399025	JX399056
P. colombiensis	CBS 118553^b	Eucalyptus urograndis	Colombia	KM199307	KM199421	KM199488
*P. cyclobalanopsidis*	CFCC 54328 ^b	*Cyclobalanopsis glauca*	China	OK339735	OK358511	OK358496
	CFCC 55891	*Cyclobalanopsis glauca*	China	OK339736	OK358512	OK358497
P. digitalis	MFLU 14-0208^b	Digitalis purpurea	New Zealand	KP781879	KP781883	NA
P. dilucida	LC3232	Camellia sinensis	China	KX894961	KX895293	KX895178
P. dilucida	LC8184	Camellia sinensis	China	KY464138	KY464158	KY464148
P. diploclisiae	CBS 115449	Psychotria tutcheri	China	KM199314	KM199416	KM199485
	CBS 115587^b	Diploclisia glaucescens	China	KM199320	KM199419	KM199486
P. disseminata	CBS 143904	Persea americana	New Zealand	MH554152	MH554825	MH554587
	MEAN 1165	Pinus pinea	Portugal	MT374687	MT374712	MT374699
P. diversiseta	MFLUCC 12-0287^b	Rhododendron sp.	China	JX399009	JX399040	JX399073
P. doitungensis	MFLUCC 14-0115^b	Dendrobium sp.	Thailand	MK993574	MK975837	MK975832
P. dracaenicola	MFLUCC 18-0913^b	Dracaena sp.	Thailand	MN962731	MN962733	MN962732
P. dracontomelonis	MFLU 14-0207^b	Dracontomelon dao	Thailand	KP781877	NA	KP781880
P. ericacearum	IFRDCC 2439^b	Rhododendron delavayi	China	KC537807	KC537821	KC537814
P. etonensis	BRIP 66615^b	Sporobolus jacquemontii	Australia	MK966339	MK977634	MK977635
P. formosana	NTUCC 17-009^b	Poaceae sp.	China	MH809381	MH809385	MH809389
P. furcata	MFLUCC 12-0054^b	Camellia sinensis	Thailand	JQ683724	JQ683708	JQ683740
	LC6691	Camellia sinensis	China	KX895030	KX895363	KX895248
*P. foliicola*	CFCC 54440 ^b	*Castanopsis faberi*	China	ON007029	ON005057	ON005046
	CFCC 57359	*Castanopsis faberi*	China	ON007030	ON005058	ON005047
	CFCC 57360	*Castanopsis faberi*	China	ON007031	ON005059	ON005048
P. gaultheriae	IFRD 411-014^b	Gaultheria forrestii	China	KC537805	KC537819	KC537812
P. gibbosa	NOF 3175^b	Gaultheria shallon	Canada	LC311589	LC311590	LC311591
P. grevilleae	CBS 114127^b	Grevillea sp.	Australia	KM199300	KM199407	KM199504
*P. guangxiensis*	CFCC 54308 ^b	*Quercus griffithii*	China	OK339737	OK358513	OK358498
	CFCC 54300	*Quercus griffithii*	China	OK339738	OK358514	OK358499
*P. guizhouensis*	CFCC 54803 ^b	*Cyclobalanopsis glauca*	China	ON007035	ON005063	ON005052
	CFCC 57364	*Cyclobalanopsis glauca*	China	ON007036	ON005064	ON005053
P. hawaiiensis	CBS 114491^b	Leucospermum sp.	USA	KM199339	KM199428	KM199514
P. hispanica	CBS 115391^b	Protea sp.	Spain	MH553981	MH554640	MH554399
P. hollandica	CBS 265.33^b	Sciadopitys verticillata	Netherlands	KM199328	KM199388	KM199481
P. humicola	CBS 336.97^b	Soil	Papua New Guinea	KM199317	KM199420	KM199484
P. hunanensis	CSUFTCC15^b	Camellia oleifera	China	OK493599	OK562374	OK507969
	CSUFTCC18	Camellia oleifera	China	OK493600	OK562375	OK507970
P. inflexa	MFLUCC 12-0270^b	Unidentified tree	China	JX399008	JX399039	JX399072
P. intermedia	MFLUCC 12-0259^b	Unidentified tree	China	JX398993	JX399028	JX399059
P. italiana	MFLU 14-0214^b	Cupressus glabra	Italy	KP781878	KP781882	KP781881
P. jesteri	CBS 109350^b	Fragraea bodenii	Papua New Guinea	KM199380	KM199468	KM199554
P. jiangxiensis	LC4399^b	Camellia sp.	China	KX895009	KX895341	KX895227
P. jinchanghensis	LC6636^b	Camellia sinensis	China	KX895028	KX895361	KX895247
	LC8190	Camellia sinensis	China	KY464144	KY464164	KY464154
P. kaki	KNU-PT-1804^b	Diospyros kaki	Korea	LC552953	LC552954	LC553555
P. kandelicola	NCYU 19-0355^b	Kandelia candel	China	MT560723	MT563100	MT563102
P. kenyana	CBS 442.67^b	Coffea sp.	Kenya	KM199302	KM199395	KM199502
	LC6633	Camellia sinensis	China	KX895027	KX895360	KX895246
	CFCC 54962	*Quercus aliena*	China	OM746237	OM839910	OM840009
	CFCC 55330	*Cyclobalanopsis fleuryi*	China	OM746238	OM839911	OM840010
	CFCC 54621	**Quercus aliena var. acutiserrata**	China	OM746239	OM839912	OM840011
	CFCC 54732	*Cyclobalanopsis neglecta*	China	OM746243	OM839916	OM840015
	CFCC 54742	*Castanopsis hystrix*	China	OM746245	OM839918	OM840017
	CFCC 54618	**Quercus aliena var. acutiserrata**	China	OM746248	OM839921	OM840020
	CFCC 54805	*Cyclobalanopsis glauca*	China	OM746253	OM839926	OM840025
	CFCC 55088	*Castanopsis fissa*	China	OM746254	OM839927	OM840026
P. knightiae	CBS 111963	Knightia sp.	New Zealand	KM199311	KM199406	KM199495
	CBS 114138^b	Knightia sp.	New Zealand	KM199310	KM199408	KM199497
P. krabiensis	MFLUCC 16-0260^b	Pandanus sp.	Thailand	MH388360	MH412722	MH388395
P. leucadendri	CBS 121417^b	Leucadendron sp.	South Africa	MH553987	MH554654	MH554412
P. licualicola	HGUP 4057^b	Licuala grandis	China	KC492509	KC481683	KC481684
P. lijiangensis	CFCC 50738^b	Castanopsis carlesii var. spinulosa	China	KU860520	KU844184	KU844185
P. linearis	MFLUCC 12-0271^b	Trachelospermum sp.	China	JX398992	JX399027	JX399058
*P. lithocarpi*	CFCC 55100 ^b	*Lithocarpus chiungchungensis*	China	OK339742	OK358518	OK358503
	CFCC 55893	*Lithocarpus chiungchungensis*	China	OK339743	OK358519	OK358504
P. lushanensis	LC4344^b	Camellia sp.	China	KX895005	KX895337	KX895223
	LC8182	Camellia sp.	China	KY464136	KY464156	KY464146
	LC8183	Camellia sp.	China	KY464137	KY464157	KY464147
	CFCC 54894	*Quercus serrata*	China	OM746282	OM839955	OM840054
P. macadamiae	BRIP 63738b	Macadamia integrifolia	Australia	KX186588	KX186680	KX186621
	BRIP 63739b	Macadamia integrifolia	Australia	KX186587	KX186679	KX186620
	BRIP 637441a	Macadamia integrifolia	Australia	KX186586	KX186678	KX186619
P. malayana	CBS 102220^b	Macaranga triloba	Malaysia	KM199306	KM199411	KM199482
P. monochaeta	CBS 144.97^b	Quercus robur	Netherlands	KM199327	KM199386	KM199479
	CBS 440.83	Taxus baccata	Netherlands	KM199329	KM199387	KM199480
*P. nanjingensis*	CFCC 53882	*Quercus aliena*	China	OM746295	OM839968	OM840067
	CSUFTCC16^b	Camellia oleifera	China	OK493602	OK562377	OK507972
P. nanningensis	CSUFTCC10^b	Camellia oleifera	China	OK493596	OK562371	OK507966
P. neolitseae	NTUCC 17-011^b	Neolitsea villosa	China	MH809383	MH809387	MH809391
	CFCC 54590	*Lithocarpus amygdalifolius*	China	OK339744	OK358520	OK358505
P. novae-hollandiae	CBS 130973^b	Banksia grandis	Australia	KM199337	KM199425	KM199511
P. oryzae	CBS 111522	Telopea sp.	USA	KM199294	KM199394	KM199493
	CBS 171.26	NA	Italy	KM199304	KM199397	KM199494
	CBS 353.69^b	Oryza sativa	Denmark	KM199299	KM199398	KM199496
P. pallidotheae	MAFF 240993^b	Pieris japonica	Japan	AB482220	NA	NA
P. pandanicola	MFLUCC 16-0255^b	Pandanus sp.	Thailand	MH388361	MH412723	MH388396
P. papuana	CBS 331.96^b	Coastal soil	Papua New Guinea	KM199321	KM199413	KM199491
	CBS 887.96	Cocos nucifera	Papua New Guinea	KM199318	KM199415	KM199492
P. parva	CBS 265.37	Delonix regia	NA	KM199312	KM199404	KM199508
	CBS 278.35^b	Delonix regia	NA	KM199313	KM199405	KM199509
P. photiniicola	GZCC 16-0028^b	Photinia serrulata	China	KY092404	KY047663	KY047662
P. pini	MEAN 1092	Pinus pinea	Portugal	MT374680	MT374705	MT374693
P. pinicola	KUMCC 19-0183^b	Pinus armandii	China	MN412636	MN417507	MN417509
P. portugalica	CBS 393.48^b	NA	Portugal	KM199335	KM199422	KM199510
P. rhizophorae	MFLUCC 17-0416^b	Rhizophora mucronata	Thailand	MK764283	MK764349	MK764327
P. rhododendri	IFRDCC 2399^b	Rhododendron sinogrande	China	KC537804	KC537818	KC537811
P. rhodomyrtus	LC4458	Camellia sinensis	China	KX895010	KX895342	KX895228
	HGUP4230^b	Rhodomyrtus tomentosa	China	KF412648	KF412642	KF412645
	CFCC 54733	*Quercus aliena*	China	OM746310	OM839983	OM840082
	CFCC 55052	*Cyclobalanopsis augustinii*	China	OM746311	OM839984	OM840083
P. rosea	MFLUCC 12-0258^b	Pinus sp.	China	JX399005	JX399036	JX399069
P. scoparia	CBS 176.25^b	Chamaecyparis sp.	China	KM199330	KM199393	KM199478
P. sequoiae	MFLUCC 13-0399^b	Sequoia sempervirens	Italy	KX572339	NA	NA
*P. shaanxiensis*	CFCC 54958 ^b	*Quercus variabilis*	China	ON007026	ON005054	ON005043
	CFCC 57356	*Quercus variabilis*	China	ON007027	ON005055	ON005044
*P. silvicola*	CFCC 55296 ^b	*Cyclobalanopsis kerrii*	China	ON007032	ON005060	ON005049
	CFCC 54915	*Cyclobalanopsis kerrii*	China	ON007033	ON005061	ON005050
	CFCC 57363	*Cyclobalanopsis kerrii*	China	ON007034	ON005062	ON005051
P. spathulata	CBS 356.86^b	Gevuina avellana	Chile	KM199338	KM199423	KM199513
P. spathuliappendiculata	CBS 144035^b	Phoenix canariensis	Australia	MH554172	MH554845	MH554607
P. telopeae	CBS 114137	Protea sp.	Australia	KM199301	KM199469	KM199559
	CBS 114161^b	Telopea sp.	Australia	KM199296	KM199403	KM199500
	CBS 113606	Telopea sp.	Australia	KM199295	KM199402	KM199498
P. terricola	CBS 141.69^b	Soil	Pacific Islands	MH554004	MH554680	MH554438
P. thailandica	MFLUCC 17-1616^b	Rhizophora mucronata	Thailand	MK764285	MK764351	MK764329
P. trachycarpicola	OP068^b	Trachycarpus fortunei	China	JQ845947	JQ845945	JQ845946
	IFRDCC 2403	Podocarpus macrophyllus	China	KC537809	KC537823	KC537816
	LC4523	Camellia sinensis	China	KX895011	KX895344	KX895230
P. unicolor	MFLUCC 12-0276^b	Rhododendron sp.	China	JX398999	JX399030	NA
	MFLUCC 12-0275	Unidentified tree	China	JX398998	JX399029	JX399063
P. verruculosa	MFLUCC 12-0274^b	Rhododendron sp.	China	JX398996	NA	JX399061
P. yanglingensis	LC4553^b	Camellia sinensis	China	KX895012	KX895345	KX895231
	LC3412	Camellia sinensis	China	KX894980	KX895312	KX895197
P. yunnanensis	HMAS 96359^b	Podocarpus macrophyllus	China	AY373375	NA	NA

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Isolates and sequences generated during the present study are in bold. NA, not available.

Ex-type strain.

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PERMALINK

Morphology and Phylogeny of Pestalotiopsis (Sporocadaceae, Amphisphaeriales) from Fagaceae Leaves in China

Ning Jiang

Hermann Voglmayr

Han Xue

Chun-Gen Piao

Yong Li

Roles

ABSTRACT

INTRODUCTION

RESULTS

FIG 1.

TAXONOMY

FIG 2.

TABLE 1.

FIG 3.

FIG 4.

FIG 5.

FIG 6.

FIG 7.

FIG 8.

FIG 9.

FIG 10.

FIG 11.

DISCUSSION

MATERIALS AND METHODS

Sample collection and isolation.

DNA extraction, sequencing, and phylogenetic analyses.

Morphology.

Data availability.

TABLE 2.

ACKNOWLEDGMENTS

Contributor Information

REFERENCES

Associated Data

Data Availability Statement

TABLE 2.

ACTIONS

PERMALINK

RESOURCES

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