Novel genus and species of Diaporthostomataceae (Diaporthales) in China

Ning Jiang; Han Xue; Yong Li

doi:10.3897/imafungus.16.145422

Research Article

Novel genus and species of Diaporthostomataceae (Diaporthales) in China

Ning Jiang^‡, Han Xue^‡, Yong Li^‡

‡ Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Beijing, China

Corresponding author: Yong Li ( lylx@caf.ac.cn )

Academic editor: Jadson Bezerra

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Jiang N, Xue H, Li Y (2025) Novel genus and species of Diaporthostomataceae (Diaporthales) in China. IMA Fungus 16: e145422. https://doi.org/10.3897/imafungus.16.145422

Abstract

Diaporthales is a significant fungal order comprising species that predominantly inhabit plant tissues, being pathogens, endophytes, and saprobes. Recent studies have uncovered extensive species diversity across various hosts, utilizing both morphological characteristics and molecular phylogenetic analyses. In this study, samples of leaf spots and branch cankers were collected from China, and fungal isolations were established. Species identification was conducted using a phylogenetic approach based on combined sequence data from the internal transcribed spacer (ITS) region, large subunit ribosomal DNA (LSU), the DNA-directed RNA polymerase II second largest subunit (rpb2), and translation elongation factor 1-alpha (tef1) genes, together with morphological observations. As a result, the novel genus Tiania is proposed, with three newly described species: T. chinensis, T. lithocarpicola, and T. quercicola. These species are validated by pairwise homoplasy index (PHI) analysis, ensuring robust support for their distinction. This study explores the rare family Diaporthostomataceae, providing the first descriptions of their anamorphic forms. By offering detailed morphological and molecular data, this research lays a foundation for future taxonomic and systematic studies of the Diaporthales.

Key words:

Ascomycota, biodiversity, phylogeny, systematics, taxonomy

Introduction

The order Diaporthales (Sordariomycetes, Ascomycota) comprises a diverse and ecologically significant group of fungi, including pathogens, endophytes, and saprotrophs, primarily associated with plant tissues, especially woody hosts (Sogonov et al. 2008; Walker et al. 2010; Voglmayr et al. 2012; Rossman et al. 2015; Zhang et al. 2023). Members of this order are distinguished by their unique teleomorphic features, including solitary or aggregated, immersed or erumpent, orange, brown, or black perithecial ascomata located in stromatic tissues or on substrates, often with a defined centrum; unitunicate asci with a prominent refractive ring; short to elongate, aseptate or septate, hyaline or pigmented ascospores (Castlebury et al. 2002; Rossman et al. 2007; Senanayake et al. 2017; Jaklitsch and Voglmayr 2020). The anamorphic states of this order are highly diverse, encompassing acervular, pycnidial, and synnematal conidiomata; usually phialidic and rarely annellidic conidiogenous cells; unicellular to septate, hyaline to pigmented, various shaped conidia (Castlebury et al. 2002; Senanayake et al. 2017; Fan et al. 2018; Jiang et al. 2021b).

In the traditional morphological classification system, families and genera within Diaporthales are primarily distinguished based on the morphology of stromata, including stromatic development and tissue types, the position of ascomata and perithecial necks, and the shape of ascospores. However, taxonomists have held differing views on the classification of diaporthalean fungi into families, e.g., Diaporthaceae and Melanosporaceae in Luttrell (1951), Diaporthaceae, Gnomoniaceae and Melanconidaceae in Chadefaud (1960), Diaporthaceae, Gnomoniaceae and Valsaceae in Wehmeyer (1975), Coryneaceae, Cytosporaceae, Gnomoniaceae and Melanconidaceae in Barr (1978).

Castlebury et al. (2002) initiated the molecular phylogenetic analysis of the order Diaporthales, focusing on the evaluation of its familial relationships. Their study included four families, Diaporthaceae, Gnomoniaceae, Melanconidaceae, and Valsaceae, along with two complexes: the Schizoparme complex and the Cryphonectria-Endothia complex, based on large subunit nuclear ribosomal DNA (LSU) sequences. Subsequently, additional families, including Cryphonectriaceae, Juglanconidaceae, Harknessiaceae, Lamproconiaceae, Macrohilaceae, Pseudoplagiostomaceae, Schizoparmaceae, Stilbosporaceae, and Sydowiellaceae, were incorporated into Diaporthales supported by morphological studies and phylogenetic analyses using the internal transcribed spacer (ITS) and LSU sequences (Gryzenhout et al. 2006; Rossman et al. 2007; Cheewangkoon et al. 2010; Crous et al. 2012b, 2015a; Voglmayr and Jaklitsch 2014; Alvarez et al. 2016; Norphanphoun et al. 2016; Voglmayr et al. 2017; Yang et al. 2017). Senanayake et al. (2017) further advanced the classification of Diaporthales by investigating its phylogenetic relationships using a combined dataset of ITS, LSU, DNA-directed RNA polymerase II second largest subunit (rpb2), and translation elongation factor 1-alpha (tef1) gene regions. Their work expanded the recognized families within this order to 21. Subsequently, additional families were incorporated into Diaporthales based on molecular phylogenetic frameworks based on ITS, LSU, rpb2, and tef1 sequence data, as well as morphological characteristics (Senanayake et al. 2018; Fan et al. 2018; Guterres et al. 2019; Jiang et al. 2020, 2021a).

The family Diaporthostomataceae was initially proposed by Fan et al. (2018) to include a single genus and species. When first described, it was considered phylogenetically sister to Diaporthosporellaceae (Fan et al. 2018). However, Jiang et al. (2020) sequenced the tef1 gene of Diaporthosporella cercidicola and determined that these two families are not sister clades, a finding later corroborated by Mu et al. (2024). Diaporthostomataceae is characterized by a typical diaporthalean teleomorph and can be distinguished from other families in Diaporthales by the absence of stromatic tissues and their fusoid, multiguttulate ascospores with an inconspicuous median septum (Fan et al. 2018).

Species of Diaporthales are well-known for causing plant diseases (Shuttleworth and Guest 2017; Jiang et al. 2019a; Udayanga et al. 2021; Zhu et al. 2024). For example, Cryphonectria parasitica is the causal agent of chestnut blight (Jiang et al. 2019b), while Gnomoniopsis fragariae, Paragnomonia fragariae, and Paraphomopsis obscurans are responsible for strawberry leaf spot (Udayanga et al. 2021). Additionally, many species of Cytospora are associated with tree canker diseases (Lawrence et al. 2018). Recent studies combining morphological and molecular phylogenetic analyses have uncovered numerous cryptic species within Diaporthales associated with tree disease symptoms (Liu et al. 2024; Mu et al. 2024).

During the extensive investigations conducted to collect forest pathogens in China, several diaporthalean taxa exhibiting branch canker and leaf spot symptoms were successfully isolated. The primary objective of this study was to accurately identify these newly collected diaporthalean species through morphological and molecular methods, while also elucidating their phylogenetic relationships within Diaporthales.

Materials and methods

Surveys and isolations

Samples, including leaf spots and branch cankers, were collected between 2019 and 2024 in China. Leaf samples were placed in self-sealing bags and transported to the laboratory for fungal isolation. Branch samples with visible fruiting bodies were cut into approximately 15 cm segments and preserved in paper sample bags for fungal isolation in the laboratory.

Leaves exhibiting spots were washed under tap water for 20 s, then dried on sterilized absorbent cotton. The leaves were surface sterilized by immersing them for 1 min in 75% ethanol, followed by 3 min in 1.25% sodium hypochlorite, and then for 1 min in 75% ethanol. After a 2-min rinse in distilled water, they were dried again on sterilized absorbent cotton. The leaves were then cut into 0.5 × 0.5 cm pieces using a sterile double-edged blade. Pieces with diseased and healthy tissues were transferred onto the surface of potato dextrose agar (PDA; 200 g potatoes, 20 g dextrose, and 20 g agar per liter) and incubated at 25 °C to obtain pure fungal cultures. Branches with fresh fruiting bodies were rinsed in tap water for 30 s to remove surface dust and then dried on sterilized absorbent cotton. Conidiomata and perithecia were carefully sectioned with a sterile blade to expose the spore masses, which were then transferred onto the surface of PDA plates using a sterile needle. The plates were incubated at 25 °C to establish fungal cultures. Type specimens were deposited in the herbarium of the Chinese Academy of Forestry (CAF, http://museum.caf.ac.cn/), and isolates were stored at the China Forestry Culture Collection Center (CFCC, https://cfcc.caf.ac.cn/).

Morphological analyses

The morphology of the new species identified in this study was analyzed based on fruiting bodies naturally formed on branches and PDA plates. Pseudostromata and conidiomata were sectioned using a double-edged blade, and their structures were examined under a Zeiss Discovery V8 stereomicroscope (Jena, Germany). Microscopic features, including asci, ascospores, conidiophores, conidiogenous cells, and conidia, were observed and photographed with an Olympus BX51 microscope (Tokyo, Japan). For spore measurements, 50 spores were randomly selected. The results are presented as maximum and minimum values (in parentheses), along with the range expressed as the mean ± standard deviation.

Colony characteristics were observed on three media types: potato dextrose agar (PDA), malt extract agar (MEA; 30 g malt extract, 5 g mycological peptone, 15 g agar per liter), and synthetic nutrient-deficient agar (SNA; 0.2 g glucose, 0.2 g sucrose, 1 g potassium dihydrogen phosphate, 1 g potassium nitrate, 0.25 g magnesium sulfate anhydrous, 0.5 g potassium chloride, 14 g agar per liter). Colony colors were documented following Rayner (1970).

Molecular analyses

Genomic DNA was extracted from colonies grown on PDA plates for 10 d using the Wizard® Genomic DNA Purification Kit (Promega, Madison, WI, USA), following the manufacturer’s protocol. To amplify the ITS, LSU, rpb2 and tef1 gene loci, the following primer pairs were used: ITS1/ITS4, LR0R/LR5, RPB2-5F/fRPB2-7cR, and EF1-728F/EF1-986R or EF1-728F/EF2, respectively (White et al. 1990; Glass and Donaldson 1995; Carbone and Kohn 1999; Liu et al. 1999; Rehner et al. 2001). Polymerase chain reaction (PCR) conditions included an initial denaturation at 94 °C for 5 min, followed by 35 cycles of 30 s at 94 °C, 50 s at 48 °C (for ITS and LSU) or 54 °C (for tef1) or 55 °C (for rpb2), and 1 min at 72 °C, with a final elongation step of 7 min at 72 °C. Amplicons were sequenced in both directions by Ruibo Xingke Biotechnology Company Limited (Beijing, China).

Sequences were assembled using Seqman v. 7.1.0 (DNASTAR Inc., Madison, WI, USA) and deposited in GenBank, and reference sequences were selected from recent studies on Diaporthales (Table 1). Sequence alignments of the four loci (ITS, LSU, rpb2, and tef1) were performed in MAFFT v. 7 (Katoh and Standley 2023) and manually edited in MEGA v. 7.0.21.

Table 1.

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XLSX

Details of isolates included in the molecular study.

Species	Strain	GenBank accession numbers				References
Species	Strain	ITS	LSU	rpb2	tef1	References
Apiognomonia errabunda	AR 2813	DQ313525	NA	DQ862014	DQ313565	Sogonov et al. 2007
Apiosporopsis carpinea	CBS 771.79	NA	AF277130	NA	NA	Zhang and Blackwell 2001
Apoharknessia insueta	CBS 111377*	JQ706083	AY720814	NA	MN271820	Crous et al. 2012b; Jiang et al. 2020
Apoharknessia insueta	CBS 114575	MN172402	MN172370	NA	MN271821	Jiang et al. 2020
Asterosporium asterospermum	MFLU 15-3555	NA	MF190062	NA	NA	Senanayake et al. 2017
Auratiopycnidiella tristaniopsis	CBS 132180*	JQ685516	JQ685522	NA	MN271825	Crous et al. 2012a; Jiang et al. 2020
Auratiopycnidiella tristaniopsis	CPC 16371	MN172405	MN172374	NA	MN271826	Jiang et al. 2020
Aurifilum marmelostoma	CBS 124928*	FJ890495	MH874934	MN271788	MN271827	Begoude et al. 2010; Jiang et al. 2020
Chrysofolia barringtoniae	TBRC 5647*	KU948046	KU948045	NA	NA	Suwannarach et al. 2016
Chrysofolia colombiana	CBS 139909*	KR476738	KR476771	NA	MN271829	Crous et al. 2015b; Jiang et al. 2020
Coniella africana	CBS 114133*	AY339344	AY339293	KX833421	KX833600	Alvarez et al. 2016
Coniella eucalyptorum	CBS 112640*	AY339338	AY339290	KX833452	KX833637	Alvarez et al. 2016
Coniella fusiformis	CBS 141596*	KX833576	KX833397	KX833481	KX833674	Alvarez et al. 2016
Coryneum gigasporum	CFCC 52319*	MH683565	MH683557	MH685729	MH685737	Jiang et al. 2018b
Coryneum umbonatum	D201	MH674329	MH674329	MH674333	MH674337	Jiang et al. 2018b
Cryphonectria citrine	CBS 109758*	MN172407	EU255074	EU219342	MN271843	Sogonov et al. 2008; Jiang et al. 2020
Cryphonectria decipens	CBS 129351	EU442657	MN172385	MN271796	MN271844	Jiang et al. 2020
Cytospora chrysosperma	CFCC 89982	KP281261	KP310805	KU710952	KP310848	Yang et al. 2015
Cytospora elaeagni	CFCC 89633	KF765677	KF765693	KU710956	KU710919	Yang et al. 2015
Cytospora viridistroma	CBS 202.36*	MN172408	MN172388	NA	MN271853	Jiang et al. 2020
Dendrostoma castaneae	CFCC 52745*	MH542488	MH542644	MH545395	MH545437	Jiang et al. 2019a
Dendrostoma chinense	CFCC 52755*	MH542500	MH542648	MH545407	MH545449	Jiang et al. 2019a
Diaporthe eres	LC3198	KP267873	KY011845	NA	KP267947	Gao et al. 2016
Diaporthe hongkongensis	LC0784	KC153104	KY011876	NA	KC153095	Gao et al. 2016
Diaporthosporella cercidicola	CFCC 51994*	KY852492	KY852515	NA	MN271855	Yang et al. 2017; Jiang et al. 2020
Diaporthosporella macarangae	NCYU 19-0359*	MW114354	MW114415	NA	NA	Tennakoon et al. 2021
Diaporthosporella macarangae	NCYU 19-0363	MW114355	MW114416	NA	NA	Tennakoon et al. 2021
Diaporthostoma machili	CFCC 52100*	MG682080	MG682020	MG682040	MG682060	Fan et al. 2018
Diaporthostoma machili	CFCC 52101	MG682081	MG682021	MG682041	MG682061	Fan et al. 2018
Disculoides eucalyptorum	CBS 132184	JQ685518	JQ685524	MH545414	MH545456	Crous et al. 2012a
Dwiroopa lythri	CBS 109755*	MN172410	MN172389	MN271801	MN271859	Jiang et al. 2020
Dwiroopa punicae	CBS 143163*	MK510676	MK510686	MK510692	NA	Xavier et al. 2019
Foliocryphia eucalypti	CBS 124779*	GQ303276	GQ303307	MN271802	MN271861	Cheewangkoon et al. 2009; Jiang et al. 2020
Foliocryphia eucalyptorum	CBS 142536*	KY979772	KY979827	MN271803	MN271862	Crous et al. 2013; Jiang et al. 2020
Gnomonia gnomon	CBS 199.53	DQ491518	AF408361	EU219295	NA	Castlebury et al. 2002; Sogonov et al. 2007
Harknessia australiensis	CBS 132119*	JQ706085	JQ706211	NA	MN271863	Crous et al. 2012b; Jiang et al. 2020
Harknessia capensis	CBS 111829*	AY720719	AY720816	NA	MN271864	Crous et al. 2012b; Jiang et al. 2020
Harknessia gibbosa	CBS 120033*	EF110615	EF110615	NA	MN271868	Crous et al. 2012b; Jiang et al. 2020
Juglanconis juglandina	CBS 121083	KY427148	KY427148	KY427198	KY427217	Voglmayr et al. 2017
Juglanconis oblonga	MAFF 410216	KY427153	KY427153	KY427203	KY427222	Voglmayr et al. 2017
Juglanconis pterocaryae	MAFF 410079	KY427155	KY427155	KY427205	KY427224	Voglmayr et al. 2017
Lamproconium desmazieri	MFLUCC 15-0870	KX430134	KX430135	MF377605	MF377591	Norphanphoun et al. 2016; Senanayake et al. 2017
Lamproconium desmazieri	MFLUCC 15-0872	KX430138	KX430139	MF377606	MF377593	Norphanphoun et al. 2016; Senanayake et al. 2017
Luteocirrhus shearii	CBS 130776*	KC197021	KC197019	MN271807	MN271890	Crane and Burgess 2013; Jiang et al. 2020
Macrohilum eucalypti	CPC 10945	DQ195781	DQ195793	MN271809	NA	Crous et al. 2006; Jiang et al. 2020
Macrohilum eucalypti	CPC 19421	KR873244	KR873275	MN271810	NA	Crous et al. 2006; Jiang et al. 2020
Mastigosporella anisophylleae	CBS 136421*	KF779492	KF777221	NA	MN271892	Crous et al. 2013; Jiang et al. 2020
Mastigosporella pigmentata	COAD 2370*	MG587929	MG587928	NA	NA	Crous et al. 2018
Melanconiella ellisii	BPI 878343	JQ926271	JQ926271	JQ926339	JQ926406	Voglmayr et al. 2012
Melanconiella spodiaea	MSH	JQ926298	JQ926298	JQ926364	JQ926431	Voglmayr et al. 2012
Melanconis betulae	CFCC 50471	KT732952	KT732971	KT732984	KT733001	Fan et al. 2016
Melanconis itoana	CFCC 50474	KT732955	KT732974	KT732987	KT733004	Fan et al. 2016
Melanconis stilbostoma	CFCC 50475	KT732956	KT732975	KT732988	KT733005	Fan et al. 2016
Nakataea oryzae	CBS 243.76	KM484861	DQ341498	NA	NA	Klaubauf et al. 2014
Neocryphonectria chinensis	CFCC 53025*	MN172414	MN172397	MN271812	MN271893	Jiang et al. 2020
Neopseudomelanconis castaneae	CFCC 52787*	MH469162	MH469164	NA	NA	Jiang et al. 2018a
Phaeoappendicospora thailandensis	MFLU 12-2131	MF190157	MF190102	NA	NA	Senanayake et al. 2017
Prosopidicola albizziae	CPC 27478	KX228274	KX228325	NA	NA	Crous et al. 2013
Prosopidicola Mexicana	CBS 113529	AY720709	NA	NA	NA	Lennox et al. 2004
Pseudomelanconis caryae	CFCC 52110*	MG682082	MG682022	MG682042	MG682062	Fan et al. 2018
Pseudoplagiostoma corymbiae	CPC 14161	GU973510	GU973604	NA	GU973540	Cheewangkoon et al. 2010
Pseudoplagiostoma oldie	CBS 115722	GU973535	GU973610	NA	GU973565	Cheewangkoon et al. 2010
Pseudoplagiostoma variabile	CBS 113067	GU973536	GU973611	NA	GU973566	Cheewangkoon et al. 2010
Pyricularia grisea	Ina168	NA	AB026819	NA	NA	Sone et al. 2000
Pyrispora castaneae	CFCC 54349*	MW208108	MW208105	MW218535	MW227340	Jiang et al. 2021a
Pyrispora castaneae	CFCC 54350	MW208109	MW208106	MW218536	MW227341	Jiang et al. 2021a
Sillia karstenii	MFLU 16-2864	KY523482	KY523500	KY501636	NA	Senanayake et al. 2017
Sirococcus tsugae	CBS 119626	EU199203	EU199136	EU199159	EF512534	Mejía et al. 2008
Stegonsporium acerophilum	CBS 117025	EU039982	EU039993	KF570173	EU040027	Voglmayr and Jaklitsch 2008, 2014
Stilbospora longicornuta	CBS 122529*	KF570164	KF570164	KF570194	KF570232	Voglmayr and Jaklitsch 2014
Synnemasporella aculeans	CFCC 52094	MG682086	MG682026	MG682046	MG682066	Fan et al. 2018
Synnemasporella toxicodendri	CFCC 52097*	MG682089	MG682029	MG682049	MG682069	Fan et al. 2018
*Tiania chinensis*	CFCC 59134*	PQ781258	PQ781255	PQ786769	PQ786772	In this study
*Tiania chinensis*	CFCC 59135	PQ781259	PQ781256	PQ786770	PQ786773	In this study
*Tiania chinensis*	CFCC 71190*	PQ781260	PQ781257	PQ786771	PQ786774	In this study
*Tiania lithocarpicola*	CFCC 55331*	OK339758	OK339787	OK358595	OK358599	In this study
*Tiania lithocarpicola*	CFCC 55882	OK339759	OK339788	OK358596	OK358600	In this study
*Tiania quercicola*	CFCC 54435*	OK339756	OK339785	OK358593	OK358597	In this study
*Tiania quercicola*	CFCC 55885	OK339757	OK339786	OK358594	OK358598	In this study
Tubakia iowensis	CBS 129012*	MG591879	MG591971	NA	MG592064	Braun et al. 2018
Tubakia seoraksanensis	CBS 127490*	MG591907	KP260499	NA	MG592094	Braun et al. 2018

Phylogenetic analyses were conducted on the combined dataset of the four loci using Maximum Likelihood (ML) and Bayesian Inference (BI). ML analysis was performed with the GTR substitution model and 1000 bootstrap replicates via the CIPRES Science Gateway portal (https://www.phylo.org/; Miller et al. 2010) using RAxML-HPC BlackBox v. 8.2.10 (Stamatakis 2008). BI analysis applied partition-specific evolutionary models selected by MrModeltest v. 2.3 using the Akaike Information Criterion (AIC). Markov Chain Monte Carlo (MCMC) simulations in MrBayes v. 3.1.2 (Ronquist and Huelsenbeck 2003) were run for 10 million generations with two chains initiated from random trees. Convergence was confirmed by an average standard deviation of split frequencies below 0.01, and trees were sampled every 1000 generations. The first 25% of sampled trees were discarded as burn-in, and posterior probabilities (BPP) were calculated from the remaining trees. Bootstrap (BS) support in ML analyses was assessed with 1000 replicates, and phylogenetic trees were visualized in FigTree v. 1.4.4 (Rambaut 2018).

The pairwise homoplasy index (PHI, Φ_w) test was conducted using the SplitsTree App to evaluate recombination among closely related phylogenetic species (Huson and Bryant 2024). The analysis used a concatenated four-locus dataset (ITS, LSU, rpb2, and tef1), and Φ_w-statistic below 0.05 (p-value < 0.05) indicated no significant evidence of recombination. Relationships among closely related taxa were illustrated with split graphs generated using the Log-Det transformation and split decomposition methods, providing a clear visualization of phylogenetic associations.

Results

Phylogenetic analyses

The combined dataset of ITS, LSU, rpb2, and tef1 comprised 81 strains, with Nakataea oryzae (CBS 243.76) and Pyricularia grisea (Ina168) designated as the outgroup taxa. The final alignment consisted of 3,229 characters (ITS: 660; LSU: 789; rpb2: 1,065; tef1: 715), including gaps. The ML optimization likelihood value for the best RAxML tree was -53,777.19, with the matrix containing 2,052 distinct alignment patterns and 38.12% undetermined characters or gaps. The estimated base frequencies were A = 0.238237, C = 0.263656, G = 0.272107, and T = 0.226000. Substitution rates were calculated as follows: AC = 1.555398, AG = 2.732713, AT = 1.808604, CG = 1.204566, CT = 6.521498, and GT = 1.000000. The gamma distribution shape parameter (α) was 0.269658. For Bayesian inference (BI) analysis, the most appropriate models for each locus were confirmed using MrModeltest. The selected models were SYM+I+G4 for ITS, SYM+R3 for LSU, TN+F+I+G4 for rpb2, and TIM2+F+I+G4 for tef1. The Bayesian analysis results aligned with the ML tree topology. ML bootstrap support values (BS) ≥ 50% and Bayesian posterior probabilities (BPP) ≥ 0.90 are indicated on the branches in Fig. 1. The phylogram, constructed using four gene markers, delineated 31 distinct lineages corresponding to 31 families within Diaporthales. Phylogenetic analysis revealed that Diaporthostomataceae forms a sister clade to Tubakiaceae, together constituting a well-supported clade with Pseudomelanconidaceae and Melanconiellaceae. Notably, the new strains in this study form a robustly supported clade that is sister to Diaporthostoma, representing a newly discovered genus which we propose to name Tiania.

Figure 1.

Phylogram of Diaporthales resulting from a maximum likelihood analysis based on the ITS, LSU, rpb2 and tef1 gene sequence. Numbers above the branches indicate ML bootstrap values (left, ML BS ≥ 50%) and Bayesian posterior probabilities (right, BPP ≥ 0.90). The tree is rooted with Nakataea oryzae (CBS 243.76) and Pyricularia grisea (Ina168). Isolates from the present study are marked in blue.

PHI Analysis

In the genus Tiania, seven isolates were grouped into three distinct clades with high support values (Fig. 1). To validate the species delineation, PHI analysis was performed. The analysis revealed no significant evidence of genetic recombination among these species (p = 0.1139; Fig. 2).

Figure 2.

The split graphs of a PHI test result of Tiania species using the LogDet transformation and split decomposition options based on the ITS, LSU, rpb2 and tef1 gene sequence (p = 0.1139).

Taxonomy

Tiania Ning Jiang, gen. nov.

MycoBank No: 857019

Etymology.

In honor of Chinese taxonomist Prof. Dr. Chengming Tian for his contributions for forest pathogens.

Type species.

Tiania chinensis Ning Jiang.

Description.

Pseudostromata immersed to semi-immersed in the bark, scattered, conical, with perithecia arranged irregular. Ectostromatic disc grey to brown, circular to ovoid. Ostioles brown to black. Perithecia flask-shaped to spherical. Asci hyaline, with chitinoid, refractive ring, clavate to elongate-obovoid, 8-spored. Ascospores biseriate, cylindrical to allantoid, thin-walled, hyaline, 0–1 septate. Conidiomata acervular in tree branches and sporodochial in culture, aggregated, immersed to semi-immersed, pulvinate. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, phialidic. Conidia aseptate, hyaline, smooth, multi-guttulate, fusoid, cylindrical to allantoid, constricted at the middle or not.

Notes.

The newly proposed genus Tiania is phylogenetically closely related to Diaporthostoma within the family Diaporthostomaceae (Fig. 1). Morphologically, Tiania can be distinguished from Diaporthostoma by its formation of stromatic tissue and its cylindrical to allantoid ascospores (Fan et al. 2018).

Tiania chinensis Ning Jiang, sp. nov.

MycoBank No: 857020

Figs 3, 4

Etymology.

Named after the collection country of the type specimen, China.

Diagnosis.

Distinct from its phylogenetically related species T. lithocarpicola by longer conidia.

Typus.

CHINA • Xizang Autonomous Region, Rikaze City, Jilong County, Jilong Town, Rema Village, on diseased branches of Quercus semecarpifolia, 20 August 2022, Ning Jiang, Min Liu & Peng Jin (holotype CAF800088; ex-holotype culture CFCC 59134); • Xizang Autonomous Region, Linzhi City, Gongbujiangda County, Gongbujiangda Town, on diseased branches of Quercus spinosa, 7 July 2024, Ning Jiang, Jiangrong Li, Jieting Li & Liangna Guo (paratype CAF800141; ex-paratype culture CFCC 71190).

Figure 3.

Tiania chinensis sp. nov. from Quercus spinosa (CAF800141). A, B Habit of ascostromata on branch; C, D transverse section through ascostroma; E longitudinal section through ascostroma; F asci; G ascospores. Scale bars: 500 μm (B–E); 20 μm (F); 10 μm (G).

Description.

Pseudostromata immersed to semi-immersed in the bark, scattered, conical, 630–1240 μm diam, 330–480 μm high, with 5–10 perithecia arranged irregularly. Ectostromatic disc grey to brown, circular to ovoid, 300–470 μm diam. Ostioles brown to black, 90–150 μm diam. Perithecia flask-shaped to spherical, 490–620 μm diam. Asci hyaline, with chitinoid, refractive ring, clavate to elongate-obovoid, (38.5–)41.5–47.5(–52) × (7–)8–9.5(–10) μm, 8-spored. Ascospores biseriate, cylindrical to allantoid, thin-walled, hyaline, 0–1 septate, (11–)11.5–14.5(–15) × (2.5–)3–3.5 (n = 50) μm, L/W ratio = 3.7–5. Conidiomata acervular, aggregated, immersed to semi-immersed in the bark, pulvinate, dark brown to black, 250–600 μm high, 350–1000 μm diam. Conidiophores indistinct, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical, phialidic, 9.2–14.5 × 2–2.9 μm. Conidia aseptate, hyaline, smooth, multi-guttulate, cylindrical to allantoid, straight or slightly curved, (7.5–)8–9(–10) × (2–)2.5–3(–3.5) μm (n = 50), L/W = 2.7–3.6.

Figure 4.

Tiania chinensis sp. nov. from Quercus semecarpifolia (CAF800088). A, C Habit of conidiomata on branch; B transverse section through conidioma; D longitudinal section through conidioma; E conidiogenous cells with attached conidia; F, G conidia. Scale bars: 500 μm (A); 200 μm (B, C); 100 μm (D); 10 μm (E–G).

Culture characteristics.

Colonies on PDA flat, spreading, with abundant flocculent aerial mycelium and even margin, white to sky grey, reaching 90 mm diam after 2 wk at 25 °C. Colonies on MEA flat, spreading, lavender grey to grey olivaceous, reaching 90 mm diam after 2 wk at 25 °C. Colonies on SNA flat, spreading, with sparse flocculent aerial mycelium and feathery margin.

Additional material examined.

CHINA • Xizang Autonomous Region, Rikaze City, Jilong County, Jilong Town, Rema Village, from cankered barks of Quercus semecarpifolia, 21 August 2022, Ning Jiang, Min Liu & Peng Jin (living culture CFCC 59135).

Distribution.

China, Xizang Autonomous Region.

Ecology.

Associated with branch canker disease with Quercus semecarpifolia and Q. spinosa.

Notes.

Three isolates obtained from diseased branches of Quercus semecarpifolia and Q. spinosa formed a distinct clade, separate from Tiania lithocarpicola and T. quercicola, and are identified as T. chinensis sp. nov. This species can be distinguished from T. lithocarpicola by its longer conidia (8–9 × 2.5–3 μm in T. chinensis vs. 5–6.5 × 2–2.5 μm in T. lithocarpicola) and from T. quercicola by its cylindrical to allantoid conidia.

Tiania lithocarpicola Ning Jiang, sp. nov.

MycoBank No: 857021

Fig. 5

Etymology.

Named after the host genus Lithocarpus and “-cola” = “inhabiting”.

Diagnosis.

Distinct from its sister species T. chinensis by shorter conidia.

Typus.

CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Lithocarpus elaeagnifolius, 12 November 2018, Yong Li (holotype CAF800042; ex-holotype culture CFCC 55331).

Description.

Conidiomata in culture sporodochial, aggregated, erumpent, pulvinate, light brown, 150–650 μm diam., exuding light brown conidial masses. Conidiophores hyaline, smooth, cylindrical, branched. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, attenuate towards the apex, phialidic, 6.5–22.5 × 1.5–2.5 μm. Conidia aseptate, hyaline, smooth, multi-guttulate, fusoid to ellipsoid, straight or slightly curved, (4.5–)5–6.5(–7.5) × 2–2.5(–3) μm (n = 50), L/W = 1.9–3.5.

Figure 5.

Tiania lithocarpicola sp. nov. from Lithocarpus elaeagnifolius (CAF800042). A Colonies on PDA, MEA and SNA after 10 d at 25 °C; B conidiomata formed on PDA; C conidiogenous cells; D–G conidia. Scale bars: 300 μm (B); 10 μm (C–G).

Culture characteristics.

Colonies on PDA flat, spreading, with moderate flocculent aerial mycelium and even margin, forming concentric rings, white to straw, reaching 90 mm diam after 2 wk at 25 °C. Colonies on MEA flat, spreading, with moderate flocculent aerial mycelium and undulating margin, forming salmon irregular center area and smoke grey to ochreous outer area, reaching 70 mm diam after 2 wk at 25 °C. Colonies on SNA flat, dense, white, slowly growing.

Additional material examined.

CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, from leaf spots of Lithocarpus elaeagnifolius, 12 November 2018, Yong Li (living culture CFCC 55882).

Distribution.

China, Hainan Province.

Ecology.

Associated with leaf spot disease with Lithocarpus elaeagnifolius.

Notes.

Tiania lithocarpicola is phylogenetically closely related to T. chinensis but can be distinguished by its shorter conidia (5–6.5 × 2–2.5 μm in T. lithocarpicola vs. 8–9 × 2.5–3 μm in T. chinensis).

Tiania quercicola Ning Jiang, sp. nov.

MycoBank No: 857020

Fig. 6

Etymology.

Named after the host genus Quercus and “-cola” = “inhabiting”.

Diagnosis.

Distinct from T. chinensis and T. lithocarpicola by conidia that are constricted at the middle.

Typus.

CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, on diseased leaves of Quercus macrocalyx, 30 March 2019, Yong Li (holotype CAF800035; ex-holotype culture CFCC 54435).

Description.

Conidiomata in culture sporodochial, aggregated, erumpent, pulvinate, light orange, 150–700 μm diam., exuding light orange conidial masses. Conidiophores hyaline, smooth, cylindrical, branched, usually reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, cylindrical to ampulliform, attenuate towards the apex, phialidic, 10.5–21.5 × 1–2.5 μm. Conidia aseptate, hyaline, smooth, multi-guttulate, cylindrical, constricted at the middle, straight or slightly curved, base truncate, 5.5–7(–8) × 2–2.5 μm (n = 50), L/W = 1.6–2.7.

Figure 6.

Tiania quercicola sp. nov. from Quercus macrocalyx (CAF800035). A Colonies on PDA, MEA and SNA after 10 d at 25 °C; B conidiomata formed on PDA; C conidiogenous cells; D–G conidia. Scale bars: 300 μm (B); 10 μm (C–G).

Culture characteristics.

Colonies on PDA flat, spreading, with abundant flocculent aerial mycelium and even margin, initially white, becoming umber after 1 wk, reaching 90 mm diam after 2 wk at 25 °C. Colonies on MEA flat, spreading, with abundant flocculent aerial mycelium and undulating margin, white to smoke grey, reaching 90 mm diam after 2 wk at 25 °C. Colonies on SNA flat, spreading, with sparse flocculent aerial mycelium and feathery margin, white, reaching 90 mm diam after 3 wk at 25 °C.

Additional material examined.

CHINA • Hainan Province, Changjiang Li Autonomous County, Bawangling National Forest Park, from leaf spots of Quercus macrocalyx, 30 March 2019, Yong Li (living culture CFCC 55885).

Distribution.

China, Hainan Province.

Ecology.

Associated with leaf spot disease with Quercus macrocalyx.

Notes.

Tiania quercicola, isolated from Quercus macrocalyx, is phylogenetically closely related to T. chinensis from Quercus semecarpifolia and Q. spinosa, and T. lithocarpicola from Lithocarpus elaeagnifolius (Fig. 1). However, T. quercicola can be distinguished from these two species by its conidia, which are constricted at the middle.

Key to genera and species of Diaporthostomataceae

1	Stromata well-developed	*Diaporthostoma machili*
–	Stromata absent	2
2	Conidia constricted at the middle	*Tiania quercicola*
–	Conidia not constricted at the middle	3
3	Conidia cylindrical to allantoid, 8–9 × 2.5–3 μm	*T. chinensis*
–	Conidia fusoid to ellipsoid, 5–6.5 × 2–2.5 μm	*T. lithocarpicola*

Discussion

Diaporthales is a well-studied order within Ascomycota, both morphologically and phylogenetically (Barr 1978; Voglmayr and Jaklitsch 2014; Alvarez et al. 2016; Senanayake et al. 2017, 2018; Voglmayr et al. 2017). Since the first molecular study on this order, which was based on the single locus LSU (Castlebury 2002), a more robust classification system has been established primarily based on the morphology of naturally formed fruiting bodies on hosts and the phylogeny of combined loci, including ITS, LSU, rpb2, and tef1 (Senanayake et al. 2017; Fan et al. 2018; Jiang et al. 2020). In this study, we introduce a new genus, Tiania, in the family Diaporthostomataceae, based on newly collected samples exhibiting typical diaporthalean characteristics. Additionally, we propose three new species within this genus, named T. chinensis, T. lithocarpicola, and T. quercicola.

The family Diaporthostomataceae was established with a single genus and species, Diaporthostoma machili (Fan et al. 2018). It is characterized by a teleomorph resembling Diaporthe and can be distinguished from its phylogenetic sister clade, Diaporthosporellaceae, by having discrete perithecia and fusoid, straight to curved ascospores with a median septum (Yang et al. 2017; Fan et al. 2018). Unfortunately, no anamorph has been reported for this fungus to date.

As the second genus in the Diaporthostomataceae family, Tiania is characterized by both teleomorphic and anamorphic states. This genus features pseudostromata resembling those of Cytospora, but it is distinguished by its aseptate or septate ascospores, differing from Cytospora (Fan et al. 2020). In its anamorphic state, Tiania exhibits conidia of various shapes. For instance, T. chinensis produces cylindrical to allantoid conidia, similar to those of Cytospora (Fan et al. 2020); T. lithocarpicola forms fusoid to ellipsoid conidia, resembling those of Diaporthe (Dissanayake et al. 2024); and T. quercicola develops cylindrical conidia constricted in the middle, akin to those of Micromelanconis (Jiang et al. 2021c). Additionally, its dark acervular conidiomata bear a striking resemblance to those of Coryneum (Senanayake et al. 2017). Thus, the new genus Tiania exhibits morphological similarities to several taxa, encompassing features from different families within Diaporthales.

The taxonomy of Diaporthales has undergone significant advancements over the past decade, uncovering numerous fascinating taxa (Voglmayr and Jaklitsch 2014; Alvarez et al. 2016; Senanayake et al. 2017, 2018; Voglmayr et al. 2017; Fan et al. 2018; Jiang et al. 2020, 2021b). At the family level, several families, including Cytosporaceae, Diaporthaceae, and Gnomoniaceae, are large groups containing a high number of species (Senanayake et al. 2018). In contrast, some families, such as Dwiroopaceae, Mastigosporellaceae, and Diaporthosporellaceae, are relatively less studied and comprise only a small number of species (Fan et al. 2018; Xavier et al. 2019; Jiang et al. 2020). The family Diaporthostomataceae stands out as a particularly rare and understudied fungal group. This rarity may be attributed to their specific habitat requirements, as they thrive in environments undisturbed by adverse conditions. Currently, all known species of Diaporthostomataceae have been discovered exclusively in primeval forests or natural reserves.

Species of Diaporthostomataceae may act as pathogens on their original hosts, as suggested by symptoms observed during investigations. However, due to their rarity, there is currently no need for active management of the disease, even if they are confirmed as pathogens. Nonetheless, comprehensive pathogenicity tests are required in the future to confirm their role and impact as pathogens.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Adherence to national and international regulations

All the fungal strains used in this study have been legally obtained, respecting the Convention on Biological Diversity (Rio Convention).

Funding

This study was supported by Fundamental Research Funds of CAF (CAFYBB2023PA002), and the National Microbial Resource Center of the Ministry of Science and Technology of the People’s Republic of China (NMRC-2024-7).

Author contributions

Conceptualization (NJ, HX and YL), Methodology (NJ and HX), Software (NJ), Validation (NJ), Formal analysis (NJ), Investigation (NJ and YL), Resources (NJ, HX and YL), Data Curation (NJ, HX and YL), Writing – Original draft (NJ), Writing – Review and Editing (NJ, HX and YL), Visualization (NJ), Supervision (YL), Project administration (NJ, HX and YL), Funding Acquisition (NJ, HX and YL).

Author ORCIDs

Ning Jiang https://orcid.org/0000-0002-9656-8500

Han Xue https://orcid.org/0000-0003-0414-6237

Yong Li https://orcid.org/0000-0002-4406-1329

Data availability

All of the data that support the findings of this study are available in the main text.

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﻿Abstract

Key words:

﻿Introduction

﻿Materials and methods

﻿Surveys and isolations

﻿Morphological analyses

﻿Molecular analyses

﻿Results

﻿Phylogenetic analyses

﻿PHI Analysis

﻿Taxonomy

﻿ Tiania Ning Jiang, gen. nov.

Etymology.

Type species.

Description.

Notes.

﻿ Tiania chinensis Ning Jiang, sp. nov.

Etymology.

Diagnosis.

Typus.

Description.

Culture characteristics.

Additional material examined.

Distribution.

Ecology.

Notes.

﻿ Tiania lithocarpicola Ning Jiang, sp. nov.

Etymology.

Diagnosis.

Typus.

Description.

Culture characteristics.

Additional material examined.

Distribution.

Ecology.

Notes.

﻿ Tiania quercicola Ning Jiang, sp. nov.

Etymology.

Diagnosis.

Typus.

Description.

Culture characteristics.

Additional material examined.

Distribution.

Ecology.

Notes.

﻿Key to genera and species of Diaporthostomataceae

﻿Discussion

﻿Additional information

Conflict of interest

Ethical statement

Adherence to national and international regulations

Funding

Author contributions

Author ORCIDs

Data availability

﻿References

Abstract

Introduction

Materials and methods

Surveys and isolations

Morphological analyses

Molecular analyses

Results

Phylogenetic analyses

PHI Analysis

Taxonomy

Tiania Ning Jiang, gen. nov.

Tiania chinensis Ning Jiang, sp. nov.

Tiania lithocarpicola Ning Jiang, sp. nov.

Tiania quercicola Ning Jiang, sp. nov.

Key to genera and species of Diaporthostomataceae

Discussion

Additional information

References