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Research Article
Morpho-phylogenetic evidence reveals four novel species of Coniella (Diaporthales, Schizoparmaceae) from southern China
expand article infoDuhua Li‡§, Zixu Dong‡, Qiyun Liu‡, Yaling Wang‡, Zhaoxue Zhang‡, Xiuguo Zhang‡, Jiwen Xia§
‡ Shandong Agricultural University, Taian, China
§ Linyi University, Linyi, China

Corresponding author: Jiwen Xia ( xiajiwen@lyu.edu.cn )

Academic editor: Danushka Sandaruwan Tennakoon
© 2025 Duhua Li, Zixu Dong, Qiyun Liu, Yaling Wang, Zhaoxue Zhang, Xiuguo Zhang, Jiwen Xia.
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: Li D, Dong Z, Liu Q, Wang Y, Zhang Z, Zhang X, Xia J (2025) Morpho-phylogenetic evidence reveals four novel species of Coniella (Diaporthales, Schizoparmaceae) from southern China. MycoKeys 116: 1-23. https://doi.org/10.3897/mycokeys.116.145857
Open Access

Abstract

Coniella species are distributed worldwide and have been reported as plant pathogens, endophytes, or saprobes. In our ongoing survey of terrestrial plant fungi in southern China, we obtained Coniella isolates from diseased plant leaf tissues in Fujian, Hainan, and Yunnan provinces. Maximum likelihood and Bayesian inference based on four loci (ITS, LSU, rpb2, and tef1-α) were used to clarify the taxonomic placement of the species. We confirmed that they represent four new species, namely Coniella diaoluoshanensis, C. dongshanlingensis, C. grossedentatae, and C. veri based on both morphology and phylogeny support. The new species are compared with other Coniella species, comprehensive descriptions and micrographs are provided.

Key words:

Morphology, multigene phylogeny, new taxa, taxonomy

Introduction

Coniella was formally introduced by Von Höhnel (1918) with C. pulchella (= C. fragariae (Oudem.) B. Sutton) as the type species (Von Höhnel 1918; Sutton 1977; Crous et al. 2014a). Samuels et al. (1993) initially recognized the uniqueness of Schizoparme and its relationship to Coniella and Pilidiella, these were initially placed in the Melanconidaceae. Both Castlebury et al. (2002) and Van Niekerk et al. (2004) revealed that these species within the Diaporthales, which they collectively designated as the Schizoparme complex. Rossman et al. (2007) introduced a new family, Schizoparmaceae, which comprises the distinctive teleomorph genus Schizoparme, its asexual state Pilidiella, and the closely related anamorph genus Coniella. These genera are cosmopolitan fungal pathogens associated with foliar, fruit, stem, and root diseases on a wide variety of hosts, including some economically important hosts (Van Niekerk et al. 2004; Alvarez et al. 2016). They occur as parasites on unrelated dicotyledonous hosts (Samuels et al. 1993) or sometimes as secondary invaders of injured plant tissues (Ferreira et al. 1997).

Coniella has undergone comprehensive morpho-molecular studies and experienced several taxonomic adjustments over the years. Petrak and Sydow (1927) classified Coniella into two subgenera: Euconiella (dark conidia), typified by C. pulchella, and Pseudoconiella (hyaline to pale conidia), typified by C. granati. Von Arx (1973, 1981) classified Coniella and Pilidiella as distinct genera, with Coniella characterized by dark brown conidia and Pilidiella by hyaline conidia that darken to a pale brown when mature. Nonetheless, Sutton (1980) and Nag Raj (1993) disregarded conidial pigmentation as a defining trait and still opted to employ the earlier name Coniella. Samuels et al. (1993) stated Schizoparme as the sexual morph and positioned it in Melanconidaceae. Castlebury et al. (2002) classified Pilidiella and Coniella as members of the Schizoparme complex. Van Niekerk et al. (2004) demonstrated that these taxa form a distinct evolutionary lineage within the Diaporthales based on ITS, LSU, and tef1-α sequences. Subsequently, Rossman et al. (2007) established a new family, Schizoparmaceae, including the above three genera, viz. Coniella, Pilidiella, and Schizoparme. Alvarez et al. (2016) demonstrated that Coniella, Pilidiella, and Schizoparme formed a monophyletic clade in Schizoparmaceae and suggested adopting Coniella (the older asexual typified name) instead of Pilidiella and Schizoparme, in accordance with Article 59.1 of the International Code of Nomenclature for Algae, Fungi, and Plants (ICN, Melbourne Code; McNeill et al. 2012). Additionally, due to the many numbers of species and the similarity in morphological characteristics, they suggested that the identification of new species within Coniella must be based on a combination of DNA sequence data and morphological characteristics. Chethana et al. (2017) used a combination of morphological analysis and multigene phylogeny with the genealogical concordance phylogenetic species recognition (GCPSR) method to delineate species boundaries. Hyde et al. (2020) and Tennakoon et al. (2021) conducted the recent phylogenetic analyses for Coniella species within the Schizoparmaceae. Currently, there are 66 accepted Coniella species (Index Fungorum: https://indexfungorum.org; MycoBank: http://www.mycobank.org; Mu et al. 2024).

In this study, we conducted extensive sample collection in southern China, primarily collecting plant leaves with obvious fungal necrosis or typical blight spot symptoms. Several Coniella fungi were collected from the diseased leaves of Ampelopsis grossedentata, Cinnamomum verum, Kadsura longipedunculata, and Lygodium circinnatum. Based on morphological and multi-locus analysis employing internal transcribed spacer (ITS), 28S large subunit ribosomal RNA gene (LSU), partial RNA polymerase II second largest subunit (rpb2), and translation elongation factor 1-alpha gene (tef1-α), four new Coniella species, namely C. diaoluoshanensis, C. dongshanlingensis, C. grossedentatae, and C. veri, were proposed.

Materials and methods

Sample collection and isolation

During 2022 to 2024, a large number of plant leaves that exhibited obvious signs of fungal necrosis or typical blight spot symptoms were collected from Fujian, Hainan, and Yunnan provinces in China. This study used tissue isolation methods to isolate fungi (Li et al. 2024). These diseased leaves were cut into small pieces of about 25 mm2 and surface sterilized by immersion in a 75% ethanol solution for 60 s, washed one time in sterile deionized water for 20 s, transferred to 5% sodium hypochlorite (NaOCl) for 90 s, and then washed three times in sterile deionized water for 60 s, subsequently dried on sterilized filter paper. The tissue pieces were transferred to the potato dextrose agar (PDA, 200 g potato, 20 g dextrose, 20 g agar, add deionized water and fill to 1000 mL, natural pH) plates and placed in a biological incubator at 25 °C for 3–4 days. The hyphal tips of individual colonies were transferred to new PDA plates to obtain pure cultures, which were then cut into 25 mm2 pieces using a sterile scalpel and stored in 2 mL frozen tubes containing 20% sterilized glycerin, with 8–10 pieces placed in each tube, for fungal strain preservation at -20 °C for further study.

Morphological and cultural characterization

The culture characteristics of the colonies were observed and photographed using a Sony Alpha 6400L digital camera (Sony Group Corporation, Tokyo, Japan) on 7 and 14 days, respectively. The micromorphological characteristics of the colonies were observed with the Olympus SZX10 stereomicroscope and Olympus BX53 microscope (Olympus Corporation, Tokyo, Japan), along with the BioHD-A20c color digital camera (FluoCa Scientific, China, Shanghai). Structural measurements were carried out using Digimizer software (v5.6.0) with a minimum of 30 measurements taken for each structure, such as conidiophores, conidiogenous cells, and conidia. The voucher specimens have been deposited in the Herbarium of the Department of Plant Pathology, Shandong Agricultural University, Taian, China (HSAUP). Additionally, the ex-type living cultures were deposited in the Shandong Agricultural University Culture Collection (SAUCC) and the China General Microbiological Culture Collection Center (CGMCC). The taxonomic information of the new taxa were submitted to MycoBank (http://www.mycobank.org, accessed on 2 Jan. 2025).

DNA extraction, PCR amplification, and sequencing

The DNA of the fungal genome was extracted using the modified cetyltrimethylammonium bromide (CTAB) method (Guo et al. 2000; Wang et al. 2023) or the magnetic bead kit method (OGPLF-400, GeneOnBio Corporation, Changchun, China) (Zhang et al. 2023). PCR amplifications of four genes (ITS, LSU, rpb2, and tef1-α) were done, and the corresponding primer pairs and PCR conditions were listed in Table 1. The PCR reaction was conducted in a 12 μL reaction volume, with a composition of 6 μL of 2 × Hieff Canace® Plus PCR Master Mix (with dye) (Cat. No. 10154ES03, Yeasen Biotechnology, Shanghai, China), 0.5 μL each of forward and reverse primer (10 μM TsingKe, Qingdao, China), and 0.5 μL of template genomic DNA (about 10 ng/μL), with the volume adjusted to 12 μL using distilled deionized water. PCR products were separated using 1% agarose gel and GelRed (TsingKe, Qingdao, China). Gel extraction was purified using a Gel Extraction Kit (Cat. No. AE0101-C, Shandong Sparkjade Biotechnology Co., Ltd., Jinan, China). The purified PCR products were subjected to bidirectional sequencing by Sangon Biotech Company Limited (Shanghai, China). The raw data were analyzed using MEGA v. 7.0 to obtain consistent sequences (Kumar et al. 2016). The sequence data have been deposited in GenBank, and their accession numbers were listed in Table 2.

Table 1.
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The primer sequences and PCR programs in this study.

Locus Primers Sequence (5’ – 3’) PCR cycles References
ITS ITS5 GGA AGT AAA AGT CGT AAC AAG G (94 °C: 30 s, 55 °C: 30 s, 72 °C: 45 s) × 29 cycles White et al. 1990
ITS4 TCC TCC GCT TAT TGA TAT GC
LSU LR0R GTA CCC GCT GAA CTT AAG C (94 °C: 30 s, 48 °C: 50 s, 72 °C: 1 min 30 s) × 35 cycles Vilgalys and Hester 1990; Rehner and Samuels 1994
LR5 TCC TGA GGG AAA CTT CG
rpb2 RPB2-5F2 GGG GWG AYC AGA AGA AGG C (94 °C: 45 s, 60 °C: 45 s, 72 °C: 2 min) × 5 cycles, (94 °C: 45 s, 54 °C: 45 s, 72 °C: 2 min) × 30 cycles Liu et al. 1999; Sung et al. 2007
RPB2-7CR CCC ATR GCT TGY TTR CCC AT
tef1-α EF1-728F CAT CGA GAA GTT CGA GAA GG (95 °C: 30 s, 51 °C: 30 s, 72 °C: 1 min) × 35 cycles O’Donnell et al. 1998; Carbone and Kohn 1999
EF2 GGA RGT ACC AGT SAT CAT GTT
Table 2.
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Species names, strain numbers, hosts or substrates, regions, and corresponding GenBank accession numbers of DNA sequences used in this study.

Species Strain numbers Host/Substrate Region GenBank accession numbers References
ITS LSU rpb2 tef1-α
Coniella africana CBS 114133* = CPC405 Eucalyptus nitens South Africa AY339344 AY339293 KX833421 KX833600 Van Niekerk et al. 2004; Alvarez et al. 2016
Coniella castanea SAUCC200313* Castanea mollissima China OL757537 OL757563 OL770463 OL780610 Wang et al. 2022
SAUCC200314 Castanea mollissima China OL757538 OL757564 OL770464 OL780611 Wang et al. 2022
Coniella cili GUCC 194020.1 Rosa roxburghii China ON791171 ON791212 ON815908 ON815944 Zhang et al. 2024b
GUCC 196007.1* Rosa roxburghii China ON791172 ON791213 ON815909 ON815945 Zhang et al. 2024b
Coniella crousii NFCCI 2213 Terminalia chebula India HQ264189 NA NA NA Rajeshkumar et al. 2011
Coniella diaoluoshanensis CGMCC3.27786* = SAUCC 7481-1 Kadsura longipedunculata China PQ357094 PQ357134 PQ361030 PQ404804 This study
SAUCC 7481-4 Kadsura longipedunculata China PQ357095 PQ357135 PQ361031 PQ404805 This study
Coniella diospyri CBS 145071* = CPC 34674 Diospyros mespiliformis South Africa MK047439 MK047489 MK047543 MK047562 Crous et al. 2018
Coniella diplodiella CBS 111858* = CPC3708 Vitis vinifera France AY339323 KX833335 KX833423 KX833603 Van Niekerk et al. 2004; Alvarez et al. 2016
CBS 112729 = CPC3927 Vitis vinifera South Africa KX833520 KX833345 KX833433 KX833613 Alvarez et al. 2016
Coniella diplodiopsis CBS 109.23 = CPC 3933 Vitis vinifera Switzerland NA AY339287 KX833440 KX833624 Van Niekerk et al. 2004; Alvarez et al. 2016
CBS 590.84* = CPC 3940 Vitis vinifera Italy AY339334 AY339288 NA NA Van Niekerk et al. 2004
CBS 116310 = CPC 3793 Vitis vinifera Italy KX833532 KX833357 KX833443 KX833627 Alvarez et al. 2016
Coniella dongshanlingensis CGMCC3.27785* = SAUCC 7265-5 Lygodium circinnatum China PQ357090 PQ357130 PQ361026 PQ404800 This study
SAUCC 7265-6 Lygodium circinnatum China PQ357091 PQ357131 PQ361027 PQ404801 This study
Coniella duckerae CBS 142045*= VPRI 13689 Lepidospermum concavum Australia KY924929 NA NA NA Marin-Felix et al. 2017
Coniella erumpens CBS 523.78* Rotten wood Chile KX833535 KX833361 KX833446 KX833630 Alvarez et al. 2016
Coniella eucalyptigena CBS 139893* = CPC 24793 Eucalyptus brassiana Malaysia KR476725 KR476760 NA NA Crous et al. 2015a
Coniella eucalyptorum CBS 112640* = CPC 3904 = DFR 100185 Eucalyptus grandis × E. tereticornis Australia AY339338 AY339290 KX833452 KX833637 Van Niekerk et al. 2004; Alvarez et al. 2016
CBS 114852 Eucalyptus sp. Australia KX833556 KX833380 KX833464 KX833652 Alvarez et al. 2016
Coniella fici MFLU 18-2578* Ficus septica China MW114356 MW114417 NA NA Tennakoon et al. 2021
Coniella fragariae CBS 172.49* = CPC 3930 Fragaria sp. Belgium AY339317 AY339282 KX833472 KX833663 Van Niekerk et al. 2004; Alvarez et al. 2016
CBS 454.68 Malus sylvestris Denmark KX833571 KX833393 KX833477 KX833670 Alvarez et al. 2016
Coniella fujianensis CGMCC3.25353 Canarium album China OR623057 OR623054 OR637413 OR637415 Mu et al. 2024
CGMCC3.25354* Canarium album China OR623058 OR623055 OR637414 OR637416 Mu et al. 2024
Coniella fusiformis CBS 141596* = CPC 19722 Eucalyptus sp. Indonesia KX833576 KX833397 KX833481 KX833674 Alvarez et al. 2016
CBS 114850 Eucalyptus pellita Australia KX833574 KX833395 KX833479 KX833672 Alvarez et al. 2016
Coniella granati CBS 132860 Punica granatum Turkey KX833577 KX833400 KX833484 KX833677 Alvarez et al. 2016
CBS 252.38 = ATCC 12685 = CPC 3714 Vitis vinifera Italy KX833581 AY339291 KX833488 KX833681 Van Niekerk et al. 2004; Alvarez et al. 2016
Coniella grossedentatae SAUCC 1354-1 Ampelopsis grossedentata China PQ357062 PQ357102 PQ361000 PQ404774 This study
CGMCC3.27783*= SAUCC 1354-3 Ampelopsis grossedentata China PQ357063 PQ357103 PQ361001 PQ404775 This study
Coniella heterospora CBS 143031* = FMR 15231 Herbivorous dung Spain LT800501 LT800500 LT800502 LT800503 Crous et al. 2017
Coniella hibisci CBS 109757* = AR 3534 Hibiscus sp. Africa KX833589 AF408337 NA KX833689 Castlebury et al. 2002; Marin-Felix et al. 2017
Coniella javanica CBS 455.68* Hibiscus sabdariffai Indonesia KX833583 KX833403 KX833489 KX833683 Alvarez et al. 2016
Coniella koreana CBS 143.97* NA South Korea KX833584 AF408378 KX833490 KX833684 Alvarez et al. 2016
Coniella lanneae CBS 141597* = CPC 22200 Lannea sp. Zambia KX833585 KX833404 KX833491 KX833685 Alvarez et al. 2016
Coniella limoniformis CBS 111021* = PPRI 3870 = CPC 3828 Fragaria sp. South Africa KX833586 KX833405 KX833492 KX833686 Alvarez et al. 2016
Coniella lustricola DAOMC 251731* NA America MF631778 MF631799 MF651900 MF651899 Raudabaugh et al. 2018
DAOMC 251732 NA America MF631779 MF631800 NA NA Raudabaugh et al. 2018
DAOMC 251733 NA America MF631780 MF631801 NA NA Raudabaugh et al. 2018
DAOMC 251734 NA America MF631781 MF631802 NA NA Raudabaugh et al. 2018
Coniella macrospora CBS 524.73* = CPC 3935 Terminalia ivoriensisstem Ivory Coast KX833587 AY339292 KX833493 KX833687 Alvarez et al. 2016
Coniella malaysiana CBS 141598* = CPC 16659 Corymbia torelliana Malaysia KX833588 KX833406 KX833494 KX833688 Alvarez et al. 2016
Coniella nicotianae CBS 875.72* = PD 72/793 Nicotiana tabacum Jamaica KX833590 KX833407 KX833495 KX833690 Alvarez et al. 2016
Coniella nigra CBS 165.60* = IMI 181519 = IMI 181599 = CPC 4198 Soil India AY339319 KX833408 KX833496 KX833691 Van Niekerk et al. 2004; Alvarez et al. 2016
Coniella obovata CBS 111025 = CPC 4196 = IMI 261318 Leaves South Africa AY339313 KX833409 KX833497 KX833692 Van Niekerk et al. 2004; Alvarez et al. 2016
Coniella paracastaneicola CBS 141292* = CPC 20146 Eucalyptus sp. Australia KX833591 KX833410 KX833498 KX833693 Alvarez et al. 2016
Coniella peruensis CBS 110394* = RMF 74.01 Soil of rain forest Peru KJ710463 KJ710441 KX833499 KX833695 Crous et al. 2015b; Alvarez et al. 2016
Coniella pseudodiospyri CBS 145540* = CPC 35725 Eucalyptus microcorys Australia MK876381 MK876422 MK876479 MK876493 Crous et al. 2019
Coniella pseudogranati CBS 137980* = CPC 22545 Terminalia stuhlmannii Zambia KJ869132 KJ869189 NA NA Crous et al. 2014b
Coniella pseudokoreana MFLU 13-0282* = MFLUCC 12-0427 Leaves Thailand MF190145 NA NA NA Senanayake et al. 2017
Coniella pseudostraminea CBS 112624* = IMI 233050 Fragaria sp. South Africa KX833593 KX833412 KX833500 KX833696 Alvarez et al. 2016
Coniella quercicola CBS 283.76 Excrements of Glomerus, which had eaten forest soil The Netherlands KX833594 KX833413 KX833501 KX833697 Alvarez et al. 2016
CBS 904.69* Quercus robur The Netherlands KX833595 KX833414 KX833502 KX833698 Alvarez et al. 2016
Coniella solicola CBS 766.71* Soil South Africa KX833597 KX833416 KX833505 KX833701 Alvarez et al. 2016
Coniella straminea CBS 149.22 = CPC 3932 Fragaria sp. USA AY339348 AY339296 KX833506 KX833704 Van Niekerk et al. 2004; Alvarez et al. 2016
Coniella tibouchinae CBS 131594* = CPC 18511 Tibouchina granulosa Brazil JQ281774 KX833418 KX833507 JQ281778 Miranda et al. 2012; Alvarez et al. 2016
Coniella veri CGMCC3.27787* = SAUCC 8877-4 Cinnamomum verum China PQ357098 PQ357138 PQ361034 PQ404810 This study
SAUCC 8877-7 Cinnamomum verum China PQ357099 PQ357139 PQ361035 PQ404811 This study
Coniella vitis MFLUCC 16-1399* = JZB3700001 Vitis vinifera China KX890008 KX890083 NA KX890058 Chethana et al. 2017
Coniella wangiensis CBS 132530* = CPC 19397 Eucalyptus sp. Australia JX069873 JX069857 KX833509 KX833705 Crous et al. 2012; Alvarez et al. 2016
Dwiroopa lythri CBS 109755* = AR 3383 Lythrum salicaria USA MN172410 MN172389 MN271801 MN271859 Jiang et al. 2020

Notes: New species established in this study are shown in bold. Those marked “*” in the table are represented as ex-type or ex-epitype strains. NA: Not available.

Sequence alignment and phylogenetic analyses

The nucleotide sequences of four new species were submitted to the NCBI’s GenBank nucleotide database (https://www.ncbi.nlm.nih.gov/, accessed on 2 Jan. 2025), and all related species were retrieved for phylogenetic analysis. Multiple sequences were aligned using MAFFT version 7 (http://mafft.cbrc.jp/alignment/server/index.html, accessed on 2 Jan. 2025) with default settings, and manual correction was applied if necessary (Katoh et al. 2019). For phylogenetic analyses, single and concatenated sequences were subjected to analysis by Maximum Likelihood (ML) and Bayesian Inference (BI) algorithms, respectively. Both ML and BI were executed on the CIPRES Science Gateway portal (https://www.phylo.org/, accessed on 2 Jan. 2025) or offline software (ML was executed in RaxML-HPC2 on XSEDE v8.2.12 and BI analysis was executed in MrBayes v3.2.7a with 64 threads on Linux) (Miller et al. 2012; Ronquist et al. 2012; Stamatakis 2014). For the ML analysis, the default parameters were used, and 1,000 rapid bootstrap replicates were run with the GTR+G+I model of nucleotide evolution; for BI, it was performed using a rapid bootstrapping algorithm with an automatic stop option and utilized MrModeltest v.2.3 to determine the best evolutionary model for each partition (Nylander 2004; Zhang et al. 2024a). Bayesian Inference posterior probabilities (BIPP) were evaluated by Markov Chain Monte Carlo (MCMC) (Rannala and Yang 1996; Zhaxybayeva and Gogarten 2002). The BI analyses encompassed two parallel runs spanning 5,000,000 generations, with a stop rule incorporated and a sampling frequency of 50 generations. The burn-in fraction was set at 0.25, and posterior probabilities were calculated from the remaining trees. The resulting trees were generated using FigTree v. 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree, accessed on 2 Jan. 2025) or ITOL: Interactive Tree of Life (https://itol.embl.de/, accessed on 2 Jan. 2025) (Letunic and Bork 2021), and the final layout of the trees was refined in Adobe Illustrator CC 2019. The names of the isolates in this study are marked in red in the phylogenetic tree.

Results

Molecular phylogeny

Initially, based on the ITS sequence data, we preliminarily determined that the eight strains belong to Coniella. Subsequently, based on ML and BI methods, we conducted a combined analysis of ITS, LSU, rpb2, and tef1-α gene data to construct phylogenetic trees for further determination of the phylogenetic position of these strains. The phylogenetic analysis of Coniella strains included 63 sequences, with Dwiroopa lythri (CBS 109755) serving as the outgroup. The final alignment comprised 2800 concatenated characters, viz. 1–600 (ITS), 601–1380 (LSU), 1381–2140 (rpb2), and 2141–2800 (tef1-α). The ML optimization likelihood was calculated to be -23461.791405. The matrix exhibited 1116 distinct alignment patterns, with 18.42% of characters or gaps remaining undetermined. The optimal models, evaluated by MrModeltest and selected in the BI, are as follows: the SYM+I+G model for ITS and the GTR+I+G model for LSU, rpb2, and tef1-α. The alignment exhibited a total of 1121 unique site patterns (ITS: 211, LSU: 78, rpb2: 322, tef1-α: 510). The topology of the ML tree concurred with that derived from BI; thus, only the ML tree is presented (Fig. 1). Combining morphological characteristics and molecular phylogenetic analyses, the eight strains in this study were introduced as four new species, namely Coniella diaoluoshanensis, C. dongshanlingensis, C. grossedentatae, and C. veri.

Figure 1. 

Phylogenetic relationship of Coniella based on concatenated sequences of ITS, LSU, rpb2, and tef1-α sequence data with Dwiroopa lythri (CBS 109755) as the outgroup. The Maximum Likelihood Bootstrap Value (left, MLBV ≥ 75%) and the Bayesian Inference Posterior Probability (right, BIPP ≥ 0.90) are shown as MLBV/BIPP above the nodes. The ex-type strains are marked with “*” and indicated in boldface. Strains from this study are shown in red. The scale bar at the bottom left represents 0.05 substitutions per site. Some branches are shortened according to the indicated multipliers to fit the page size, and these are indicated by the symbol (//).

Taxonomy

Coniella diaoluoshanensis D.H. Li, J.W. Xia & X.G. Zhang, sp. nov.

MycoBank No: 856520
Fig. 2

Holotype.

China • Hainan Province: Diaoluoshan National Forest Park, on diseased leaves of Kadsura longipedunculata (Schisandraceae), 18.660546°N, 109.936445°E, 94.1 m asl., 27 Mar. 2024, D.H. Li, holotype HSAUP 7481-1, ex-type living culture SAUCC 7481-1 = CGMCC3.27786.

Figure 2. 

Coniella diaoluoshanensis (CGMCC3.27786) a leaves of Kadsura longipedunculata b, c surface and reverse sides of colony after 14 days on PDA (b) and OA (c) d conidiomata forming on OA e, f conidiophores and conidiogenous cells with developing conidia g, h conidia. Scale bars: 10 μm (e–h).

Etymology.

Named after the collection site of the type specimen, Diaoluoshan National Forest Park.

Description.

Hypha immersed, 1.9–6.5 μm wide, branched, multi-septate, enlarged towards septum and terminal, hyaline. Asexual morph: Conidiomata nearly spherical, separate, scarce, immersed or superficial, surface uneven, sizes inconsistent, black. Conidiophores cylindrical, aseptate, straight or slightly curved, densely aggregated, simple, smooth, usually reduced to conidiogenous cells. Conidiogenous cells phialidic, simple, aggregative, hyaline, smooth, 8.1–11 × 1.4–2.6 μm (mean ± SD = 9.6 ± 0.8 × 2.1 ± 0.4 μm, n = 30), with apical periclinal thickening, blastospore at the apex. Conidia elliptical or fusiform, apices tapering, subobtuse, apically rounded, widest at the middle, bases tapering to a truncate hilum, multi-guttulate, immature conidia hyaline, mature conidia pale olivaceous, wall darker than pale olivaceous body of conidium, smooth, 7.5–9.3 × 4.7–5.5 μm (mean ± SD = 8.4 ± 0.5 × 5.1 ± 0.3 μm, n = 30). Sexual morph unknown.

Culture characteristics.

Colonies on PDA after 14 days of cultivation in the dark at 25 °C, reaching 75–77 mm in diam., with a growth rate of 5.4–5.5 mm/day; from above: white to cream-colored with age, sparse aerial mycelium at the center, irregularly circular, slightly low; peripheral mycelium dense, concentric rings, flat; colony edge irregular, sparse aerial mycelium, dispersed, striped; reverse: similar in color. Colonies on OA covering entire plate after 14 days of cultivation in the dark at 25 °C; from above: white, devoid of aerial mycelium at the center, with dispersed and sparse aerial mycelium at the edges; reverse: even white texture.

Additional material studied.

China • Hainan Province: Diaoluoshan National Forest Park, on diseased leaves of Kadsura longipedunculata (Schisandraceae), 18.660546°N, 109.936445°E, 94.1 m asl., 27 Mar. 2024, D.H. Li, HSAUP 7481-4, living culture SAUCC 7481-4.

Notes.

Phylogenetic analyses showed that Coniella diaoluoshanensis formed an independent clade (Fig. 1) and was closely related to C. eucalyptigena (CBS 139893), C. eucalyptorum (CBS 112640 and CBS 114852), and C. malaysiana (CBS 141598). Coniella diaoluoshanensis was distinguished from C. eucalyptigena by 4/573 and 7/791 base-pair differences in ITS and LSU sequences, from C. eucalyptorum (CBS 112640) by 19/565, 7/793, 68/765, and 164/539 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, and from C. malaysiana by 16/553, 7/783, 67/767, and 154/488 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, respectively. Morphologically, C. eucalyptigena lacks asexual sporulation description, making it impossible to compare microscopic structures with C. diaoluoshanensis. However, their macroscopic colony colors differ greatly: on PDA, C. diaoluoshanensis is cream-colored while C. eucalyptigena is salmon; on OA, C. diaoluoshanensis is white on the surface, whereas C. eucalyptigena is rosy buff. Morphologically, since C. eucalyptigena only had a description of sexual morphology, it could not be directly compared with the asexual morphology in this study. Then, C. eucalyptorum and C. malaysiana, which were closely related on the evolutionary tree, were selected for comparison. The conidiogenous cells of C. diaoluoshanensis (8.1–11 × 1.4–2.6 μm) shorter than those of C. eucalyptorum (10–17 × 3–3.5 μm) and C. malaysiana (8.5–18 × 1.5–3.5 μm); the conidia of C. diaoluoshanensis (7.5–9.3 × 4.7–5.5 μm) shorter than those of C. eucalyptorum (9–14 × 6–8 μm) and C. malaysiana (8–11.5 × 3–5 μm); and the mature conidial color of C. diaoluoshanensis (pale olivaceous) was lighter than that of C. eucalyptorum (medium to dark red-brown) and C. malaysiana (pale brown) (Van Niekerk et al. 2004; Crous et al. 2015a; Alvarez et al. 2016; Zhang et al. 2024b). Therefore, we describe our collection as a novel species.

Coniella dongshanlingensis D.H. Li, J.W. Xia & X.G. Zhang, sp. nov.

MycoBank No: 856519
Fig. 3

Holotype.

China • Hainan Province: Dongshanling Scenic Area, on diseased leaves of Lygodium circinnatum (Lygodiaceae), 18.802153°N, 110.421473°E, 18.8 m asl., 26 Mar. 2024, D.H. Li, holotype HSAUP 7265-5, ex-type living culture SAUCC 7265-5 = CGMCC3.27785.

Figure 3. 

Coniella dongshanlingensis (CGMCC3.27785) a a leaf of Lygodium circinnatum b, c surface and reverse sides of colony after 14 days on PDA (b) and OA (c) d, e conidiomata forming on PDA f, g conidiophores and conidiogenous cells with developing conidia h, i conidia. Scale bars: 10 μm (f–i).

Etymology.

Named after the collection site of the type specimen, Dongshanling Scenic Area.

Description.

Hypha superficial, 1.1–3.2 μm wide, less branched, multi-septate, hyaline to pale yellow. Asexual morph: Conidiomata pycnidial to nearly spherical, separate, superficial, surface enveloped in a gelatinous sheath, sizes inconsistent, initially appearing hyaline, becoming black with mature. Conidiophores cylindrical, aseptate, straight or slightly curved, densely aggregated, simple, smooth, usually reduced to conidiogenous cells. Conidiogenous cells phialidic, simple, aggregative, hyaline, smooth, 7.3–19.2 × 1.5–3.3 μm (mean ± SD = 12.6 ± 2.6 × 2.4 ± 0.5 μm, n = 30), with apical periclinal thickening, blastospore at the apex. Conidia elliptical to fusiform, apices tapering, subobtuse, apically rounded, bases tapering to a truncate hilum, immature conidia hyaline, multi-guttulate, mature conidia olivaceous, 1–2 guttulate, wall darker than olivaceous body of conidium, smooth, 7.8–10 × 5.1–7 μm (mean ± SD = 8.7 ± 0.6 × 6.2 ± 0.4 μm, n = 30). Sexual morph unknown.

Culture characteristics.

Colonies on PDA after 14 days of cultivation in the dark at 25 °C, reaching 47–50 mm in diam., with a growth rate of 3.4–3.6 mm/day; from above: white to pale orange with age, medium aerial mycelium, circular, slightly low at the center, slightly higher at the edges; reverse: similar in color. Colonies on OA covering entire plate after 14 days of cultivation in the dark at 25 °C; from above: pale orange, interspersed with extensive black pycnidia, medium aerial mycelium, flat; reverse: similar in color.

Additional material studied.

China • Hainan Province: Dongshanling Scenic Area, on diseased leaves of Lygodium circinnatum (Lygodiaceae), 18.802153°N, 110.421473°E, 18.8 m asl., 26 Mar. 2024, D.H. Li, HSAUP 7265-6, living culture SAUCC 7265-6.

Notes.

Phylogenetic analyses showed that Coniella dongshanlingensis formed an independent clade (Fig. 1) and was closely related to C. fujianensis (CGMCC3.25353 and CGMCC3.25354). Coniella dongshanlingensis was distinguished from C. fujianensis (CGMCC3.25354) by 5/589, 9/657, and 19/306 base-pair differences in ITS, rpb2, and tef1-α sequences, respectively. Morphologically, the conidiogenous cells of C. dongshanlingensis (7.3–19.2 × 1.5–3.3 μm) are longer than those of C. fujianensis (3.5–8 × 2.5–3.5 μm); the conidia of C. dongshanlingensis (7.8–10 × 5.1–7 μm) slightly shorter than those of C. fujianensis (8–10.5 × 5.5–7.5 μm), and the mature conidial color of C. dongshanlingensis (olivaceous) is lighter than that of C. fujianensis (brown) (Mu et al. 2024). Therefore, we describe our collection as a novel species.

Coniella grossedentatae D.H. Li, J.W. Xia & X.G. Zhang, sp. nov.

MycoBank No: 856518
Fig. 4

Holotype.

China • Fujian Province: Wuyishan City, Xingcun Town, on diseased leaves of Ampelopsis grossedentata (Vitaceae), 27.749556°N, 117.679038°E, 751.68 m asl., 15 Oct. 2022, D.H. Li, holotype HSAUP 1354-3, ex-type living culture SAUCC 1354-3 = CGMCC3.27783.

Figure 4. 

Coniella grossedentatae (CGMCC3.27783) a leaves of Ampelopsis grossedentata b, c surface and reverse sides of colony after 14 days on PDA (b) and OA (c) d colony on PDA e conidiomata forming on pine needle f, g conidiophores and conidiogenous cells with developing conidia h, i conidia. Scale bars: 10 μm (f–i).

Etymology.

Named after the species epithet of the host plant, Ampelopsis grossedentata.

Description.

Hypha superficial, 1.3–3.5 μm wide, branched, multi-septate, hyaline to pale orange. Asexual morph: Conidiomata spherical or narrowly ellipsoid, separate, immersed or superficial, some surfaces enveloped in a gelatinous sheath, some surface uneven, sizes inconsistent, black. Conidiophores cylindrical, aseptate, straight or slightly curved, densely aggregated, simple, usually reduced to conidiogenous cells. Conidiogenous cells phialidic, simple, aggregative, hyaline, smooth, 10.6–23.1 × 1.7–3.8 μm (mean ± SD = 16.8 ± 3 × 2.5 ± 0.6 μm, n = 30), with apical periclinal thickening, blastospore at the apex. Conidia nearly spherical, apices acute, widest at the middle, bases tapering to a truncate hilum, multi-guttulate, immature conidia hyaline, mature conidia medium brown, wall darker than medium brown body of conidium, smooth, 8–10.5 × 7.5–9.5 μm (mean ± SD = 9.4 ± 0.6 × 8.4 ± 0.5 μm, n = 30). Sexual morph unknown.

Culture characteristics.

Colonies on PDA after 14 days of cultivation in the dark at 25 °C, reaching 86–90 mm in diam., with a growth rate of 6.1–6.4 mm/day; from above: orange in the middle and edges, with white in between, medium aerial mycelium, granular, circular, flat; reverse: similar in color. Colonies on OA covering entire plate after 14 days of cultivation in the dark at 25 °C; from above: white in the middle and edges, with orange in between, sparse aerial mycelium, flat; reverse: similar in color.

Additional material studied.

China • Fujian Province: Wuyishan City, Xingcun Town, on diseased leaves of Ampelopsis grossedentata (Vitaceae), 27.749556°N, 117.679038°E, 751.68 m asl., 15 Oct. 2022, D.H. Li, HSAUP 1354-1, living culture SAUCC 1354-1.

Notes.

Phylogenetic analyses showed that Coniella grossedentatae formed an independent clade (Fig. 1) basal to C. dongshanlingensis (CGMCC3.27785, SAUCC 7265-6), C. fujianensis (CGMCC 3.25353, CGMCC 3.25354), and C. wangiensis (CBS 132530). Coniella grossedentatae can be distinguished from C. dongshanlingensis by 4/604, 1/793, 52/902, and 80/532 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, and from C. fujianensis by 8/588, 1/798, 34/657, and 64/313 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, and from C. wangiensis by 2/603, 5/798, 35/767, and 79/329 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, respectively. Morphologically, the conidiogenous cells of C. grossedentatae (10.6–23.1 × 1.7–3.8 μm) are longer than those of C. dongshanlingensis (7.3–19.2 × 1.5–3.3 μm), C. fujianensis (3.5–8 × 2.5–3.5 μm), and C. wangiensis (15–20 × 3–4 μm); the conidia of C. grossedentatae (8–10.5 × 7.5–9.5 μm) are wider than those of C. dongshanlingensis (7.8–10 × 5.1–7 μm) and C. fujianensis (8–10.5 × 5.5–7.5 μm), and shorter than those of C. wangiensis (9–13 × 7–10 μm) (Crous et al. 2012; Alvarez et al. 2016). Therefore, we describe our collection as a novel species.

Coniella veri D.H. Li, J.W. Xia & X.G. Zhang, sp. nov.

MycoBank No: 856521
Fig. 5

Holotype.

China • Yunnan Province: Pu’er City, Yixiang Town, Pu’er Sun River Forest Park, on diseased leaves of Cinnamomum verum (Lauraceae), 22.593953°N, 101.086217°E, 1596.44 m asl., 15 May 2024, D.H. Li, holotype HSAUP 8877-4, ex-type living culture SAUCC 8877-4 = CGMCC3.27787.

Figure 5. 

Coniella veri (CGMCC3.27787) a leaves of Cinnamomum verum b, c surface and reverse sides of colony after 14 days on PDA (b) and OA (c) d conidiomata forming on OA e, f conidiophores and conidiogenous cells with developing conidia g, h conidia. Scale bars: 10 μm (e–h).

Etymology.

Named after the species epithet of the host plant, Cinnamomum verum.

Description.

Hypha superficial, 1.3–3.3 μm wide, branched, multi-septate, hyaline. Asexual morph: Conidiomata spherical, aggregated or solitary, immersed or superficial, some surfaces enveloped in a gelatinous sheath, some surface uneven, sizes inconsistent, initially appearing hyaline, becoming black with mature. Conidiophores cylindrical, septate, branched, straight or slightly curved, densely aggregated, simple, usually reduced to conidiogenous cells. Conidiogenous cells phialidic, simple, aggregative, or solitary, hyaline, smooth, 9.5–17.5 × 1.2–2.5 μm (mean ± SD = 12.5 ± 1.5 × 1.8 ± 0.4 μm, n = 30), with apical periclinal thickening, blastospore at the apex. Conidia elliptical to fusiform, apices acute, widest at the middle, bases tapering to a truncate hilum, multi-guttulate gather at both ends, hyaline, thick-walled, smooth, 6.2–8.8 × 3.6–4.7 μm (mean ± SD = 7.7 ± 0.6 × 4 ± 0.3 μm, n = 30). Sexual morph unknown.

Culture characteristics.

Colonies on PDA after 14 days of cultivation in the dark at 25 °C, reaching 81–85 mm in diam., with a growth rate of 5.8–6.1 mm/day; from above: white, medium aerial mycelium, slightly higher at the center, circular, radial, flat; reverse: pale orange in the middle, orange in the edges. Colonies on OA after 14 days of cultivation in the dark at 25 °C, reaching 72–77 mm in diam., had a growth rate of 5.1–5.5 mm/day; from above: white, sparse aerial mycelium, black pycnidia formed in the center, flat; reverse: similar in color.

Additional material studied.

China • Yunnan Province: Pu’er City, Yixiang Town, Pu’er Sun River Forest Park, on diseased leaves of Cinnamomum verum (Lauraceae), 22.593953°N, 101.086217°E, 1596.44 m asl., 15 May 2024, D.H. Li, HSAUP 8877-7, living culture SAUCC 8877-7.

Notes.

Phylogenetic analyses showed that Coniella veri formed an independent clade (Fig. 1) and was closely related to C. cili (GUCC 194020.1 and GUCC 196007.1). Coniella veri can be distinguished from C. cili (GUCC 196007.1) by 31/597, 8/791, 52/869, and 125/516 base-pair differences in ITS, LSU, rpb2, and tef1-α sequences, respectively. Morphologically, the conidiogenous cells of C. veri (9.5–17.5 × 1.2–2.5 μm) are shorter than those of C. cili (13–23.5 × 1–2 μm); the conidia of C. veri (6.2–8.8 × 3.6–4.7 μm) are shorter than those of C. cili (5.5–17.5 × 2.5–5 μm); the conidial shape of C. veri is elliptical to fusiform, whereas the conidial size and shape of C. cili exhibit considerable variation, including limoniform, fusoid, clavate, cylindrical, and elongated elliptical forms (Zhang et al. 2024b). Therefore, we describe our collection as a novel species.

Discussion

Coniella species have a worldwide distribution, reported in countries across all continents (Van Niekerk et al. 2004; Alvarez et al. 2016). They have been found in Asia (e.g., China, India, Indonesia, Malaysia, South Korea, and Thailand), Europe (e.g., Belgium, Denmark, France, Italy, the Netherlands, Switzerland, and Spain), Africa (e.g., Ivory Coast, South Africa, and Zambia), the Americas (e.g., the United States, Brazil, Peru, Jamaica, and Chile), and Oceania (e.g., Australia). These countries, ranging from landlocked nations such as Zambia and Switzerland to coastal countries like China, Brazil, and Australia, as well as island nations including Jamaica and Indonesia, are geographically diverse. They are distributed on both sides of the equator and span multiple climatic zones, from tropical to frigid, coastal to inland, and plain to mountain, encompassing diverse climate types such as tropical, temperate, and alpine. Many countries, including most of Africa, northern Brazil, Indonesia, and Malaysia, have tropical climates with high temperatures and abundant precipitation year-round. China, with its vast territory, large latitudinal span, wide longitudinal extent, and complex and diverse topography, nearly covers all major climate types, providing favorable conditions for the formation of Coniella species diversity (Castlebury et al. 2002; Van Niekerk et al. 2004; Alvarez et al. 2016; Raudabaugh et al. 2018; Wang et al. 2022; Mu et al. 2024; Zhang et al. 2024b).

Currently, Coniella has accepted 66 species, many of which were introduced solely based on morphological studies (Index Fungorum: https://indexfungorum.org; MycoBank: http://www.mycobank.org; Alvarez et al. 2016; Mu et al. 2024). Morphological characteristics of some conidia are highly similar and can be classified into two categories: one comprises olivaceous brown to brown conidia that are ellipsoid or globose, while the other category consists of hyaline conidia that are fusiform or clavate, often with very similar shapes and sizes. Rendering precise identification of Coniella species difficult solely on morphological characteristics (Crous et al. 2014a). Consequently, there is a strong current trend towards integrating morphological and molecular methods to assess or clarify the taxonomic placement and phylogenetic relationships of Coniella species (Alvarez et al. 2016). Based on phylogenetic analyses of ITS, LSU, and tef1-α sequence data, Van Niekerk et al. (2004) demonstrated that Coniella represents a distinct evolutionary lineage within the Diaporthales (Van Niekerk et al. 2004). Based on phylogenetic analyses of ITS, LSU, rpb2, and tef1-α sequence data, Alvarez et al. (2016) conducted a taxonomic revision of the genus. Since then, phylogenetic analyses of Coniella have largely continued to use these four genetic loci (Alvarez et al. 2016).

According to previous studies, Coniella species have been recorded as plant pathogens, endophytes, and saprobes (Samuels et al. 1993; Ferreira et al. 1997; Alvarez et al. 2016; Chethana et al. 2017). Their hosts encompass multiple categories, including plants (such as trees, shrubs, herbs, and ferns), animal excreta, and soils (Crous et al. 2015b; Alvarez et al. 2016). In recent years, several Coniella species have been reported and described in China. For example, Fröhlich and Hyde (2000) discovered C. calamicola on both living and dead leaves of Daemonorops margaritae in Hong Kong. Chen et al. (2014) first reported that C. granati can cause fruit rot and twig blight in pomegranate (Punica granatum) in Anhui Province. Chethana et al. (2017) reported that C. vitis is the pathogenic fungus causing white rot in grapes (Vitis vinifera) in Beijing Municipality, Guangxi, Hebei, Henan, and Jilin Provinces. Tennakoon et al. (2021) isolated a new species, C. fici, from dead leaves of Ficus septica (Moraceae) on the island of Taiwan. Wang et al. (2022) isolated a new species, C. castanea, from symptomatic leaves of Castanea mollissima (Fagaceae) in an orchard in Shandong Province. Mu et al. (2024) isolated a new species, C. fujianensis, from diseased leaves of Canarium album (Burseraceae) in Fujian Province. Zhang et al. (2024b) isolated the endophytic species C. cili from healthy fruits and seeds of Rosa roxburghii (Rosaceae) in Guizhou Province.

During a continuous survey of terrestrial plant fungi in certain regions of southern China, four new species of Coniella were discovered from diseased leaf tissues of infected plants in Fujian, Hainan, and Yunnan provinces. These new species are named Coniella diaoluoshanensis, C. dongshanlingensis, C. grossedentatae, and C. veri. Among them, C. grossedentatae utilizes Ampelopsis grossedentata (Vitaceae) as its host. Van Niekerk et al. (2004) have previously reported species of C. diplodiopsis isolated from Vitis vinifera (Vitaceae) collected in Italy. In contrast, C. diaoluoshanensis, C. dongshanlingensis, and C. veri are the first reports that are associated with the hosts Kadsura longipedunculata, Lygodium circinnatum, and Cinnamomum verum, respectively. This will further broaden the host range of Coniella species and contribute to the fields of plant pathology and fungal taxonomy. With the increasing number of Coniella species, we believe that comprehensive research on this genus will uncover more hidden Coniella species from terrestrial plants.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by the National Natural Science Foundation of China (nos. 32370001, 32270024, 31900014, U2002203), the Key Technological Innovation Program of Shandong Province, China (no. 2022CXGC020710), the Jinan City’s ‘New University 20 Policies’ Initiative for Innovative Research Teams Project (no. 202228028) and the Innovative Agricultural Application Technology Project of Jinan City (no. CX202210).

Author contributions

Sampling, molecular biology analysis: Duhua Li and Zixu Dong; fungal isolation: Qiyun Liu and Yaling Wang; description and phylogenetic analysis: Duhua Li and Zhaoxue Zhang; microscopy: Duhua Li and Jiwen Xia; writing-original draft preparation: Duhua Li; writing-review and editing: Xiuguo Zhang and Jiwen Xia. All authors read and approved the final manuscript.

Author ORCIDs

Duhua Li https://orcid.org/0009-0006-5200-2034

Qiyun Liu https://orcid.org/0009-0009-9545-7962

Zhaoxue Zhang https://orcid.org/0000-0002-4824-9716

Xiuguo Zhang https://orcid.org/0000-0001-9733-8494

Jiwen Xia https://orcid.org/0000-0002-7436-7249

Data availability

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

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