Unveiling species diversity within early-diverging fungi from China V: Five new species of Absidia (Cunninghamellaceae, Mucoromycota)

Xin-Yu Ji; Zi-Ying Ding; Yong Nie; Heng Zhao; Shi Wang; Bo Huang; Xiao-Yong Liu

doi:10.3897/mycokeys.117.149185

Research Article

Unveiling species diversity within early-diverging fungi from China V: Five new species of Absidia (Cunninghamellaceae, Mucoromycota)

Xin-Yu Ji^‡, Zi-Ying Ding^‡, Yong Nie^§, Heng Zhao^§, Shi Wang^‡, Bo Huang^|, Xiao-Yong Liu^‡¶

‡ Shandong Normal University, Jinan, China

§ Anhui University of Technology, Anhui, China

| Anhui Agricultural University, Hefei, China

¶ Chinese Academy of Sciences, Beijing, China

Corresponding author: Bo Huang ( bhuang@ahau.edu.cn )

Corresponding author: Xiao-Yong Liu ( liuxy@sdnu.edu.cn )

Academic editor: Ajay Kumar Gautam

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: Ji X-Y, Ding Z-Y, Nie Y, Zhao H, Wang S, Huang B, Liu X-Y (2025) Unveiling species diversity within early-diverging fungi from China V: Five new species of Absidia (Cunninghamellaceae, Mucoromycota). MycoKeys 117: 267-288. https://doi.org/10.3897/mycokeys.117.149185

Abstract

Absidia is widely distributed in soil, koji, and various types of feces. A multi-locus phylogeny covering the small subunit (SSU), internal transcribed spacer (ITS), and large subunit of ribosomal RNA gene (LSU rDNA), translation elongation factor 1-alpha (TEF1α), and actin (Act), combined with morphological characteristics, revealed five new species in this genus. This study provides their descriptions and illustrations and discusses their differences from morphological allies and phylogenetic relatives. Absidia collariata sp. nov. is distinguished from other species in terms of the length of collars. A. hainanensis sp. nov. is named after the geographical location Hainan, distinctive with a higher maximum growing temperature. A. pyriformis sp. nov. is different from other species in terms of sporangial shape. A. tardiva sp. nov. is characterized by slow growth. And A. tibetensis sp. nov. is named after the geographical location Tibet and differentiated by more ampulliform swellings. This study further enriches the species diversity of Absidia as the latest discovery of early-diverging fungi in China.

Key words:

Fungal diversity, molecular phylogeny, Mucorales, soil-borne fungi, taxonomy

Introduction

Absidia Tiegh. belongs to Mucoromycota, Mucoromycotina, Mucoromycetes, Mucorales, and Cunninghamellaceae (http://www.indexfungorum.org/, accessed on 1 November 2024). This genus was founded in 1876 and typified by A. reflexa Tiegh (Tieghem 1878). This genus is ubiquitous and widely distributed (Zhang et al. 2018; Crous et al. 2020; Hurdeal et al. 2021; Urquhart and Idnurm 2021; Zhao et al. 2021b; Zhao et al. 2022a); for example, the Czech Republic (688 records), Australia (862), Estonia (1621), Lithuania (422), Italy (372), South Africa (198), and Argentina (515) (https://www.gbif.org/, accessed on 2 November 2024). Absidia are most frequently encountered in July, August, and September, and relatively high temperatures favor Absidia members (https://www.gbif.org/, accessed on 2 November 2024). In China, they are mainly found in tropical regions such as Yunnan and Hainan provinces. They are occasionally isolated in places with low temperatures, such as Tibet and Jilin (Zhao et al. 2022a). The species of Absidia are usually isolated mainly in soil samples. They are also found in plant debris, herbivorous feces, decaying substrates, and air (Lima et al. 2020). The genus Absidia is very important in industry and medicine because it produces steroids, laccases, fatty acids, and other useful substances (Davoust and Persson 1992; Kristanti et al. 2016; Zhao et al. 2021b). Absidia is also frequently used in biotransformation of various natural products, including reduction reactions, hydroxylation, and glycosylation (Zhao et al. 2022a). Moreover, Absidia plays a vital role as the causative agent for human mucormycoses (Constantinides et al. 2008). And it is a contaminant in wine production (Zhao et al. 2022a).

Currently, there are 139 records of Absidia species, variants, and subspecies in the Index Fungorum database (http://www.indexfungorum.org/, accessed November 2, 2024). Absidia usually produces erect or slightly bent sporangia. There is a septum under sporangia. The sporangia are mostly nearly spherical to pyriform, deliquescent-walled, and multi-spored. Sporangiophores arise singly or in whorls. A small protuberance sometimes appears at the apex of columellae. Collars are evident if present. And zygospores have many appendages (Tieghem 1878; Guan et al. 2021; Zhao et al. 2021b; Zong et al. 2021).

Over the past few years, Absidia has experienced a rapid influx of proposed new species. (Lima et al. 2020; Zong et al. 2021; Cordeiro et al. 2020; Zhao et al. 2023). In this paper, five new species, A. collariata sp. nov., A. hainanensis sp. nov., A. pyriformis sp. nov., A. tardiva sp. nov., and A. tibetensis sp. nov., are described from the soils in Yunnan Province, Hainan Province, and Tibet, based on evidence of molecular phylogeny, morphogenetic characteristics, and maximum growth temperature. This is the fifth report of a serial work on the diversity of Chinese early-diverging fungi (Tao et al. 2024; Wang et al. 2024; Zhao et al. 2024; Ding et al. 2025).

Materials and methods

Isolation and morphology

In 2024, soil samples were collected from Yunnan, Tibet, and Hainan in China, following the methods of Yu Li and his colleagues (Liu et al. 2019; Zou et al. 2022). Each sample (approximately 50 g) was placed in a sterile Whirl-Pack plastic bag, labeled with a number, date, vegetation type, altitude, latitude, and longitude. All samples were preserved at 4 °C upon delivery to the laboratory. By combining soil dilution plate and moist-chamber culture methods, pure strains were isolated from soil samples (Constantinides et al. 2008; Kristanti et al. 2016; Zhao et al. 2021a). About 1 g of soil was put into a 10 mL centrifuge tube with 10 mL of sterile water and shaken on a shaker for 25 minutes to prepare a soil suspension. One milliliter of the starting suspension was transferred into 9 mL of sterile water to obtain a 10^-2 soil suspension. The above steps were repeated to get 10^-3 and 10^-4 soil suspensions. Approximately 200 μL of the 10^-3 and 10^-4 soil suspensions were pipetted onto the Rose Bengal Chloramphenicol agar (RBC: peptone 5.00 g/L, KH₂PO₄ 1.00 g/L, MgSO₄·7H₂O 0.50 g/L, Rose Bengal 0.05 g/L, glucose 10.00 g/L, chloramphenicol 0.10 g/L, agar 15.00 g/L) and dispersed evenly with a sterilized triangular glass coating rod. The plates were cultivated at 26 °C in the dark for 2–5 days (Corry et al. 1995). Subsequently, the hyphae at the edge of the colony were transferred to a potato dextrose agar (PDA: glucose 100 g, potato 1000 g, agar 100 g, sterilized water 5000 mL, and pH 7). Upon colony forming, a macro shot was taken with a digital camera (Canon PowerShot G7X, Canon, Tokyo, Japan). For the moist chamber, 1 g of soil was evenly sprinkled onto the surface of PDA plates, sealed with Parafilm, and cultivated inverted at 26 °C away from light. After three days, the purification of target strains was performed using an inoculation ring streak. After two days, the agar with hyphae located at the edge of the colony was transferred to a new potato dextrose agar and cultivated as above. Microscopic morphological characteristics of fungi were observed with a stereoscope (SZX10, OLYMPUS, Tokyo, Japan) and a light microscope (BX53, OLYMPUS, Tokyo, Japan) and photographed with a high-definition color digital camera (DP80, OLYMPUS, Tokyo, Japan). Structural measurements were performed using Digimizer software (v5.6.0), and each trait (sporangiospores, stolons, apophyses, and so on) was measured 25 or more times. A gradient method was used to determine its maximum growth temperature (Hoffmann et al. 2007; Hoffmann and Voigt 2009; Hoffmann 2010). Cultured colonies were incubated at 25 °C for two days, and then the temperature was increased by one degree Celsius per day until no further growth was observed. This temperature was recorded as the maximum growth temperature. All strains were stored in 10% sterilized glycerin at -20 °C and preserved in the Shandong Normal University Culture Collection (XG). The living ex-holotype cultures were stored in the China Microbiological Culture Collection Center, Beijing, China (CGMCC). Dry cultures of holotypes were submitted to the Herbarium Mycologicum Academiae Sinicae, Beijing, China (HMAS). The taxonomic information was uploaded to the Fungal Names repository (https://nmdc.cn/fungalnames/).

DNA extraction, PCR amplification, and sequencing

Fungal genomic DNA was extracted using a DNA extraction kit (Cat. No.: 70409-20; BEAVER Biomedical Engineering Co., Ltd.) (Doyle et al. 1990; Wang et al. 2023). The ITS, LSU, TEF1α, Act, and SSU regions were amplified using the primer pairs and polymerase chain reaction (PCR) programs specified in Table 1. The final volume of the reaction mixture was 25 μL, containing 12.5 μL 2 × Hieff Canace Plus PCR Master Mix with dye (Yeasen Biotechnology, Cat. No. 10154ES03), 9.5 μL ddH₂O, 1 μL forward primer (10 μM), 1 μL reverse primer (10 μM), and 1 µL of genomic DNA template (1 ng/μL). The PCR programs are listed in Table 1. PCR amplification products were observed on a 2% agarose electrophoresis gel. Fragments were visualized at 254 nm UV light (Zhang et al. 2022). The amplified product was purified using a gel extraction kit (Cat# AE0101-C, Shandong Sparkjade Biotechnology Co., Ltd.). Primer synthesis and DNA sequencing were carried out by Tsingke Biotechnology (Beijing, China). MEGA v. 7.0 (Mega Limited, Auckland, New Zealand) was used to obtain consensus sequences. Finally, all sequences generated in this study were deposited in GenBank.

Table 1.

Download as

CSV

XLSX

PCR information used in this study.

Loci	PCR primers	Primer sequence (5’ – 3’)	PCR cycles	References
ITS	ITS5	GGA AGT AAA AGT CGT AAC AAG G	95 °C 5 min; (95 °C: 30 s, 55 °C: 30 s, 72 °C: 1 min) × 35 cycles; 72 °C 10 min	White et al. (1990)
ITS	ITS4	TCC TCC GCT TAT TGA TAT GC		White et al. (1990)
LSU	LR0R	GTA CCC GCT GAA CTT AAG C	95 °C 5 min; (94 °C: 30 s, 52 °C: 45 s, 72 °C: 1.5 min) × 30 cycles; 72 °C 10 min	Hurdeal et al. (2023)
LSU	LR5	TCC TGA GGG AAA CTT CG		Hurdeal et al. (2023)
TEF1α	EF1-983F	GCYCCYGGHCAYCGTGAYTTYAT	95 °C 5 min; (95 °C: 30 s, 55 °C: 60 s, 72 °C: 60 s) × 30 cycles; 72 °C 10 min	Jaklitsch et al. (2005)
TEF1α	TEF1LLErev	AACTTGCAGGCAATGTGG		Jaklitsch et al. (2005)
Act	ACT-1	TGG GAC GAT ATG GAI AAI ATC TGG CA	95 °C 3 min; (95 °C: 60 s, 55 °C: 60 s, 72 °C: 1 min) × 30 cycles; 72 °C 10 min	Voigt and Wöstemeyer (2000)
Act	ACT-4R	TC ITC GTA TIC TIG CTI IGA IAT CCA CA T		Voigt and Wöstemeyer (2000)
SSU	NS1	GTA GTC ATA TGC TTG TCT CC	95 °C 5 min; (94 °C: 60 s, 54 °C: 50 s, 72 °C: 1 min) × 37 cycles; 72 °C 10 min	(White et al. 1990)
SSU	NS4	CTT CCG TCA ATT CCT TTA AG		(White et al. 1990)

Phylogenetic analyses

Reference sequences were downloaded according to the latest articles (Zhao et al. 2021b; Zhao et al. 2022a; Tao et al. 2024; Zhao et al. 2024). Phylogenetic analyses were performed for each marker individually, followed by a combined analysis (ITS-LSU-SSU-Act-TEF1α). The sequences newly obtained in this study were compared with reference sequences in GenBank using MEGA v.7.0 software (Larsson 2014; Kumar et al. 2016). The phylogeny was inferred using maximum likelihood (ML) and Bayesian inference (BI) algorithms (Nie et al. 2020a; Nie et al. 2020b), which were integrated with the CIPRES Science Portal (https://www.phylo.org/, accessed November 5, 2024). ML analysis was performed using RaxML 8.2.4 (https://www.phylo.org/) in CIPRES Science Gateway V. 3.3 with 1,000 bootstrap replicates (Miller et al. 2010; Nguyen et al. 2015). BI analysis was performed using the GTR + I + G model with a sampling frequency of every 1,000 generations, and eight cold Markov chains were run simultaneously for two million generations (Ronquist et al. 2012; Stamatakis 2014). Utilizing the iTOL website (https://itol.embl.de, accessed November 5, 2024), all trees that resulted were plotted and optimized. Finally, Adobe Illustrator CC 2019 was used to beautify the phylogenetic tree (Jiang et al. 2024).

Results

Molecular phylogeny

Phylogenetic analyses were performed on a dataset containing 103 isolates, including 93 strains retrieved from GenBank and 10 acquired herein. Of these, 99 isolates were classified as the ingroup Absidia, while four strains, Cunninghamella elegans (CBS 160.28), C. elegans (CBS 167.53), C. blakesleeana (CBS 133.27), and C. blakesleeana (CBS 782.68), were employed as outgroups. In total, it consisted of 5,087 concatenated characters: 1–1,102 (ITS), 1,103–2,072 (LSU), 2,073–3,157 (TEF1α), 3,158–4,118 (Act), and 4,119–5,087 (SSU). Among these, 2,766 characters remained constant, 724 were variable and parsimony-uninformative, and 1,597 were parsimony-informative. The maximum likelihood (ML) tree (Fig. 1) and Bayesian tree showed comparable topological structures.

Figure 1.

Phylogram of the genus Absidia based on a concatenated ITS, LSU, TEF1α, Act, and SSU sequence alignment, with Cunninghamella elegans and C. blakesleeana serving as outgroups. The robustness of branches is marked at the node with the Maximum Likelihood Bootstrap Value (left, MLBV ≥ 70%) and Bayesian Inference Posterior Probability (right, BIPP ≥ 0.90), which are separated by a slash “/”. Ten newly isolated strains are indicated in red bold. Branches shortened to fit the page are indicated by a double slash “//”. Bold strains marked with a star marker “*” are ex-types or ex-holotypes. The scale at the bottom left indicates 0.1 substitutions per site.

Taxonomy

Absidia collariata X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Fungal Names: FN 572256

Fig. 2

Type.

China • Yunnan Province, Yuxi City, Xinping Yi Dai Autonomous County, Ancient Tea Horse Road (23°57'28"N, 101°30'38"E, 2196.56 m), from soil, 5 Jul. 2024, X.Y. Ji and X.Y. Liu, holotype HMAS 353360, ex-holotype living culture CGMCC 3.28536 (=XG08666-10-1).

Figure 2.

Absidia collariata ex-holotype CGMCC 3.28536 a, b colonies on PDA (a obverse b reverse) c–e sporangia f, g swelling on sporangiophores h–j columellae k rhizoids l sporangiospores. Scale bars: 10 μm (c–l).

Etymology.

The collariata (Lat.) refers to its long collars.

Description.

Colonies on PDA at 26 °C for 5 days, reaching 68 mm in diameter, moderately fast growing with a rate of 13.6 mm/d, higher in the center than at margin, at first white, becoming grayish brown when mature, regular shape at reverse. Hyphae light-colored at first, becoming brown when mature, 4.7–9.4 µm (x– = 6.4 µm, n = 20) wide. Stolons branched, hyaline to light brown, smooth, septate, 5.1–6.3 µm (x– = 5.6 µm, n = 15) in diameter. Rhizoids well developed, root-like, branched. Sporangiophores growing on stolons, erect or slightly bent, mostly unbranched or simply branched, smooth, single or 2–4 in whorls, monopodial or sympodial, 21.7–213.5 × 2.3–5.4 µm (x– = 112.8 × 4.1 µm, n = 15). Sporangia globose to pyriform, smooth, hyaline, deliquescent-walled, 16.2–37.0 × 14.0–31.1 µm (x– = 24.2 × 23.2 µm, n = 15), and with a septum 8.8–17.9 µm (x– = 13.9 µm, n = 15) below apophyses; the septum is not obvious when young. Apophyses distinct, funnel-shaped, 6.0–9.5 µm (x– = 8.5 µm, n = 15) high, 5.1–8.8 µm (x– = 6.2 µm, n = 15) wide at the base, and 8.4–13.9 µm (x– = 10.7 µm, n = 15) wide at the top, darker brown when old. Collars present, obvious. Columellae nearly conical, sometimes subspherical to hemispherical, 8.1–13.8 × 7.7–14.9 µm (x– = 10.6 × 10.9 µm, n = 15). Projections absent or present, hyaline, single. Sporangiospores hyaline, smooth, mostly oval, 2.1–3.8 × 1.8–2.7 µm (x– = 3.1 × 2.3 µm, n = 20). Chlamydospores absent. Zygospores not found.

Maximum growth temperature.

29 °C.

Additional specimen examined.

China • Yunnan Province, Yuxi City (23°57'28"N, 101°30'38"E, 2196.56 m), from soil, 5 Jul. 2024, X.Y. Ji and X.Y. Liu, living culture XG08666-10-2.

Notes.

In the molecular phylogeny, A. collariata was closely related to A. psychrophilia (Zhao et al. 2022b). Morphologically, the width of stolons in A. collariata was smaller than that in A. psychrophilia (5.1–6.3 µm vs. 5.5–11 µm). Furthermore, the A. collariata has smaller sporangiophores (2.1–3.8 × 1.8–2.7 µm vs. 3.8–5 × 2.2–3.5 µm). The length and width of sporangiophore were also smaller in A. collariata than those in A. psychrophilia (21.7–213.5 × 2.3–5.4 µm vs. 193–288 × 4.5–9 µm). And the A. psychrophilia has larger sporangia (20–50 μm vs. 16.2–37.0 × 14.0–31.1 μm). Zygospores were not observed in A. collariata.

Absidia hainanensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Fungal Names: FN 572258

Fig. 3

Type.

China • Hainan Province, Danzhou City, Tropical Botanical Garden (19°30'42"N, 109°30'3"E, 168.7 m), from soil, 26 Jun. 2024, X.Y. Ji and X.Y. Liu, holotype HMAS 353362, ex-holotype living culture CGMCC 3.28535 (=XG06908-1).

Figure 3.

Absidia hainanensis ex-holotype CGMCC 3.28535 a, b colonies on PDA (a obverse b reverse) c, d sporangia e–g columellae h rhizoids i sporangiospores; Scale bars: 10 µm (c–i).

Etymology.

The hainanensis (Lat.) refers to Hainan Province of China, where the type was collected.

Description.

Colonies on PDA at 26 °C for 5 days, reaching 75 mm in diameter, fast growing with a rate of 15 mm/d, at first white, becoming grayish-brown when old. Hyphae hyaline at first, becoming light brown when mature, 2.5–10.2 µm (x– = 5.2 µm, n = 20) in diameter. Rhizoids root-like, simply branched. Stolons hyaline, smooth, branched, 2.9–10.1 µm (x– = 6.1 µm, n = 15) in diameter. Sporangiophores erect or slightly bent, mostly unbranched or simply branched, smooth, monopodial or sympodial, single or 2–4 in whorls, 18.8–159.2 × 1.9–3.7 µm (x– = 75.4 × 2.8 µm, n = 15). Sporangia spherical to subspherical, smooth, hyaline, deliquescent-walled, 15.1–35.2 × 14.3–29.4 µm (x– = 24.2 × 22.1 µm, n = 15), and with a septum 11.5–24.2 µm (x– = 17.0 µm, n = 15) below apophyses. Apophyses obvious, funnel-shaped, 3.3–5.1 µm (x– = 4.6 µm, n = 15) high, 2.5–8.5 µm (x– = 4.3 µm, n = 15) wide at the base, and 7.5–17.4 µm (x– = 11.0 µm, n = 15) wide at the top, light brown, hyaline. Collars present. Columellae mostly oval, 3.9–13.9 × 8.6–19.5 µm (x– = 6.3 × 10.7 µm, n = 15). Projections absent or present, hyaline, single. Sporangiospores ovoid to cylindrical, smooth, hyaline, 3.1–3.7 × 1.9–2.5 µm (x– = 3.6 × 2.2 µm, n = 20). Chlamydospores absent. Zygospores not found.

Maximum growth temperature.

34 °C.

Additional specimen examined.

China • Hainan Province, Danzhou City (19°30'42"N, 109°30'3"E, 168.7 m), from soil, 26 June 2024, X.Y. Ji and X.Y. Liu, living culture XG06908-4.

Notes.

In the molecular phylogeny, A. hainanensis was closely related to A. oblongispora (Zong et al. 2021). Morphologically, the sporangiophores of A. hainanensis were at most four in whorls, while those of A. oblongispora were at most five in whorls. Additionally, the maximum length of the sporangiophores in A. oblongispora was significantly greater than that in A. hainanensis (300 µm vs. 159.2 µm). The sporangiospore size was smaller in A. hainanensis (3.1–3.7 × 1.9–2.5 μm vs. 3.5–4.5 × 2.5–8.5 μm). The A. hainanensis has wider columellae (8.6–19.5 µm vs. 8.5–16.5 µm). Physiologically, the maximum growth temperature of A. hainanensis was higher (34 °C vs. 32 °C).

Absidia pyriformis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Fungal Names: FN 572255

Fig. 4

Type.

China, Yunnan Province, Pu’er City, Mojiang Hani Autonomous County, Lianzhu Town (23°25'34"N, 101°40'58"E, 1338.32 m), from soil, 4 July 2024, X.Y. Ji and X.Y. Liu, holotype HMAS 353359, ex-holotype living culture CGMCC 3.28538 (=XG09540-14-1).

Figure 4.

Absidia pyriformis ex-holotype CGMCC 3.28538 a, b colonies on PDA (a obverse b reverse) c, d sporangia e–g columellae h rhizoids i sporangiospores; Scale bars: 10 µm (c–i).

Etymology.

The epithet pyriformis (Lat.) refers to the shape of the sporangia.

Description.

Colonies on PDA at 26 °C for 5 days, attaining 76 mm in diameter, moderately fast growing with a rate of 15.2 mm/d, white at first, gradually light gray, irregularly at reverse. Hyphae branched, hyaline at first, sometimes brownish when mature, aseptate when juvenile, septate with age, 3.6–15.2 µm (x– = 6.3 µm, n = 20) wide. Stolons branched, hyaline, smooth, septate, 4.5–11.9 µm (x– = 6.9 µm, n = 15) in diameter. Rhizoids well developed, root-like, branched, tapering at the end. Sporangiophores arising from stolons, erect or slightly bent, 2–5 in whorls, monopodial, mostly unbranched or simply branched, smooth, 21.7–279.8 × 1.4–7.4 µm (x– = 97.9 × 4.4 µm, n = 15), with one septum 11.5–26.8 µm (x– = 18.5 µm, n = 15) below apophyses. Sporangia are mostly pyriform, deliquescent-walled, smooth, multi-spored, colorless when young, brownish when old, 11.7–38.8 × 11.0–29.7 µm (x– = 27.9 × 22.1 µm, n = 15). Apophyses distinct, subhyaline, usually brownish when old, 5.0–8.4 µm (x– = 7.0 µm, n = 15) high, 2.4–6.1 µm (x– = 4.4 µm, n = 15) wide at the base, and 8.9–19.2 µm (x– = 12.5 µm, n = 15) wide at the top. Collars distinct. Columellae hemispherical, subglobose to globose, smooth, subhyaline or brownish, 8.8–21.4 × 16.7–20.7 µm (x– = 10.9 × 16.7 µm, n = 15). Projections at the apex, when smaller, with an oval projection. Sporangiospores hyaline, smooth, almost cylindrical, 3.2–4.5 × 1.7–2.8 µm (x– = 3.9 × 2.2 µm, n = 20). Chlamydospores absent. Zygospores absent.

Maximum growth temperature.

33 °C.

Additional specimen examined.

China • Yunnan Province, Pu’er City, from soil (23°25'34"N, 101°40'58"E, 1338.32 m), 4 July 2024, X.Y. Ji and X.Y. Liu, living culture XG09540-14-5.

Notes.

Phylogenetically, A. pyriformis was closely related to A. soli (Hurdeal et al. 2021). Compared with A. soli, the A. pyriformis presented a smaller sporangia size (11.7–38.8 × 11.0–29.7 µm vs. 16–51 × 15–45.5 µm), and the septum showed at a shorter distance from apophyses (11.5–26.8 µm vs. 21.5–37.5 µm); conversely, sporangiophores exhibited a larger size (3.2–4.5 µm vs. 1.7–2.8 µm), and columellae had a longer length (8.8–21.4 µm vs. 7.5–12.5 µm).

Absidia tardiva X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Fungal Names: FN 572254

Fig. 5

Type.

China, Yunnan Province, Yuxi County, Jinshan National Forest (23°38'15"N, 101°16'30"E, 2397.53 m), from soil, 14 May 2024, X.Y. Ji and X.Y. Liu, holotype HMAS 353358, ex-holotype living culture CGMCC 3.28537 (=XG08757-4).

Figure 5.

Absidia tardiva ex-holotype CGMCC 3.28537 a, b colonies on PDA (a obverse b reverse) c, e sporangia f, g a swelling on sporangiophores and hyphae h, i columellae j sporangiospores. Scale bars: 10 µm (c–j).

Etymology.

The epithet tardiva (Lat.) refers to this species growing more slowly than other strains.

Description.

Colonies on PDA at 26 °C for 4 days, reaching 41 mm in diameter, slow-growing with a rate of 10.25 mm/d; it begins white and gradually turns pale yellow to grayish-brown, irregular at reverse. Hyphae branched, hyaline at first, brownish when mature, 2.6–10.7 µm (x– = 5.6 µm, n = 20) in diameter, sometimes swollen. Stolons hyaline to brownish, smooth, branched, 3.7–8.4 µm (x– = 5.7 µm, n = 15) in diameter. Rhizoids not observed. Sporangiophores erect or slightly bent, single or 2–4 in whorls, unbranched or simply branched, monopodial or sympodial, with a septum 10.6–23.1 µm (x– = 13.8 µm, n = 15) below apophyses, 7.9–141.9 × 1.9–7.4 µm (x– = 70.7 × 4.2 µm, n = 15), sometimes with a swelling beneath sporangia. Sporangia subspherical to spherical, smooth, multi-spored, 12.9–48.3 × 9.3–34 µm (x– = 30.3 × 22.2 µm, n = 15), deliquescent-walled. Apophyses distinct, subhyaline, small, slightly pigmented, 3.2–10.4 µm (x– = 5.2 µm, n = 15) high, 2.9–7.2 µm (x– = 4.4 µm, n = 15) wide at the base, and 8.0–20.5 µm (x– = 13.2 µm, n = 15) wide at the top. Collars absent. Columellae hemispherical, subhyaline to hyaline, smooth, 2.9–13.8 × 4.9–16.3 µm (x– = 8.7 × 9.8 µm, n = 15). Projections present, shaped like a grain of rice. Sporangiospores variously shaped, mostly ovoid; a few are cylindrical or subglobose, smooth, hyaline, 3.4–4.6 × 2.1–2.8 µm (x– = 3.9 × 2.3 µm, n = 20). Chlamydospores absent. Zygospores not observed.

Maximum growth temperature.

27 °C.

Additional specimen examined.

China • Yunnan Province, Yuxi County, from soil (23°38'15"N, 101°16'30"E, 2397.53 m), 14 May 2024, X.-Y. Ji and X.-Y. Liu, living culture XG08757-6.

Notes.

Phylogenetic analysis of five genes showed that A. tardiva was closely related to A. psychrophilia (Zhao et al. 2022b). Morphologically, the sporangia shape of A. psychrophilia was pyriform, while the sporangia of A. tardiva were hemispherical to spherical in shape. And the distance between the septum and apophysis was shorter in A. psychrophilia than in A. tardiva (10–17 µm vs. 10.6–23.1 µm). The columellae of A. psychrophilia were larger than those of A. tardiva (6.5–30 μm in diameter vs. 2.9–13.8 × 4.9–16.3 μm). The overall size of spores in A. psychrophilia was slightly larger than that in A. tardiva (long: 3.8–5 × 2.2–3.5 µm vs. wide: 3.4–4.6 × 2.1–2.8 µm); the shape of sporangiospores in A. psychrophilia was cylindrical, whereas the shape of spores in A. tardiva was oval.

Absidia tibetensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Fungal Names: FN 572257

Fig. 6

Type.

China • Tibet, Xigaze City, Yadong Country (27°21'53"N, 88°58'26"E, 2827 m), from soil, 1 Oct 2024, X.Y. Ji and X.Y. Liu, holotype HMAS 353361, ex-holotype living culture CGMCC 3.28534 (=XG00415-1).

Figure 6.

Absidia tibetensis ex-holotype CGMCC 3.28534 a, b colonies on PDA (a obverse b reverse) c, d sporangia e–g columellae h, i a swelling on sporangiophores and hyphae j rhizoids k sporangiospores. Scale bars: 10 μm (c–k).

Etymology.

The tibetensis (Lat.) refers to the Tibet Autonomous Region of China, where the type was collected.

Description.

Colonies on PDA at 26 °C for 5 days, reaching 53 mm in diameter, slow-growing with a rate of 10.6 mm/d, white at first and gradually turning to light brown; the reverse side of the colony resembles a petal-shaped, regularly at reverse. Rhizoids root-like, always branched, with a septum at the top. Hyphae hyaline to slightly gray, 5.0–10.0 µm (x– = 7.1 µm, n = 20) in diameter, sometimes ampulliform-shaped swollen. Stolons hyaline, slightly brownish, branched, smooth, 3.2–11.0 µm (x– = 6.0 µm, n = 15) in diameter. Sporangiophores erect or slightly bent, unbranched or simple branched, smooth, single or 2–5 in whorls, monopodial or sympodial, 14.7–144.0 × 2.5–5.7 µm (x– = 78.2 × 4.0 µm, n = 15), sometimes with a swelling beneath sporangia. Sporangia globose to pyriform, smooth, multi-spored, deliquescent-walled, 11.0–30.2 × 11.1–26.6 µm (x– = 21.5 × 17.1 µm, n = 15), and with a septum 8.4–20.0 µm (x– = 15.6 µm, n = 15) below apophyses. Apophyses obvious, funnel-shaped, gradually widening from the base to the top, 2.5–9.6 µm (x– = 6.7 µm, n = 15) high, 3.2–8.3 µm (x– = 4.2 µm, n = 15) wide at the base, and 7.4–19.0 µm (x– = 11.2 µm, n = 15) wide at the top, hyaline, light brown. Collars absent or present. Columellae conical, nearly globose, occasionally oval, 8.5–19.9 × 10.1–16.5 µm (x– = 11.3 × 11.8 µm, n = 15). Projections present or absent, hyaline when present, needle-pointed. Sporangiospores smooth, hyaline, mostly oval, 2.6–3.9 × 1.6–2.4 µm (x– = 3.6 × 2.1 µm, n = 20). Chlamydospores absent. Zygospores not found.

Maximum growth temperature.

30 °C.

Additional specimen examined.

China • Tibet, Xigaze City, Yadong County (27°21'53"N, 88°58'26"E, 2827 m), from soil, 1 October 2024, X.Y. Ji and X.Y. Liu, living culture XG00415-3.

Notes.

In the molecular phylogeny, A. tibetensis was closely related to A. sichuanensis (Zhao et al. 2022b). Morphologically, the maximum width of the hyphae in A. yunnanensis was greater than that in A. tibetensis (15.5 µm vs. 10.0 µm). Apophyses had a wider base width and top width in A. tibetensis (3.2–8.3 × 7.4–19.0 μm vs. 3.0–5.0 × 5.5–12.0 μm). The sporangiophore size was smaller in A. tibetensis (2.6–3.9 × 1.6–2.4 μm vs. 3.0–4.5 × 2.0–2.5 μm). The swelling on sporangiophores and hyphae was observed in A. tibetensis. The collars were not observed in A. sichuanensis. Physiologically, the maximum growth temperature of A. tibetensis was higher (30 °C vs. 28 °C).

Discussion

Absidia is widely distributed. Some soil samples in Yunnan Province, Tibet Autonomous Region, and Hainan Province were investigated in this study. The cities Pu’er and Yuxi in Yunnan Province have a subtropical monsoon climate with complex terrain, mild and humid climate, and abundant precipitation. The climatic environment is conducive to the growth of various microorganisms. Yadong County of the Tibet Autonomous Region has a plateau and mountainous climate, with significant seasonal changes and extreme weather phenomena. Danzhou City, in Hainan Province, has a tropical humid monsoon climate with abundant sunshine and abundant rainfall. Five new species of the genus Absidia were discovered in these places (Wu et al. 2020; Wang et al. 2021; Zhao et al. 2022b; Jiang et al. 2024).

Based on morphology, growth temperature dynamics, and molecular phylogenetic analyses, five novel species were identified in the genus Absidia, namely A. collariata sp. nov., A. hainanensis sp. nov., A. pyriformis sp. nov., A. tardiva sp. nov., and A. tibetensis sp. nov. In this study, phylogenetic analysis was performed for these five novel species based on five loci, namely ITS, LSU, TEF1α, Act, and SSU. By analyzing these data, strong support was obtained for the clades of these species (A. tardiva 100% MLBV and 1.00 BIPP; A. pyriformis 100% MLBV and 1.00 BIPP; A. collariata 100% MLBV and 1.00 BIPP; A. tibetensis 100% MLBV and 1.00 BIPP; A. hainanensis 100% MLBV and 1.00 BIPP; Fig. 1). At the same time, in terms of morphological structure and physiology, we also found some differences between these five newly discovered species and their closely related species (Benny 2008; Zheng et al. 2009; Hurdeal et al. 2021; Urquhart and Idnurm 2021). They have great differences in sporangiospore size, stolon width, sporangia size, base and top width of the apophyses, and so on (Ellis and Hesseltine 1965; Davoust and Persson 1992; Lima et al. 2020). Besides, the maximum growth temperature between them is also different. These differences laid the foundation for the identification of the five novel species.

Table 2.

Download as

CSV

XLSX

GenBank accession numbers of sequences used in this study.

Species	Strains	GenBank accession numbers
Species	Strains	ITS	LSU	TEF-1α	Act	SSU
Absidia abundans	XY09265	ON074697	ON074681	NA	NA	NA
A. abundans	CGMCC 3.16255*	NR_182590	ON074683	NA	NA	NA
A. abundans	XY09274	ON074696	ON074682	NA	NA	NA
A. aguabelensis	URM 8213*	NR_189383	NG_241934	NA	NA	NA
A. alpina	CGMCC 3.16104	OL678133	NA	NA	NA	NA
A. ampullacea	CGMCC 3.16054	MZ354138	MZ350132	NA	NA	NA
A. anomala	CBS 125.68*	MH859085	MH870799	NA	NA	NA
A. anomala	FSU5798	EF030523	NA	NA	EF030535	NA
A. biappendiculata	CBS 187.64	MZ354153	MZ350147	MZ357420	MZ357438	NA
A. bonitoensis	URM 7889*	MN977786	MN977805	NA	NA	NA
A. brunnea	CGMCC 3.16055*	MZ354139	MZ350133	MZ357403	MZ357421	NA
A. caatinguensis	URM 7156*	NR_154704	NG_058582	NA	NA	NA
A. caerulea	XY00608	OL620081	NA	NA	NA	NA
A. caerulea	XY00729	OL620082	NA	NA	NA	NA
A. caerulea	CBS101.36	MH855718	MH867230	NA	NA	NA
A. caerulea	FSU767	AY944870	NA	NA	NA	NA
A. californica	CBS 314.78	JN205816	MH872902	NA	NA	NA
A. californica	FSU4748	AY944873	EU736301	EU736247	EU736224	EU736274
A. californica	FSU4747	AY944872	EU736300	EU736246	AY944758	EU736273
A. chinensis	CGMCC 3.16057	MZ354141	MZ350135	NA	MZ357422	NA
A. chinensis	CGMCC 3.16056*	MZ354140	MZ350134	NA	NA	NA
A. cinerea	CGMCC 3.16062	MZ354146	MZ350140	MZ357407	MZ357427	NA
*A. collariata*	CGMCC 3.28536*	PQ610533	PQ605104	PQ613269	PQ613279	PQ605114
*A. collariata*	XG08666-10-2	PQ610534	PQ605105	PQ613270	PQ613280	PQ605115
A. cornuta	URM 6100*	NR_172976	MN625255	NA	NA	NA
A. cuneospora	CBS 101.59*	MH857828	MH869361	NA	NA	NA
A. cylindrospora	CBS 100.08	JN205822	JN206588	NA	NA	NA
A. digitula	CGMCC 3.16058*	MZ354142	MZ350136	MZ357404	MZ357423	NA
A. edaphica	MFLUCC 20-0088	NR_172305	NG_075367	NA	MT410739	NG_074951
A. frigida	CGMCC 3.16201*	NR_182565	OM030223	NA	NA	NA
A. fusca	CBS 102.35*	NR_103625	NG_058552	NA	NA	NA
A. gemella	CGMCC 3.16202*	OM108488	OM030224	NA	NA	NA
A. glauca	CBS 129233	MH865253	MH876693	NA	NA	NA
A. glauca	CBS 101.08*	MH854573	MH866105	NA	NA	NA
A. glauca	FSU660	AY944879	EU736302	EU736248	EU736225	EU736275
A. globospora	CGMCC 3.16031*	NR_189829	MW671544	MZ357412	MZ357431	NA
A. globospora	CGMCC 3.16035	MW671538	MW671545	MZ357413	MZ357432	NA
A. globospora	CGMCC 3.16036	MW671539	MW671546	MZ357414	MZ357433	NA
*A. hainanensis*	CGMCC 3.28535*	PQ610537	PQ605108	PQ613273	PQ613283	PQ605118
*A. hainanensis*	XG06908-4	PQ610538	PQ605109	PQ613274	PQ613284	PQ605119
A. heterospora	SHTH021	JN942683	JN982936	NA	NA	JQ004928
A. heterospora	CBS101.29*	JN206595.1	MH866483.1	NA	NA	NA
A. jiangxiensis	CGMCC 3.16105*	OL678134	PP780377	PP790569	PP790577	PP779719
A. jindoensis	CNUFC-PTI1-1	MF926622	MF926616	MF926513	MF926510	MF926626
A. koreana	EML-IFS45-1*	KR030062	KR030056	KR030060	KR030058	KT321298
A. koreana	XY00816	OL620083	ON123771	NA	NA	NA
A. koreana	XY00596	OL620084	NA	NA	NA	NA
A. lobata	CGMCC 3.16256	ON074690	ON074679	NA	NA	NA
A. longissima	CGMCC 3.16203*	NR_182566	OM030225	NA	NA	NA
A. macrospora	FSU4746	AY944882	EU736303	EU736249	AY944760	EU736276
A. macrospora	CBS 697.68*	HM849704.1	NA	NA	NA	NA
A. medulla	CGMCC 3.16034	NR_189832	MW671549	MZ357417	MZ357436	NA
A. montepascoalis	URM 8218	NR_172995	NA	NA	NA	NA
A. multispora	URM 8210*	MN953780	MN953782	NA	NA	NA
A. nigra	CBS 127.68*	NR_173068	MZ350146	MZ357419	MZ357437	NA
A. nigra	CGMCC 3.16059	MZ354143	MZ350137	MZ357405	MZ357424	NA
A. nigra	CGMCC 3.16060	MZ354144	MZ350138	MZ357406	MZ357425	NA
A. oblongispora	CGMCC 3.16061	MZ354145	MZ350139	NA	MZ357426	NA
A. ovalispora	CGMCC 3.16019	NR_176748	MW264131	NA	NA	NA
A. panacisoli	SYPF 7183*	MF522181	MF522180	MF624251	NA	MF522179
A. pararepens	XY00631	OL620085	ON123774	NA	NA	NA
A. pararepens	XY00615	OL620086	NA	NA	NA	NA
A. pararepens	XY05899	OL620087	NA	NA	NA	NA
A. pararepens	CCF 6352	MT193669	MT192308	NA	NA	NA
A. pernambucoensis	URM < BRA > 7219	MN635568	MN635569	NA	NA	NA
A. pseudocylindrospora	EML-FSDY6-2	KU923817	KU923814	NA	KU923815	KU923819
A. psychrophilia	FSU4745	AY944874	EU736306	EU736252	AY944762	EU736279
A. purpurea	CGMCC 3.16106	OL678135	NA	NA	NA	NA
*A. pyriformis*	CGMCC 3.28538*	PQ610531	PQ605102	PQ613267	PQ613277	PQ605112
*A. pyriformis*	XG09540-14-5	PQ610532	PQ605103	PQ613268	PQ613278	PQ605113
A. radiata	CGMCC 3.16257	ON074698	ON074684	NA	NA	NA
A. radiata	XY09330-1	ON074699	ON074685	NA	NA	NA
A. repens	CBS 115583*	NR_103624	NG_058551	NA	NA	NA
A. saloaensis	URM 8209*	MN953781	MN953783	NA	NA	NA
A. sichuanensis	CGMCC 3.16258*	NR_182589	ON074688	NA	NA	NA
A. soli	MFLU-20-0414*	MT396373	MT393988	NA	NA	MT394049
A. spinosa	FSU551	AY944887	EU736307	EU736253	EU736227	EU736280
A. stercoraria	EML-DG8-1*	KU168828	KT921998	KT922002	KT922000	NG_065640
A. sympodialis	CGMCC 3.16063*	MZ354147	MZ350141	NA	NA	NA
A. sympodialis	CGMCC 3.16064	MZ354148	MZ350142	MZ357408	NA	NA
*A. tardiva*	CGMCC 3.28537*	PQ610529	PQ605100	PQ613265	PQ613275	PQ605110
*A. tardiva*	XG08757-6	PQ610530	PQ605101	PQ613266	PQ613276	PQ605111
A. terrestris	FMR 14989*	LT795003	LT795005	NA	NA	NA
*A. tibetensis*	CGMCC 3.28534*	PQ610535	PQ605106	PQ613271	PQ613281	PQ605116
*A. tibetensis*	XG00415-3	PQ610536	PQ605107	PQ613272	PQ613282	PQ605117
A. turgida	CGMCC 3.16032*	NR_189830	NG_241931	MZ357415	MZ357434	NA
A. varians	CGMCC 3.16065*	MZ354149	MZ350143	MZ357409	MZ357428	NA
A. virescens	CGMCC 3.16066*	MZ354150	MZ350144	MZ357410	MZ357429	NA
A. virescens	CGMCC 3.16067	MZ354151	MZ350145	MZ357411	MZ357430	NA
A. xinjiangensis	CGMCC 3.16107*	OL678136	NA	NA	NA	NA
A. yunnanensis	XY09528	ON074701	ON074686	NA	NA	NA
A. yunnanensis	CGMCC 3.16259*	NR_182591	NG_149054	NA	NA	NA
A. zonata	CGMCC 3.16033*	NR_189831	MW671548	MZ357416	MZ357435	NA
A. zygospora	RSPG 214	KC478527	NA	NA	NA	NA
A. zygospora	ANG28	DQ914420	NA	NA	NA	NA
A. zygospora	MFLUCC 23-0016*	OR104965	OR104992	NA	NA	NA
Cunninghamella blakesleeana	CBS 782.68	JN205869	MH870950	NA	NA	NA
C. blakesleeana	CBS 133.27*	JN205865.1	MH866397.1	KJ156479.1	NA	NA
C. elegans	CBS 167.53	MH857146	HM849700	NA	NA	NA
C. elegans	CBS 160.28*	AF254928.1	NA	KJ156470.1	NA	NA

Absidia has important physiological functions, which are manifested in many aspects, such as ecology, industry, medicine, and so on. Ecologically, it helps in the decomposition of organic matter, which is essential for nutrient cycling. Industrially, it is used for the biotransformation of various natural products. However, it also has a downside: some species of the genus Absidia that can grow at 37 °C are opportunistic pathogens that cause diseases in humans and animals (Zhao et al. 2022b; Tao et al. 2024). Therefore, there is still great research value in the physiological function of the genus Absidia. As of 11 November 2024, the Global Biodiversity Information Facility (GBIF) (https://www.gbif.org/, accessed 11 November 2024) contains 8,496 globally reported georeferenced records of the genus Absidia species. The genus is most widely distributed in Europe and least in Antarctica (Ellis and Hesseltine 1965; Tran et al. 2019; Liu et al. 2021). In this study, new species of Absidia were found in regions with different climates in Yunnan, Hainan, and Tibet in China, which further revealed the species diversity of Absidia in different regions.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the National Natural Science Foundation of China (Nos. 32170012, 32470004, 32300011). 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

X.Y. Ji took charge of the drawings, DNA sequencing, and data analyses and drafted the paper; Z.Y. Ding, H. Zhao, and S. Wang collected samples and isolated cultures; Y. Nie and B. Huang revised the paper; and X.Y. Liu revised the paper and provided funding.

Author ORCIDs

Xin-Yu Ji https://orcid.org/0009-0000-4121-9103

Zi-Ying Ding https://orcid.org/0009-0003-1618-5740

Yong Nie https://orcid.org/0000-0001-8964-1661

Heng Zhao https://orcid.org/0000-0003-2938-5613

Shi Wang https://orcid.org/0000-0002-7376-7638

Bo Huang https://orcid.org/0000-0001-6032-7396

Xiao-Yong Liu https://orcid.org/0000-0002-8808-010X

Data availability

The sequences of this study have been submitted to the NCBI database (https://www.ncbi.nlm.nih.gov/, accessed November 15, 2024) with accession numbers shown in Table 2.

References

Benny GL (2008) Methods used by Dr. RK Benjamin, and other mycologists, to isolate zygomycetes. Aliso 26(1): 37–61. https://doi.org/10.5642/aliso.20082601.08

Constantinides J, Misra A, Nassab R, Wilson Y (2008) Absidia corymbifera fungal infection in burns: A case report and review of the literature. Journal of Burn Care & Research; Official Publication of the American Burn Association 29(2): 416–419. https://doi.org/10.1097/BCR.0b013e318166da78

Cordeiro TRL, Nguyen TTT, Lima DX, Da Silva SBG, De Lima CF, Leitão JDA, Gurgel LMS, Lee HB, De Azevedo Santiago ALCM (2020) Two new species of the industrially relevant genus Absidia (Mucorales) from soil of the Brazilian Atlantic Forest. Acta Botanica Brasílica 34(3): 549–558. https://doi.org/10.1590/0102-33062020abb0040

Corry JEL, Curtis GDW, Baird RM (1995) Rose bengal chloramphenicol (RBC) agar, in Progress in Industrial Microbiology. Elsevier, 431–433. https://doi.org/10.1016/S0079-6352(05)80079-7

Crous P, Wingfield M, Chooi Y, Gilchrist C, Lacey E, Pitt J, Roets F, Swart W, Cano J, Valenzuela-Lopez N, Hubka V, Shivas R, Stchigel AM, Holdom D, Jurjević Ž, Kachalkin A, Lebel T, Lock C, Martín M, Groenewald JZ (2020) Fungal Planet description sheets: 1042–1111. Persoonia 44(1): 301–459. https://doi.org/10.3767/persoonia.2020.44.11

Davoust N, Persson A (1992) Effects of growth morphology and time of harvesting on the chitosan yield of Absidia repens. Applied Microbiology and Biotechnology 37: 572–575. https://doi.org/10.1007/BF00240727

Ding ZY, Ji XY, Tao MF, Liu WX, Jiang Y, Zhao H, Meng Z, Liu XY (2025) (in review) Unveiling species diversity within early-diverging fungi from China IV: Four new species of Absidia (Cunninghamellaceae, Mucoromycota). MycoKeys. https://doi.org/10.20944/preprints202504.1116.v1

Doyle JJ, Doyle JL, Brown AHD (1990) Chloroplast DNA phylogenetic affinities of newly described species in Glycine (Leguminosae: Phaseoleae). Systematic Botany 15(3): 466–471. https://doi.org/10.2307/2419362

Ellis J, Hesseltine C (1965) The genus Absidia: Globose-spored species. Mycologia 57(2): 222–235. https://doi.org/10.1080/00275514.1965.12018205

Ellis J, Hesseltine C (1967) Species of Absidia with ovoid sporangiospores. II. Sabouraudia 5(2): 59–77. https://doi.org/10.1080/00362176785190111

Guan QX, Li YF, Zhao CL (2021) Morphological and phylogenetic evidence for recognition of two new species of Hyphoderma (Basidiomycota) from southern China, with a key to all Chinese Hyphoderma. MycoKeys 83: 145–160. https://doi.org/10.3897/mycokeys.83.69909

Hoffmann K (2010) Identification of the genus Absidia (Mucorales, Zygomycetes): A comprehensive taxonomic revision. Molecular Identification of Fungi, 439–460. https://doi.org/10.1007/978-3-642-05042-8_19

Hoffmann K, Voigt K (2009) Absidia parricida plays a dominant role in biotrophic fusion parasitism among mucoralean fungi (Zygomycetes): Lentamyces, a new genus for A. parricida and A. zychae. Plant Biology 11: 537–554. https://doi.org/10.1111/j.1438-8677.2008.00145.x

Hoffmann K, Discher S, Voigt K (2007) Revision of the genus Absidia (Mucorales, Zygomycetes) based on physiological, phylogenetic, and morphological characters; thermotolerant Absidia spp. form a coherent group, Mycocladiaceae fam. nov. Mycological Research 111: 1169–1183. https://doi.org/10.1016/j.mycres.2007.07.002

Hurdeal VG, Gentekaki E, Lee HB, Jeewon R, Hyde KD, Tibpromma S, Mortimer PE, Xu J (2021) Mucoralean fungi in Thailand: Novel species of Absidia from tropical forest soil. Cryptogamie. Mycologie 42(4): 39–61. https://doi.org/10.5252/cryptogamie-mycologie2021v42a4

Hurdeal VG, Jones EBG, Gentekaki E (2023) Absidia zygospore (Mucoromycetes), a new species from Nan province, Thailand. Studies in Fungi 8: 15. https://doi.org/10.48130/SIF-2023-0015

Jaklitsch WM, Komon M, Kubicek CP, Druzhinina IS (2005) Hypocrea voglmayrii sp. nov. from the Austrian Alps represents a new phylogenetic clade in Hypocrea/Trichoderma. Mycologia 97(6): 1365–1378. https://doi.org/10.1080/15572536.2006.11832743

Jiang Y, Zhang ZX, Zhang J, Wang S, Zhang XG (2024) Morphological and phylogenetic analyses reveal three new species of Phyllosticta (Botryosphaeriales, Phyllostictaceae) in China. Journal of Fungi (Basel, Switzerland) 10(1): 7. https://doi.org/10.3390/jof10010007

Kristanti RA, Zubir MMFA, Hadibarata T (2016) Biotransformation studies of cresolred by Absidia spinosa M15. Journal of Environmental Management 172: 107–111. https://doi.org/10.1016/j.jenvman.2015.11.017

Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870–1874. https://doi.org/10.1093/molbev/msw054

Larsson A (2014) AliView: A fast and lightweight alignment viewer and editor for large datasets. Bioinformatics (Oxford, England) 30(22): 3276–3278. https://doi.org/10.1093/bioinformatics/btu531

Lima DX, Cordeiro TRL, De Souza CAF, De Oliveira RJV, Lee HB, De Souza-Motta CM, Santiago A (2020) Morphological and molecular evidence for two new species of Absidia from Neotropic soil. Phytotaxa 446: 61–71. https://doi.org/10.11646/phytotaxa.446.1.8

Liu P, Zou Y, Li S, Stephenson SL, Wang Q, Li Y (2019) Two new species of dictyostelid cellular slime molds in high-elevation habitats on the Qinghai-Tibet Plateau, China. Scientific Reports 9: 5. https://doi.org/10.1038/s41598-018-37896-7

Liu XF, Tibpromma S, Xu JC, Kumla J, Karunarathna SC, Zhao CL (2021) Taxonomy and phylogeny reveal two new potential edible ectomycorrhizal mushrooms of Thelephora from East Asia. Diversity 13(12): 646. https://doi.org/10.3390/d13120646

Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 Gateway Computing Environments Workshop, 1–8. https://doi.org/10.1109/GCE.2010.5676129

Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300

Nie Y, Cai Y, Gao Y, Yu DS, Wang ZM, Liu XY, Huang B (2020a) Three new species of Conidiobolus sensu stricto from plant debris in eastern China. MycoKeys 73: 133–149. https://doi.org/10.3897/mycokeys.73.56905

Nie Y, Yu DS, Wang CF, Liu XY, Huang B (2020b) A taxonomic revision of the genus Conidiobolus (Ancylistaceae, Entomophthorales): Four clades including three new genera. MycoKeys 66: 55–81. https://doi.org/10.3897/mycokeys.66.46575

Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029

Stamatakis A (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics (Oxford, England) 30(9): 1312–1313. https://doi.org/10.1093/bioinformatics/btu033

Tao MF, Ding ZY, Wang YX, Zhang ZX, Zhao H, Meng Z, Liu XY (2024) Unveiling species diversity within early-diverging fungi from China II: Three new species of Absidia (Cunninghamellaceae,Mucoromycota) from Hainan Province. MycoKeys 110: 255–272. https://doi.org/10.3897/mycokeys.110.129120

Tieghem P (1878) Troisième mémoire sur les Mucorinées. Annales des Sciences Naturelles; Botanique 4: 312–399.

Tran NT, Miles AK, Dietzgen RG, Drenth A (2019) Phyllosticta capitalensis and P. paracapitalensis are endophytic fungi that show potential to inhibit pathogenic P. citricarpa on Citrus australas. Plant Pathology 48: 281–296. https://doi.org/10.1007/s13313-019-00628-0

Urquhart AS, Idnurm A (2021) Absidia healeyae: A new species of Absidia (Mucorales) isolated from Victoria, Australia. Mycoscience 62(5): 331–335. https://doi.org/10.47371/mycosci.2021.06.001

Voigt K, Wöstemeyer J (2000) Reliable amplification of actin genes facilitates deep-level phylogeny. Microbiological Research 155(3): 179–195. https://doi.org/10.1016/S0944-5013(00)80031-2

Wang K, Chen SL, Dai YC, Jia ZF, Li TH, Liu TZ, Phurbu D, Mamut R, Sun GY, Bau T, Wei SL, Yang ZL, Yuan HS, Zhang XG, Cai L (2021) Overview of China’s nomenclature novelties of fungi in the new century (2000–2020). Junwu Xuebao 40(4): 822–833. https://doi.org/10.13346/j.mycosystema.210064

Wang S, Liu X, Xiong C, Gao S, Xu W, Zhao L, Song C, Liu X, James TY, Li Z, Zhang X (2023) ASF1 regulates asexual and sexual reproduction in Stemphylium eturmiu num by DJ-1 stimulation of the PI3K/AKT signaling pathway. Fungal Diversity 123(1): 159–176. https://doi.org/10.1007/s13225-023-00528-1

Wang YX, Zhao H, Jiang Y, Liu XY, Tao MF, Liu XY (2024) Unveiling species diversity within early-diverging fungi from China III: Six new species and a new record of Gongronella (Cunninghamellaceae, Mucoromycota). MycoKeys 110: 287–317. https://doi.org/10.3897/mycokeys.110.130260

White T, Bruns T, Lee S, Taylor J, Innis M, Gelfand D, Sninsky J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics, in PCR proto cols: a guide to methods and applications, 315–322. https://doi.org/10.1016/B978 0-12-372180-8.50042-1

Wu F, Yuan HS, Zhou LW, Yuan Y, Cui BK, Dai YC (2020) Polypore diversity in South China. Junwu Xuebao 39(4): 653–682. https://doi.org/10.13346/j.mycosystema.200087

Zhang TY, Yu Y, Zhu H, Yang SZ, Yang TM, Zhang MY, Zhang YX (2018) Absidia panacisoli sp. nov., isolated from rhizosphere of Panax notoginseng. International Journal of Systematic and Evolutionary Microbiology 68(8): 2468–2472. https://doi.org/10.1099/ijsem.0.002857

Zhang ZX, Liu RY, Liu SB, Mu TC, Zhang XG, Xia JW (2022) Morphological and phylogenetic analyses reveal two new species of Sporocadaceae from Hainan, China. MycoKeys 88: 171–192. https://doi.org/10.3897/mycokeys.88.82229

Zhao H, Lv ML, Liu Z, Zhang MZ, Wang YN, Ju X, Song Z, Ren LY, Jia BS, Qiao M, Liu XY (2021a) High-yield oleaginous fungi and high-value microbial lipid resources from Mucoromycota. BioEnergy Research 14: 1196–1206. https://doi.org/10.1007/s12155-020-10219-3

Zhao H, Zhu J, Zong TK, Liu XL, Ren LY, Lin Q, Qiao M, Nie Y, Zhang ZD, Liu XY (2021b) Two new species in the family Cunninghamellaceae from China. Mycobiology 49(2): 142–150. https://doi.org/10.1080/12298093.2021.1904555

Zhao H, Nie Y, Zong TK, Dai YC, Liu XY (2022a) Three new species of Absidia (Mucoromycota) from China based on phylogeny, morphology and physiology. Diversity 14(2): 132. https://doi.org/10.3390/d14020132

Zhao H, Nie Y, Zong TK, Wang YJ, Wang M, Dai YC, Liu XY (2022b) Species diversity and ecological habitat of Absidia (Cunninghamellaceae, Mucorales) with emphasis on five new species from forest and grassland soil in China. Journal of Fungi (Basel, Switzerland) 8(5): 471. https://doi.org/10.3390/jof8050471

Zhao H, Nie Y, Zong TK, Wang K, Lv ML, Cui YJ, Tohtirjap A, Chen JJ, Zhao CL, Wu F, Cui BK, Yuan Y, Dai YC, Liu XY (2023) Species diversity, updated classification and divergence times of the phylum Mucoromycota. Fungi Diversity 123: 49–157. https://doi.org/10.1007/s13225-023-00525-4

Zhao H, Nie Y, Huang B, Liu XY (2024) Unveiling species diversity within early-diverging fungi from China I: Three new species of Backusella (Backusellaceae, Mucoromycota). MycoKeys 109: 285–304. https://doi.org/10.3897/mycokeys.109.126029

Zheng RY, Liu XY, Li RY (2009) More Rhizomucor causing human mucormycosis from China: R. chlamydosporus sp. nov. Sydowia 61(1): 135–147.

Zong TK, Zhao H, Liu XL, Ren LY, Zhao CL, Liu XY (2021) Taxonomy and phylogeny of four new species in Absidia (Cunninghamellaceae, Mucorales) from China. Frontiers in Microbiology 12: 2181. https://doi.org/10.3389/fmicb.2021.677836

Zou Y, Hou J, Guo S, Li Z, Stephenson SL, Pavlov IN, Liu P, Li Y (2022) Diversity of dictyostelid cellular slime molds, including two species new to science, in forest soils of Changbai Mountain, China. Microbiology Spectrum 10: 5. https://doi.org/10.1128/spectrum.02402-22

﻿Abstract

Key words:

﻿Introduction

﻿Materials and methods

﻿Isolation and morphology

﻿DNA extraction, PCR amplification, and sequencing

﻿Phylogenetic analyses

﻿Results

﻿Molecular phylogeny

﻿Taxonomy

﻿ Absidia collariata X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Type.

Etymology.

Description.

Maximum growth temperature.

Additional specimen examined.

Notes.

﻿ Absidia hainanensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Type.

Etymology.

Description.

Maximum growth temperature.

Additional specimen examined.

Notes.

﻿ Absidia pyriformis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Type.

Etymology.

Description.

Maximum growth temperature.

Additional specimen examined.

Notes.

﻿ Absidia tardiva X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Type.

Etymology.

Description.

Maximum growth temperature.

Additional specimen examined.

Notes.

﻿ Absidia tibetensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Type.

Etymology.

Description.

Maximum growth temperature.

Additional specimen examined.

Notes.

﻿Discussion

﻿Additional information

Conflict of interest

Ethical statement

Funding

Author contributions

Author ORCIDs

Data availability

﻿References

Abstract

Introduction

Materials and methods

Isolation and morphology

DNA extraction, PCR amplification, and sequencing

Phylogenetic analyses

Results

Molecular phylogeny

Taxonomy

Absidia collariata X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Absidia hainanensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Absidia pyriformis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Absidia tardiva X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Absidia tibetensis X.Y. Ji, H. Zhao & X.Y. Liu, sp. nov.

Discussion

Additional information

References