A phylogenetic assessment of Akanthomyces sensu lato in Cordycipitaceae (Hypocreales, Sordariomycetes): introduction of new genera, and the resurrection of Lecanicillium

A Khonsanit; D Thanakitpipattana; S Mongkolsamrit; N Kobmoo; N Phosrithong; RA Samson; PW Crous; JJ Luangsa-ard

doi:10.3114/fuse.2024.14.17

. 2024 Aug 2;14:271–305. doi: 10.3114/fuse.2024.14.17

A phylogenetic assessment of Akanthomyces sensu lato in Cordycipitaceae (Hypocreales, Sordariomycetes): introduction of new genera, and the resurrection of Lecanicillium

A Khonsanit ¹, D Thanakitpipattana ¹, S Mongkolsamrit ¹, N Kobmoo ¹, N Phosrithong ², RA Samson ³, PW Crous ^3,^4,⁵, JJ Luangsa-ard ^1,^*

PMCID: PMC11736260 PMID: 39830296

Abstract

During entomopathogenic fungal surveys conducted in Thailand, 15 specimens tentatively classified under Akanthomyces sensu lato were identified. To gain a comprehensive understanding of their taxonomy, molecular phylogenies using combined LSU, TEF1, RPB1, and RPB2 sequence data, together with morphological examination of several Akanthomyces spp. from previous studies were conducted. The analyses revealed distinct clades representing independent lineages within the Cordycipitaceae. These clades were further characterized by different asexual morph types and the respective hosts they parasitize. In this context, we resurrected the genus Lecanicillium to accommodate 12 known species previously classified under Akanthomyces sensu lato, found on diverse hosts. We propose four new genera – Corniculantispora, Corpulentispora, Zarea, and Zouia – from species previously identified as Lecanicillium. Notably, certain Akanthomyces species associated with spiders and parasitic on Ophiocordyceps sinensis were reclassified into the new genera Arachnidicola and Kanoksria, respectively. Moreover, we introduce four novel species in Akanthomyces sensu stricto found across a diverse range of moth families: Ak. buriramensis, Ak. fusiformis, Ak. niveus, and Ak. phariformis. Additionally, we provide descriptions and illustrations of the sexual morph linked to Ak. laosensis and Ak. pseudonoctuidarum, along with a second type of synnemata seen in Ak. noctuidarum and Ak. pseudonoctuidarum. To assist with their identification, keys to the genera Akanthomyces, Arachnidicola, and Lecanicillium are provided, but should not be used to replace molecular identification.

Citation: Khonsanit A, Thanakitpipattana D, Mongkolsamrit S, Kobmoo N, Phosrithong N, Samson RA, Crous PW, Luangsa-ard JJ (2024). A phylogenetic assessment of Akanthomyces sensu lato in Cordycipitaceae (Hypocreales, Sordariomycetes): introduction of new genera, and the resurrection of Lecanicillium. Fungal Systematics and Evolution 14: 271–305. doi: 10.3114/fuse.2024.14.17

Keywords: Cordycipitaceae, molecular phylogeny, new taxa, taxonomy

INTRODUCTION

Cordycipitaceae (Hypocreales, Sordariomycetes, Ascomycota) was reintroduced by Sung et al. (2007), primarily relying on the phylogenetic placement of Cordyceps militaris, the designated type species. This family contains species producing stalked, erect, fleshy, and vividly coloured stromata. Nevertheless, certain species exhibit reduced stipes, giving rise to subiculate, pallid to cream, or white stromata on their insect hosts (Sung et al. 2007, Kepler et al. 2017, Wang et al. 2020, Mongkolsamrit et al. 2022). Species within the Cordycipitaceae are commonly found on various insect hosts, displaying diverse morphological characteristics (Wang et al. 2020). Recent investigations on the phylogenetic relationships and morphological differences within Cordycipitaceae have led to the establishment of a new mycoparasitic genus Niveomyces (Araújo et al. 2022), three additional genera on spiders, Jenniferia, Parahevansia, and Polystromomyces (Mongkolsamrit et al. 2022), the introduction of a new genus Neohyperdermium on scale insects (Thanakitpipattana et al. 2022), as well as Gamszarella from soil, basidiomycete sporocarps and dead insects (Arnold & Castañeda Ruíz 1987, Zhang et al. 2020, Crous et al. 2023). In addition to these newly established genera, 17 known genera have been documented, including Akanthomyces, Ascopolyporus, Beauveria, Blackwellomyces, Cordyceps, Engyodontium, Flavocillium, Gibellula, Hevansia, Lecanicillium, Leptobacillium, Liangia, Neotorrubiella, Parengyodontium, Pseudogibellula, Samsoniella, and Simplicillium. The asexual morphs within Cordycipitaceae are diverse and have been assigned different generic names, such as Acremonium, Akanthomyces, Beauveria, Engyodontium, Gibellula, Isaria, Lecanicillium, Pseudolecanicillium, Simplicillium, and Verticillium (Zare & Gams 2001, Kuephadungphan et al. 2019, 2020, 2022, Wang et al. 2020, Alvez et al. 2022).

The genus Akanthomyces was initially established by Lebert (1858), featuring the type species A. aculeatus discovered on moths in France. Akanthomyces aculeatus is characterized by its white to cream cylindrical synnemata, covered by a hymenium-like layer of phialides that produce catenulate, short, or long chains of conidia. The phialides are hyaline, cylindrical or flask-shaped, gradually or abruptly tapering to a more or less distinct neck (Mains 1950). Currently 42 species are classified within Akanthomyces and despite the diversity within the genus, the morphological characteristics among species significantly overlap, posing challenges for accurate morphological identification to species level (Manfrino et al. 2022). Some Akanthomyces species were described with an asexual morph resembling Isaria, including Akanthomyces bashanensis, Ak. beibeiensis, Ak. araneicola, Ak. araneogenum, Ak. kanyawimiae, Ak. kunmingensis, A. neoaraneogenus, Ak. subaraneicola, Ak. sulphureus, and Ak. waltergamsii (Chen et al. 2017, 2019a, Mongkolsamrit et al. 2018, Wang et al. 2024). Others were identified with a lecanicillium-like morph, such as Ak. lepidopterorum, Ak. noctuidarum, Ak. thailandicus, Ak. tiankengensis, and Ak. tortricidarum (Mongkolsamrit et al. 2018, Aini et al. 2020, Chen et al. 2020a, Chen et al. 2022), or a verticillium-like morph, namely Ak. neocoleopterorum (Chen et al. 2020b).

Akanthomyces species are known to infect a variety of hosts, including moths, scale insects, spiders, and can also be found in soil (Kepler et al. 2017, Chen et al. 2018, Shrestha et al. 2019, Aini et al. 2020). Seven recognized Akanthomyces species were found occurring on moths, viz. Ak. aculeatus (Lebert 1858), Ak. tuberculatus (Samson & Evans 1974), Ak. noctuidarum, Ak. pyralidarum, and Ak. tortricidarum (Aini et al. 2020), as well as Ak. laosensis and Ak. pseudonoctuidarum (Wang et al. 2024). In addition, five Akanthomyces species were identified parasitizing spiders: Ak. aranearum, Ak. kanyawimiae, Ak. sulphureus, Ak. thailandicus, and Ak. waltergamsii (Mains 1950, Mongkolsamrit et al. 2018). Furthermore, through phylogenetic analyses and morphological observations, some species were reclassified into the genera Hevansia (H. arachnophila, H. longispora, H. novoguineensis, H. ovalongata, H. websteri) and Jenniferia (J. cinerea) (Kepler et al. 2017, Mongkolsamrit et al. 2022).

The genera Lecanicillium and Simplicillium were introduced for verticillium-like taxa by Gams & Zare (2001). In Lecanicillium sensu lato, the species are phylogenetically dispersed within Cordycipitaceae, failing to form a single monophyletic clade (Sukarno et al. 2009), and includes more than 30 species (Manfrino et al. 2022). The morphology of Lecanicillium consists of discrete, aculeate phialides tapering to a narrow tip, generating short to long ellipsoidal- to falcate-shaped conidia with pointed ends, which cluster into slimy heads or fascicles (Gams & Zare 2001). Kepler et al. (2017) highlighted that the sexual species Cordyceps confragosa (Torrubiella confragosa) is associated with L. lecanii, the type species of Lecanicillium. This species has a cosmopolitan distribution, being found on a diverse range of insects (Kobe & Leal 2005).

Nevertheless, Kepler et al. (2017) considered the genus Lecanicillium as synonymous with Akanthomyces. Their molecular phylogenetic investigation, using five nuclear loci (SSU, LSU, TEF1, RPB1, and RPB2), revealed that the type species, L. lecanii, was nested within Akanthomyces, thus granting priority to Akanthomyces over Lecanicillium. Consequently, Kepler et al. (2017) proposed the suppression of Lecanicillium along with Isaria, Torrubiella, and other lesser-known names in favour of Akanthomyces. Lecanicillium was therefore considered synonymous with Akanthomyces, and species such as L. attenuatum, L. lecanii, L. longisporum, L. muscarium, L. pissodis, L. sabanense, and L. uredinophilum were transferred into Akanthomyces. Some species from Lecanicillium s.l. were also reassigned to Flavocillium, Gamszarea (Ga.) and Gamszarella (Gam.) based on molecular data, including F. primulinum, F. subprimulinum, Ga. kalimantanensis, Ga. restricta, Ga. testudinea, and Ga. wallacei, Gam. antillana, Gam. magnispora (Kepler et al. 2017, Shrestha et al. 2019, Wang et al. 2020, Zhang et al. 2020, Manfrino et al. 2022, Crous et al. 2023). However, the phylogeny of several species within Lecanicillium s.l. remain unresolved, e.g. L. aphanocladii, L. aranearum, L. cauligalbarum, L. coprophilum, L. dimorphum, L. evansii, L. flavidum, L. fungicola, L. nodulosum, L. psalliotae and L. saksenae (Zare & Gams 2001, Zare & Gams 2008, Sukarno et al. 2009, Zhou et al. 2018, Su et al. 2019).

During a survey of entomopathogenic fungi in Thailand, numerous moth cadavers consistently showed infection by Akanthomyces. The objectives of this study are to clarify the phylogenetic relationships among species in Akanthomyces sensu lato and Lecanicillium sensu lato by examining both their morphology and DNA phylogeny and to offer insights into which functional characteristics could be crucial and beneficial for phenotype-based identification.

MATERIALS AND METHODS

Isolates

Samples of entomopathogenic fungi on moths were collected throughout Thailand from 2002 to 2016. Stems of living plants, underside and upper side of leaves and the leaf litter were surveyed for dead moths infected with fungi. The collected specimens were placed in plastic boxes (3 × 6 cm or 6 × 10 cm) and returned to the laboratory for further examination. Fungal isolation was done from both asexual and sexual morphs found on the specimens; for asexual morphs, conidia sporulating from synnemata were streaked onto potato dextrose agar (PDA: 200 g potato, 20 g dextrose, 15 g agar, in 1 L distilled water) plates. After 24 to 48 h, plates were checked for contamination and cultures transferred to new PDA plates. For sexual morphs, the specimen was fastened on the underside of the lid of the Petri plate over the PDA and incubated overnight in a moist chamber to collect discharged ascospores which were then transferred to new PDA plates. All pure cultures were deposited in the BIOTEC culture collection (BCC). Fungal specimens were dried in a food dehydrator and deposited in the BIOTEC Bangkok Herbarium (BBH), Thailand Science Park, Pathum Thani Province, Thailand.

Molecular phylogenetic analyses

Genomic DNA was extracted from fungal cultures following a modified cetyltrimethyl-ammonium bromide (CTAB) method as described in Doyle & Doyle (1987). The DNA of the large subunit of the nuclear ribosomal DNA (LSU), partial regions of the translation elongation factor-1α gene (TEF1), and the largest and second largest subunits of the RNA polymerase II (RPB1, RPB2) were amplified and sequenced. The PCR primers used to amplify these markers are LROR and LR5 for LSU (Vilgalys & Hester 1990, Rehner & Samuels 1994), EF-983F and EF-2218R for TEF1 (Rehner & Buckley 2005), CRPB1 and RPB1-Cr for RPB1 (Castlebury et al. 2004), and fRPB2-5F2 and fRPB2-7cR for RPB2 (Liu et al. 1999, O’Donnell et al. 2007). Thermocycler conditions for PCR amplifications followed previously described protocols by Sung et al. (2007). The PCR products were purified and then sequenced using the same primers used for PCR amplification.

DNA sequences were assembled and manually edited using BioEdit v. 7.2.5 (Hall 1999). The alignment data were edited with MUSCLE v. 3.6 software (Edgar 2004). The combined dataset was analysed by maximum likelihood (ML) and Bayesian inference (BI). Maximum likelihood was performed with RAxML-HPC2 on XSEDE v. 8.2.12 in CIPRES Science Gateway portal with 1 000 bootstrap replicates (Stamatakis 2014). Bayesian analysis was performed using MrBayes on XSEDE v. 3.2.7a (Ronquist et al. 2012), with the best fit model SYM+G model. Four chains for the MCMC variant were run with 5 000 000 generations, with sampling a tree every 1 000 generations and a burn in of 10 % from the total run.

Morphology

Photos of specimens were taken using a digital Nikon D5100 camera and Olympus DP70 Digital Camera installed on an Olympus SZX12 dissecting microscope. The micromorphological characters were photographed using a DP70 digital camera installed on an Olympus BX51 compound microscope. Morphological examination of macroscopic and microscopic features was done on microscope slides using lactophenol cotton blue. For the asexual morph, synnemata with phialides and conidia were observed from host tissue. For the sexual morph, asci and ascospores were removed from perithecia. For colony characteristics on the culture media and microscopic measurements of phialides and conidia, strains were grown on PDA and oatmeal agar (OA, Difco) in white light/dark cycles in the laboratory, after 30 d of incubation at 25 °C. The size and the shape of these morphological characters were measured with a minimum sampling size of 5−20 for perithecia and ca. 50 for phialides, conidia, asci, asci caps, part-spores to obtain an average ± standard deviation with absolute minima and maxima in parentheses. The 6^th Royal Horticultural Society (RHS) colour chart was used to describe the colours of fresh specimens and cultures (RHS Media 2015).

RESULTS

Molecular phylogenetic analyses

Sequences from a total of 166 strains in Cordycipitaceae were incorporated into this study, which included newly acquired data from 15 specimens of Akanthomyces species on moths from our collections (Table 1). Trichoderma deliquescens and T. stercorarium served as the outgroup. A final alignment of 3 541 bp from the concatenated four-loci (LSU = 892 bp, TEF1 = 1 029 bp, RPB1 = 722 bp, and RPB2 = 898 bp) was used for the phylogenetic analyses. Our results reveal that while Akanthomyces sensu lato forms a strongly supported and distinct monophyletic clade [88 % ML bootstrap (MLBS) and 0.85 BI posterior probability (BIPP)] (see Fig. 1A), it can be further divided into four monophyletic subclades with distinctive host preferences and morphologies, tentatively identified as Akanthomyces, Arachnidicola, Lecanicillium clades, and Kanoksria.

Table 1.

List of taxa included in the phylogenetic analyses and their GenBank accession numbers. Newly generated sequences in this study are marked in bold font. Ex-type strains are indicated with ^T.

Species	Strain	GenBank Accession numbers				Reference

		LSU	*TEF1*	*RPB1*	*RPB2*
Akanthomyces aculeatus	HUA 186145^T	MF416520	MF416465	—	—	Kepler et al. (2017)
	HUA 772	KC519370	KC519366	—	—	Sanjuan et al. (2014)
*Akanthomyces buriramensis*	BCC 45158	ON008556	ON013546	ON013561	—	This study
	BCC 45166	ON008544	ON013547	ON013562	—	This study
	BCC 47939^T	ON008545	ON013548	ON013563	—	This study
*Akanthomyces fusiformis*	BCC 40756^T	ON008549	ON013552	ON013567	ON013576	This study
*Akanthomyces laosensis*	BCC 49306	ON008547	ON013549	ON013564	ON013573	This study
	BCC 52398	ON008546	ON013550	ON013565	ON013574	This study
	BCC 74583	ON008548	ON013551	ON013566	ON013575	This study
	YFCC 1910941^T	OQ509510	OQ506286	OQ511535	OQ511549	Wang et al. (2024)
	YFCC 1910942	OQ509511	OQ506287	OQ511536	OQ511550	Wang et al. (2024)
*Akanthomyces niveus*	BCC 40747	ON008550	ON013553	ON013568	ON013577	This study
	BCC 79887^T	ON008551	ON013554	—	ON013578	This study
Akanthomyces noctuidarum	BBH16595	MT356085	MT477979	MT477995	MT478005	Aini et al. (2020)
	BCC28571	MT356087	MT477981	MT478009	MT478006	Aini et al. (2020)
	BCC36265^T	MT356084	MT477978	MT477994	MT477987	Aini et al. (2020)
	BCC47498	MT356086	MT477980	MT477996	MT477988	Aini et al. (2020)
*Akanthomyces phariformis*	BCC 45148^T	ON008556	ON013559	—	ON013583	This study
	BCC 76537	ON008557	ON013560	—	ON013584	This study
*Akanthomyces pseunoctuidarum*	BCC 48963	ON008552	ON013555	ON013569	ON013579	This study
	BCC 69011	ON008553	ON013556	ON013570	ON013580	This study
	BCC 69056	ON008554	ON013557	ON013571	ON013581	This study
	BCC 76483	ON008555	ON013558	ON013572	ON013582	This study
	YFCC 1808943^T	OQ509512	OQ506288	OQ511537	OQ511551	Wang et al. (2024)
	YFCC 1808944	OQ509513	OQ506289	OQ511538	OQ511552	Wang et al. (2024)
Akanthomyces pyralidarum	BCC28816^T	MT356091	MT477982	MT478000	MT478007	Aini et al. (2020)
	BCC29197	MT305694	MT508840	MT478003	MT477991	Aini et al. (2020)
	BCC40869	MT356093	MT477984	MT478002	MT477990	Aini et al. (2020)
Akanthomyces tortricidarum	BCC28583	MT356090	MT477986	MT477999	MT477993	Aini et al. (2020)
	BCC41868	MT356089	MT477985	MT477998	MT478008	Aini et al. (2020)
	BCC72638^T	MT356088	MT478004	MT477997	MT477992	Aini et al. (2020)
Akanthomyces tuberculatus	HUA 186131	MF416521	MF416466	—	—	Kepler et al. (2017)
	OSC 111002	DQ518767	DQ522338	DQ522384	DQ522435	Spatafora et al. (2007)
Arachnidicola araneicola	GY29011^T	MK942430	MK955950	MK955944	MK955947	Chen et al. (2019a)
	GY29012	MK942435	MK955951	MK955945	MK955948	Chen et al. (2019a)
Arachnidicola araneogenum	GZUIF DX1	MH978178	—	MH978181	MH978184	Chen et al. (2018)
	GZUIF SN1	MH978180	MH978188	MH978183	MH978186	Chen et al. (2018)
Arachnidicola bashanensis	CQ05621^T	OQ300420	OQ325024	—	OQ349684	Chen et al. (2023)
	CQ05622	OQ300421	OQ325025	—	OQ349685	Chen et al. (2023)
Arachnidicola beibeiensis	CQ05921^T	OQ300424	OQ325028	—	OQ349688	Chen et al. (2023)
	CQ05922	OQ300427	OQ325029	—	OQ349689	Chen et al. (2023)
Arachnidicola kanyawimiae	NHJ 6709	EU369042	EU369025	EU369067	EU369086	Johnson et al. (2009)
	CBS 541.81	MF416553	MF416498	MF416655	MF416449	Kepler et al. (2017)
	TBRC 7242	MF140718	MF140838	MF140784	MF140808	Mongkolsamrit et al. (2018)
	TBRC 7244^T	MF140716	MF140836	—	—	Mongkolsamrit et al. (2018)
Arachnidicola kunmingensis	YFCC 1708939	OQ509508	OQ506284	OQ511533	OQ511547	Wang et al. (2024)
	YFCC 1808940^T	OQ509509	OQ506285	OQ511534	OQ511548	Wang et al. (2024)
Arachnidicola subaraneicola	YFCC 2107937^T	OQ509514	OQ506290	OQ511539	OQ511553	Wang et al. (2024)
	YFCC 2107938	OQ509515	OQ506291	OQ511540	OQ511554	Wang et al. (2024)
Arachnidicola sulphurea	NHJ 5112	EU369043	EU369026	EU369066	—	Johnson et al. (2009)
	TBRC 7248^T	MF140722	MF140843	MF140787	MF140812	Mongkolsamrit et al. (2018)
	TBRC 7249	MF140721	MF140842	MF140786	MF140734	Mongkolsamrit et al. (2018)
Arachnidicola thailandica	TBRC 7245^T	—	MF140839	—	MF140809	Mongkolsamrit et al. (2018)
	TBRC 7246	MF140719	MF140840	—	MF140810	Mongkolsamrit et al. (2018)
Arachnidicola tiankengensis	KY11571^T	ON502825	ON525447	—	ON525446	Chen et al. (2022)
	KY11572	ON502827	ON525449	—	ON525448	Chen et al. (2022)
Arachnidicola waltergamsii	TBRC 7250	MF140715	MF140835	—	—	Mongkolsamrit et al. (2018)
	TBRC 7251	MF140713	MF140833	MF140781	MF140805	Mongkolsamrit et al. (2018)
Ascopolyporus polychrous	P.C. 546	DQ118737	DQ118745	DQ127236	—	Chaverri et al. (2005)
Ascopolyporus villosus	ARSEF 6355	—	AY886544	DQ118750	DQ127241	Chaverri et al. (2005)
Beauveria bassiana	ARSEF 300	—	AY531924	HQ880831	HQ880903	Rehner et al. (2011)
Beauveria staphylinidicola	ARSEF 5718	—	EF468836	EF468776	EF468881	Sung et al. (2007)
Bhushaniella rubra	BCC 47515	OQ892132	OQ914427	OQ914430	—	Mongkolsamrit et al. (2023)
	BCC 47541^T	OQ892133	OQ914428	OQ914431	OQ914433	Mongkolsamrit et al. (2023)
	BCC 47542	OQ892134	OQ914429	OQ914432	OQ914434	Mongkolsamrit et al. (2023)
Blackwellomyces pseudomilitaris	BCC 1919^T	MF416534	MF416478	—	MF416440	Kepler et al. (2017)
	BCC 2091	MF416535	MF416479	—	MF416441	Kepler et al. (2017)
Cordyceps blackwelliae	TBRC 7256^T	MF140702	MF140822	MF140771	MF140795	Mongkolsamrit et al. (2018)
Cordyceps cf. ochraceostromata	ARSEF 5691	EF468819	EF468759	EF468867	EF468921	Sung et al. (2007)
Cordyceps tenuipes	ARSEF 5135	JF415980	JF416020	JN049896	JF416000	Kepler et al. (2012)
Corniculantispora aphanocladii	CBS 797.84^T	KM283787	KM283811	KM283833	KM283853	Park et al. (2015)
Corniculantispora dimorphum	CBS 345.37	KM283788	KM283812	KM283834	KM283854	Park et al. (2015)
	CBS 363.86^T	—	AF339559	EF468784	EF468890	Sung et al. (2007)
Corniculantispora psalliotae	CBS 101270	EF469081	EF469066	EF469113	EF469095	Sung et al. (2007)
	CBS 532.81^T	AF339560	EF469067	EF469096	EF469112	Sung et al. (2007)
Corpulentispora magnispora	CGMCC3.19304^T	MK329007	MK336037	—	MK335985	Zhang et al. (2020)
	LC12469	MK329008	MK336038	—	MK335986	Zhang et al. (2020)
Engyodontium aranearum	CBS 309.85	AF339526	DQ522341	DQ522387	DQ522439	Sung et al. (2001), Kepler et al. (2017)
Engyodontium huhutii	GZUIFRhuhu^T	—	MT006068	MT006058	MT006063	Zhou et al. (2022)
Engyodontium parvisporum	IHEM 22910^T	LC092915	—	—	—	Tsang et al. (2016)
Engyodontium rectidentatum	CBS 641.74	LC092914	LC425540	—	—	Tsang et al. (2016)
Flavocillium acerosum	CBS 418.81^T	KM283786	KM283810	KM283832	KM283852	Park et al. (2015)
Flavocillium bifurcatum	YFCC 6101^T	MN576781	MN576951	MN576841	MN576897	Wang et al. (2020)
Flavocillium praecognitum	MGC39	MT247060	MT267523	—	MT267525	Zhou et al. (2022)
	MGC76	MT247061	MT267524	—	MT267526	Zhou et al. (2022)
Flavocillium primulinum	JCM 18525^T	AB712263	—	—	—	Wang et al. (2020)
	JCM 18526	AB712264	—	—	—	Wang et al. (2020)
Flavocillium subprimulinum	HKAS99548^T	MG585317	—	—	—	Wang et al. (2020)
	HKAS99549	MG585321	—	—	—	Wang et al. (2020)
Gamszarea coprophilum	CGMCC3.18986^T	MH177619	MH184587	MH177622	MH177624	Su et al. (2019)
	CGMCC3.18987	MH177618	MH184586	MH177621	MH177623	Su et al. (2019)
Gamszarea gracile	CBS 142816	—	MN602797	—	—	Ponizovskaya et al. (2020)
Gamszarea humicola	CGMCC3.19303^T	MK328997	MK336027	—	MK335979	Zhang et al. (2020)
	LC 12462	MK328998	MK336028	—	MK335980	Zhang et al. (2020)
Gamszarea lunata	CGMCC3.19315^T	MK328999	MK336029	—	MK335981	Zhang et al. (2020)
	LC12546	MK329000	MK336030	—	MK335982	Zhang et al. (2020)
Gamszarea microspora	CGMCC3.19313^T	MK329001	MK336031	—	MK335983	Zhang et al. (2020)
	LC12531	MK329002	MK336032	—	MK335985	Zhang et al. (2020)
Gamszarea restrictum	CCF 5252^T	—	LT626943	—	—	Zhou et al. (2022)
Gamszarea testudineum	UBOCC-A112180	—	LT992868	—	—	Zhou et al. (2022)
	UBOCC-A116026	—	LT992867	—	—	Zhou et al. (2022)
Gamszarea wallacei	CBS 101237^T	AY184967	EF469073	EF469102	EF469119	Sung et al. (2007)
Gamszarella buffelskloofina	CBS 150062	OR717025	—	OR683715	OR683726	Crous et al. (2023)
Gamszarella antillana	CBS 350.85^T	AF339536	DQ522350	DQ522396	DQ522450	Spatafora et al. (2007)
Gibellula nigelii	NHJ 10788	EU369036	EU369019	EU369058	EU369078	Johnson et al. (2009)
Gibellula pulchra	NHJ 10808	EU369035	EU369018	EU369056	EU369076	Johnson et al. (2009)
Gibellula sp.	NHJ 13158	EU369037	EU369020	EU369057	EU369077	Johnson et al. (2009)
Hevansia nelumboides	BCC 2093	MF416530	MF416473	—	MF416437	Kepler et al. (2017)
Hevansia novoguineensis	NHJ 4314	—	EU369012	EU369051	EU369071	Johnson et al. (2009)
Jenniferia cinerea	NHJ 3510^T	—	EU369009	EU369048	EU369070	Johnson et al. (2009)
	BCC 2191	GQ249971	GQ250029	—	—	Kuephadungphan et al. (2019)
Kanoksria zaquensis	HMAS 246915^T	MT789697	MT797812	MT797810	—	Wang et al. (2023)
	HMAS 246917	MT789696	MT797811	MT797809	—	Wang et al. (2023)
Lecanicillium aranearum	CBS 726.73a	AF339537	EF468781	EF468887	EF468934	Sung et al. (2001); Sung et al. (2007)
Lecanicillium araneogenum	GZU1031Lea^T	KX845703	KX845697	KX845699	KX845701	Chen et al. (2017)
Lecanicillium araneosum	KY11341^T	ON502832	ON525443	—	ON525442	Chen et al. (2022)
	KY11342	ON502837	ON525445	—	ON525444	Chen et al. (2022)
Lecanicillium attenuatum	CBS 170.76^T	OP752153	OP762607	OP762611	OP762615	Manfrino et al. (2022)
	KACC 42493	KM283780	KM283804	KM283826	KM283846	Park et al. (2015)
	CBS 402.78	AF339565	EF468782	EF468888	EF468935	Sung et al. (2007)
Lecanicillium fusisporum	CBS 164.70^T	AF339549	EF468783	EF468889	—	Sung et al. (2007)
Lecanicillium lecanii	CBS 101247	AF339555	DQ522359	DQ522407	DQ522466	Kepler et al. (2012)
	CBS 102067^T	KM283795	KM283818	KM283838	KM283860	Park et al. (2015)
Lecanicillium lepidopterorum	GZAC SD05151^T	MT705973	—	—	MT727044	Chen et al. (2020a)
	GZAC SD05152	MT705974	—	—	MT727045	Chen et al. (2020a)
Lecanicillium longisporum	CBS 126.27^T	AF339556	KM283820	KR064300	KR064303	Chiriví-Salomón et al. (2015), Park et al. (2015)
	CBS 102072	KM283796	KM283819	KM283839	KM283861	Park et al. (2015)
Lecanicillium muscarium	CBS 143.62^T	—	KR064305	KR064301	KR064304	Chiriví-Salomón et al. (2015)
Lecanicillium neocoleopterorum	GY11241^T	MN09396	MN097813	MN097816	MN097812	Chen et al. (2020b)
	GY11242	MN09398	MN097815	MN097817	MN097814	Chen et al. (2020b)
Lecanicillium pissodis	CBS 118231^T	KM283799	KM283822	KM283842	KM283864	Chen et al. (2020b)
Lecanicillium sabanense	ANDES-F 1024^T	KC875225	KC633266	—	KC633250	Chiriví-Salomón et al. (2015)
Lecanicillium sp.	YFCC 945	—	OQ506294	OQ511543	OQ511557	Wang et al. (2024)
Lecanicillium uredinophilum	KACC 44082^T	KM283782	KM283806	KM283828	KM283848	Park et al. (2015)
	KACC 47756	KM283783	KM283807	KM283829	KM283849	Park et al. (2015)
Liangia sinensis	YFCC 3103^T	MN576782	MN576952	MN576842	MN576898	Wang et al. (2020)
	YFCC 3104	MN576783	MN576953	MN576843	MN576899	Wang et al. (2020)
Neohyperdermium piperis	CBS 116719^T	AY466442	DQ118749	DQ127240	EU369083	Chaverri et al. (2005), Johnson et al. (2009)
Neohyperdermium pulvinatum	P.C.602	DQ118738	DQ118746	DQ127237	—	Chaverri et al. (2005)
Neotorrubiella chinghridicola	BCC39684	MK632096	MK632071	MK632181	MK632148	Thanakitpipattana et al. (2020)
	BCC80733^T	MK632097	MK632072	MK632176	MK632149	Thanakitpipattana et al. (2020)
Niveomyces coronatus	Niveo	ON493605	—	—	ON513401	Araújo et al. (2022)
	NY 04434800^T	ON493606	ON513397	ON513399	ON513400	Araújo et al. (2022)
Parengyodontium album	CBS 368.72	LC092910	—	—	—	Tsang et al. (2016)
	CBS 504.83^T	LC092899	—	—	—	Tsang et al. (2016)
Pleurodesmospora lepidopterorum	DY10501^T	—	MW834317	MW834315	MW834316	Chen et al. (2021)
	DY10502	—	—	MW834318	MW834319	Chen et al. (2021)
Polystromomyces araneae	BCC 93301^T	MZ684016	MZ707825	MZ707832	MZ707845	Mongkolsamrit et al. (2022)
Pseudogibellula formicarum	BCC 81493	MT512652	MT863566	MT533472	—	Mongkolsamrit et al. (2021)
	CBS 433.73	MH872442	MT533481	MT533475	—	Mongkolsamrit et al. (2021)
Pseudolecanicillium caatingaense	URM 8446	ON862925	OP290526	—	OP290514	Alves et al. (2022)
Pseudolecanicillium caatingaense	URM 8447 ^T	ON862926	OP290525	—	OP290513	Alves et al. (2022)
Pseudoniveomyces blattae	BCC 53567^T	ON103167	—	ON286885	ON125036	Kobmoo et al. (2023)
	BCC 53568	ON103168	ON125025	ON286886	ON125037	Kobmoo et al. (2023)
Samsoniella aurantia	TBRC 7271^T	MF140728	MF140846	MF140791	MF140818	Mongkolsamrit et al. (2018)
	TBRC 7273	MF140726	MF140844	—	MF140816	Mongkolsamrit et al. (2018)
Samsoniella inthanonensis	TBRC 7915^T	MF140725	MF140849	MF140790	MF140815	Mongkolsamrit et al. (2018)
	TBRC 7916	MF140724	MF140848	MF140789	MF140814	Mongkolsamrit et al. (2018)
Simplicillium lanosoniveum	CBS 101267	AF339554	DQ522357	DQ522405	DQ522463	Kepler et al. (2017)
	CBS 704.86	AF339553	DQ522358	DQ522406	DQ522464	Kepler et al. (2017)
Simplicillium obclavatum	CBS 311.74^T	AF339517	EF468798	—	—	Sung et al. (2007)
Trichoderma deliquescens	ATCC 208838	AF543791	AF543781	AY489662	DQ522446	Castlebury et al. (2004)
Trichoderma stercorarium	ATCC 62321	AF543792	AF543782	AY489633	EF469103	Castlebury et al. (2004)
Zarea flavidum	CBS 300.70D	KM283789	KM283813	—	KM283855	Park et al. (2015)
Zarea aleophilum	CBS 357.80^T	KM283791	KM283815	KM283835	KM283856	Park et al. (2015)
Zarea fungicola	CBS 992.69^T	KM283792	KM283816	—	KM283857	Park et al. (2015)
Zouia cauligalbarum	GZUIFRZHJ01	—	MH801920	MH801922	MH801924	Zhou et al. (2022)
	GZUIFRZHJ02	—	MH801921	MH801923	MH801925	Zhou et al. (2022)

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Fig. 1A, B. — RAxML tree of *Akanthomyces*, *Arachnidicola*, *Lecanicillium* and related genera in the *Cordycipitaceae* from a combined LSU, *TEF1*, *RPB1* and *RPB2* dataset. Number on the nodes are ML bootstrap values/Bayesian posterior probability above 70 %. Bold lines mean support for the three analyses were 100 %. In yellow boxes are species with lecanicillium-like asexual morphs. ^T = ex-type culture.

The Arachnidicola clade (A) represents a well-supported group (MLBS = 70 %, BIPP = 0.85), consisting exclusively of species found on spiders. These include Ar. sulphurea, Ar. waltergamsii, Ar. kunmingensis, Ar. thailandica, Ar. subaraneicola, Ar. araneicola, Ar. neoaraneogenus, Ar. bashanensis, Ar. beibeiensis, Ar. tiankengensis, and Ar. kanyawimiae. Most of these species exhibit an isaria-like asexual morph, apart from Ar. tiankengensis, which produces a lecanicillium-like asexual morph (Chen et al. 2022). Consequently, we propose transferring all known species in Clade A to the new genus Arachnidicola. It is noteworthy that in our analyses, Ar. coccidioperitheciatus NHJ06709 (Johnson et al. 2009) and Ar. farinosa CBS 541.81 (Kepler et al. 2017) were nested within Ar. kanyawimiae, indicating they are conspecific with Ar. kanyawimiae.

The Lecanicillium clade (B) forms a well-supported monophyletic group (BIPP = 0.79), comprising species that were originally classified in Lecanicillium but were transferred to Akanthomyces sensu Kepler et al. (2017) such as Ak. attenuatus, Ak. lecanii, Ak. dipterigenus, Ak. muscarius, Ak. sabanensis as well as Ak. neoaraneogenus (Shrestha et al. 2019), Ak. lepidopterorum, Ak. pissodis (Chen et al. 2020a), Ak. neocoleopterorum (Chen et al. 2020b), Ak. araneosus (Chen et al. 2022) and Ak. uredinophilus (Manfrino et al. 2022). Given that the type species of Lecanicillium, L. lecanii, is nested within this well-supported clade, we propose the resurrection of the genus Lecanicillium, and suggest transferring all species in this clade to Lecanicillium. Species within clade B exhibit both lecanicilliumlike and verticillium-like asexual morphs, and they have been found on various substrates, including a wide range of insects, spiders, decaying leaves, leaf litter, and mushrooms.

The Akanthomyces clade (C) is strongly supported (MLBS = 98 %, BIPP = 0.85) and distinctly separates from the Arachnidicola (A) and Lecanicillium (B) clades. Within clade C, we identify four new species: Akanthomyces phariformis, Ak. fusiformis, Ak. niveus, Ak. buriramensis, and two new records, Ak. laosensis and Ak. pseudonoctuidarum. Additionally, three Akanthomyces species previously described by Aini et al. (2020), including the type species Ak. aculeatus, are also part of this clade. All species within this clade were observed on adult moths and predominantly exhibit an akanthomyces-like asexual morph. Notably, Ak. niveus and Ak. phariformis did not show evidence of producing an asexual morph. Consequently, this clade can be designated as Akanthomyces sensu stricto.

Basal to the previously mentioned clades A–C is clade D, which exclusively includes Ak. zaquensis strains (Wang et al. 2023). This clade forms a distinct, well-supported group (BIPP = 1.00), separate from the Akanthomyces clade. As a result, it is proposed that this clade does not fall under the classification of Akanthomyces and should be designated as the new genus Kanoksria.

Significantly, Fig. 1 encompasses nearly all species within Lecanicillium. However, some species are represented solely by ITS sequences due to the absence of multigene data, preventing their inclusion in the analysis. The resulting topology in Fig. 1 aligns with earlier phylogenetic studies in Cordycipitaceae. The phylogenetic tree exposes the non-monophyletic nature of the Lecanicillium lineage, revealing it as a polyphyletic genus distributed across various clades within the Cordycipitaceae (Zare & Gams 2008, Wang et al. 2020, Zhou et al. 2022). Certain species, as illustrated in Fig. 1A, B, were transferred to other genera such as Akanthomyces, Flavocillium (clade 3), Gamszarea (clade 9) and Gamszarella (clade 6) (Kepler et al. 2017, Wang et al. 2020, Zhang et al. 2020, Zhou et al. 2022, Crous et al. 2023). In this study, we suggest transferring Lecanicillium huhutii and L. tenuipes from clade 8 to the genus Engyodontium. Additionally, we propose new genera for clades 2, 4, 5 and 7, namely Corniculantispora, Zouia, Zarea, and Corpulentispora. These proposals are moderately to strongly supported based on multilocus phylogeny. Single-locus phylogenetic analyses did not consistently recover these clades. Among the loci, only RPB1 provided all three monophyletic clades with robust support [MLBS: Clade A = 96 %, Clade B = 95 %, Clade C = 100 %], while RPB2 only recapitulated Clade A and Clade C with complete support [MLBS = 100 %], excluding Clade B, which was divided into a clade containing Lecanicillium lecanii + L. sabanense segregating from the rest of Lecanicillium (Supplementary Fig. S1).

Taxonomy

Akanthomyces buriramensis Mongkols., Khons., Thanakitp. & Luangsa-ard, sp. nov. MycoBank MB 842450. Fig. 2.

Etymology: The name is derived from the locality where the specimens were found, Buri Ram Province, Thailand.

Typus: Thailand, Buri Ram Province, Non Din Daeng District, Dong Yai Wildlife Sanctuary, Pong Kao nature trail, 14°17’46”N, 102°44’11”E, on adult moth (Noctuidae), 11 Dec. 2010, A. Khonsanit, A. Saksrikrom, B. Saracam, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (holotype BBH30240 preserved in a metabolically inactive state; ex-holotype culture BCC47939). GenBank accession numbers: ON008545 (LSU), ON013548 (TEF1), ON013563 (RPB1).

Asexual morph: Synnemata scattered, arising from the moths’ body, filiform, solitary, occasionally curved, white, pale yellow (18C), light orange yellow (22D), up to 10 mm long. Conidiophores densely compacted into a layer along the synnemata, cylindrical, monophialidic or polyphialidic. Phialides smooth-walled, ampulliform, obpyriform, lageniform, hyaline, (4–)4.5–7(–10) × 3–4 μm. Conidia 1-celled, ellipsoid, fusoid, obovoid, smooth-walled, hyaline, 3–5 × 1–2 μm. Sexual morph: Specimens found on adult moths (Noctuidae, Notodontidae), on the stem of living bamboo twigs and underside of dicotyledonous leaves. Perithecia superficial, crowded in the middle synnemata, mostly on one side of the synnema, ovoid, strong orange yellow (N163B–D), strong yellowish brown (N199D), moderate yellowish brown (N199C), (500–)560–635(–750) × (300–)320– 380(–400) μm. Asci cylindrical, hyaline, 250–450 × 3.5–4 μm. Asci-caps hemispherical 3 × 4–5 μm. Ascospores filiform, multiseptate, breaking into part-spores, hyaline, up to 250 μm long. Part-spores cylindrical, hyaline, (4–)6–14.5 × 0.5–1 μm.

Colony characteristics: Colony obverse on OA compact, raised to the agar surface, entire margin, attaining a diam of 35 mm in 20 d, white. Colony reverse on OA strong yellow (153D). Colony obverse on PDA compact, raised to the agar surface, entire margin, attaining a diam of 35 mm in 20 d, white. Colony reverse on PDA moderate yellow (160D). Conidia and reproductive structures are not observed on both media after 30 d of incubation at 25 °C.

Additional specimens examined: Thailand, Buri Ram Province, Non Din Daeng District, Dong Yai Wildlife Sanctuary, Headquarter nature trail, 14°17’46”N, 102°44’11”E, on adult moth (Noctuidae), 30 Nov. 2010, A. Khonsanit, A. Saksrikrom, B. Saracam, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH30105; BBH30106; BBH30107:BCC45157:BCC45158; BBH30108); ibid., Pa Takong nature trail, 14°17’46”N, 102°44’11”E, on adult moth, 10 Dec. 2010, A. Khonsanit, A. Saksrikrom, B. Saracam, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH30170:BCC45165); ibid., (BBH30119:BCC45166; BBH30249); ibid., Pong Kao nature trail, 14°17’46”N, 102°44’11”E, on adult moth (Notodontidae), 11 Dec. 2010, A. Khonsanit, A. Saksrikrom, B. Saracam, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH30239:BCC47938; BBH30240).

Notes: Both the asexual and sexual morph of Ak. buriramensis can be found on the same specimen. Phylogenetically, Ak. buriramensis is closely related to Ak. noctuidarum (Fig. 1A). Both species are phylogenetically well-supported and Ak. buriramensis is found in Buri Ram Province, lower northeastern Thailand, while Ak. noctuidarum strains are found throughout Thailand. Morphologically, Ak. buriramensis differs from other related species in producing crowded superficial perithecia laterally at the midsection of the short synnemata (Fig. 2C).

Akanthomyces fusiformis Khons., Thanakitp. & Luangsa-ard, sp. nov. MycoBank MB 842452. Fig. 3.

Etymology: The name is derived from the fusiform shape of its conidia.

Typus: Thailand, Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Mo Sing To nature trail, 14°26’23”N, 101°22’20”E, on adult moth (Pyralidae), 10 Dec. 2009, A. Khonsanit, K. Tasanathai, M. Sudhadham, P. Srikitikulchai, R. Ridkaew, S. Mongkolsamrit & T. Chohmee (holotype BBH27326 preserved in a metabolically inactive state; ex-holotype strain BCC40756). GenBank accession numbers: ON008549 (LSU), ON0013552 (TEF1), ON0013567 (RPB1), ON0013576 (RPB2).

Asexual morph: White mycelium covering the body of the host. Conidiophores mononematous, smooth-walled, hyaline, with single phialide. Phialides elongate-ampulliform, smooth-walled, hyaline, (4–)6–15(–27) × 1–3 μm. Conidia 1-celled, narrowly fusiform, smooth-walled, hyaline, (4–)7–10(–13) × 1–2 μm. Sexual morph: Specimens found on adult moths (Pyralidae), on the underside of dicotyledonous leaves, white mycelium covered on the host body. Perithecia superficial, scattered on the host body, ovoid, strong orange (26A), light orange yellow (19A), light yellowish pink (19B), 580–640 × 170–210 μm. Asci cylindrical, hyaline, (302–)361–420(–450) × 4–5 μm. Asci-caps abrupt cone-shaped, hyaline, 4.5–5.5 × 4–5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, (5–)7–13(–18) × 1 μm.

Colony characteristics: Colony obverse on OA compact, wrinkled, raised to the agar surface, entire margin, attaining a diam of 31– 40 mm in 20 d, white to pale yellow green (195D). Colony reverse on OA moderate yellow (160A). Colony obverse on PDA cottony, convex to the agar surface, entire margin, attaining a diam of 30–32 mm in 20 d, white. Colony reverse on PDA moderate yellow (160A), pale yellow (160D) along the margin. Conidia and reproductive structures are not observed on both media after 30 d of incubation at 25 °C.

Notes: There is only one specimen of A. fusiformis found on an adult moth (Pyralidae) from Khao Yai National Park, Nakhon Ratchasima Province, Thailand. Both asexual and sexual morph can be seen on the same specimen. However, the asexual morph of Ak. fusiformis does not produce synnemata on its host but produces the phialides scattered on white mycelium covering the host body and wings. Phylogenetically, Ak. fusiformis is closely related to Ak. pyralidarum (Fig. 1A). Morphologically, Ak. fusiformis differs from Ak. pyralidarum in producing superficial perithecia, while Ak. pyralidarum produces crowded superficial perithecia at the tip of synnemata (Aini et al. 2020). Akanthomyces fusiformis differs from other related species by producing cone-shaped asci-caps (Fig. 3E), and long narrowly fusiform conidia (Fig. 3I), see Table 2.

Table 2.

Morphological comparisons between an asexual morph of Akanthomyces species on moths.

Species	Synnemata	Phialides (μm)	Conidia (μm)	References
Ak. aculeatus	Yellowish, cylindrical, narrowing upward, 1–8 mm long, 0.1–0.5 mm wide	Subcylindric, narrowly ellipsoidal, 6–16 × 2.5–4	Ellipsoidal, obovoid, 3–6 × 2–3	Lebert (1858)
Ak. buriramensis	Scattered, arising from the moths body, white, yellow to orange, up to 10 mm long	Ampulliform, obpyriform, lageniform, 4–10 × 3–4	Ellipsoid, fusiform, obovoid, 3–5 × 1–2	This study
Ak. fusiformis	—	Elongate-ampulliform, 4–27 × 1–3	Narrowly fusiform, 4–13 × 1–2	This study
Ak. laosensis	Scattered, arising different parts of the host body, white, 3−20 mm long	Ampulliform, lageniform, obpyriform, 4–10 × 2–3	Fusiform elliptical, elliptical, 3–6 × 1.5–2	This study
Ak. noctuidarum	Long synnema arising from the thorax of the hosts, filiform, simple, solitary, occasionally curved, white, up to 30 mm long	Cylindrical with papillate end, 4−9 × 2−3	Cylindrical with rounded end, occasionally elliptical and fusiform, 3−6 × 1−2	Aini et al. (2020),
	Short synnemata scattered, arising different parts of the host body and wing veins, simple, solitary, cylindrical with rounded end, white	Cylindrical with papillate end, 4−12 × 2−4	Cylindrical with rounded end, occasionally elliptical and fusiform, 4−6 × 1−2	This study
Ak. pseudonoctuidarum	Long synnemata, filiform, occasionally curved	Ampulliform, obclavate, obpyriform, lageniform, 6–13 × 2−4	Obovoid, 3–6 × 1.5–2	This study
	Short synnemata, filiform, occasionally curved, ca. 1–30 mm long	Ampulliform, obclavate, obpyriform, lageniform, 4–17 × 2–4	Obovoid, 3–5 × 1.5–2.5
Ak. tortricidarum	Long synnemata, cylindrical to clavate with acute or blunt end, up to 5 mm long and wide ca. 120–150 μm.	Cylindrical to ellipsoidal with papillate end, 5–10 × 1.8–3	Fusoid, 2–3.2 × 1–2	Aini et al. (2020)
	Short synnemata, cylindrical with subglobose or oblong at the end, 197–300 × 15–40 μm, with diameter of the tip 43–75 μm.	Cylindrical to ellipsoidal with papillate end, 5–10 × 1.8–3	Fusoid, 1–3 × 1–2
Ak. tuberculatus	Scattered, arising from different part of the host body, white to creamish, cylindrical, clavate, 1–6 mm long, 50–300 mm wide.	Cylindrical, 7–10.5 × 2.7–3.5	Cylindrical, narrowly fusiform, 4.5–6 × 1.2–1.5	Samson & Evans (1974)

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Akanthomyces niveus Khons., Thanaktip. & Luangsa-ard, sp. nov. MycoBank MB 842453. Fig. 4.

Etymology: In Latin “nivea” means white, refers to the colour of the insect hosts of this species.

Typus: Thailand, Chiang Mai Province, Kanlayaniwatthana District, Ban Huai Baba community forest nature trail, 18°59’13”N, 98°17’06”E, on adult moth (Lepidoptera), 25 Nov. 2015, D. Thanakitpipattana, K. Tasanathai, P. Srikitikulchai, R. Promharn, S. Mongkolsamrit, S. Wongkanoun & W. Noisripoom (holotype BBH40778 preserved in a metabolically inactive state; ex-holotype strain BCC79887). GenBank accession numbers: ON008551 (LSU), ON0013554 (TEF1), ON0013578 (RPB2).

Asexual morph: Unknown. Sexual morph: Specimens found on adult moths (Lepidoptera) on the stem of living plants and underside of dicotyledonous leaves, thin whitish mycelium covering the host body. Perithecia superficial, scattered on the host body, ovoid, strong orange (26A), 400–530 × 180–200 μm. Asci cylindrical, hyaline, 273–568 × 4–5 μm. Asci-caps umbonate, hyaline, 2.5–3.5 × 4–5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, (5–)6–14(–22) × 0.5–1 μm.

Colony characteristics: Colony obverse on OA compact, raised to the agar surface, cottony, entire margin, attaining a diam of 29–31 mm in 20 d, white, occasionally light yellow (14D) in the middle. Colony reverse on OA pale yellow (11C—11D). Colony obverse on PDA closely appressed to the agar surface, compact, velvet, entire margin, attaining a diam of 29–32 mm in 20 d, white, occasionally pale greenish yellow (2D) in the middle. Colony reverse on PDA moderate orange yellow (164C) and pale yellow (164D) in the middle, pale yellow (11D). Conidia and reproductive structures are not observed on both media after 30 d of incubation at 25 °C.

Additional specimen examined: Thailand, Phetchabun Province, Nam Nao District, Nam Nao National Park, Headquarters nature trail, 16°44’25”N, 101°34’24”E, on adult moth (Pyralidae), 24 Nov. 2009, A. Khonsanit, K. Tasanathai & T. Chohmee (BBH27294:BCC40747).

Notes: Phylogenetically, Ak. niveus is closely related to Ak. tortricidarum (Aini et al. 2020). However, Ak. niveus is only known from its sexual morph, while Ak. tortricidarum is known from its asexual morph. Akanthomyces niveus differs from other related species in producing whitish mycelium covering the host body and wings (Fig. 4A, B).

Akanthomyces phariformis Khons., Thanaktip. & Luangsa-ard, sp. nov. MycoBank MB 842455. Fig. 5.

Etymology: In Latin “phari” means lighthouse, referring to the form of crowded perithecia at the apex of synnemata, like a lighthouse.

Typus: Thailand, Tak Province, Umphang District, Umphang Wildlife Sanctuary, Thi Lo Su Waterfall nature trail, 15°55’36”N, 98°45’12”E, on adult of Lemyra sp. (Erebidae), 24 Nov. 2010, A. Khonsanit, K. Sansatchanon, K. Tasanathai, P. Srikitikulchai & W. Noisripoom (holotype BBH30086 preserved in a metabolically inactive state; exholotype strain BCC45148). GenBank accession numbers: ON008556 (LSU), ON00135529 (TEF1), ON0013583 (RPB2).

Asexual morph: Unknown. Sexual morph: Specimens found on adult of Lemyra sp. (Erebidae: Lepidoptera), on the upper side of dicotyledonous leaves and the leaf litter, thin whitish mycelium covering the host body. Perithecia superficial, crowded at the apex of synnemata, ovoid, light olive brown (199B), 360–550 × 200–260 μm. Asci cylindrical, hyaline, 400–470 × 5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, (3–)4–8(–11) × 0.5–1 μm.

Colony characteristics: Colony obverse on OA flat, closely appressed to the agar surface, velvety, entire margin, attaining a diam of 37–42 mm in 20 d, white. Colony reverse on OA pale yellow (162D). Colony obverse on PDA flat, closely appressed to the agar surface, velvety, entire margin, attaining a diam of 36– 41 mm in 20 d, white, pale yellow (160D), pale greenish yellow (160C). Colony reverse on PDA moderate yellow (160A) in the middle, and bluish white (N155A) along the margin. Conidia and reproductive structures are not observed on both media after 30 d of incubation at 25 °C.

Additional specimen examined: Thailand, Chiang Mai Province, Kanlayaniwatthana District, Huai Ban Rang Fish Sanctuary nature trail, 18°59’14”N, 98°17’10”E, on adult of Othreis sp. (Noctuidae), 3 Nov. 2014, A. Khonsanit, D. Thanakitpipattana, K. Tasanathai, P. Srikitikulchai, S. Wongkanoun & W. Noisripoom (BBH40632:BCC76537).

Notes: Our Akanthomyces phariformis collections produced only the sexual morph. Phylogenetically, Ak. phariformis is closely related to Ak. laosensis (Fig. 1A). However, both specimens of Ak. phariformis were found only from northern Thailand while Ak. laosensis was found only from southern Thailand. Morphologically, Ak. phariformis differs from Ak. laosensis in producing crowded superficial perithecia at the apex of synnemata (Fig. 5A, B) while Ak. laosensis produces superficial perithecia laterally at the midsection of the synnemata (Fig. 5A–D).

Akanthomyces laosensis H. Yu bis & Y. Wang, MycoKeys 101: 123. 2024. Fig. 6.

Asexual morph: Synnemata scattered, arising from different parts of the host’s body, filiform, simple, occasionally branched, white, 3–20 mm long. Conidiophores densely compacted into a layer along the synnemata, cylindrical, monophialidic or polyphialidic. Phialides smooth-walled, ampulliform, obpyriform, lageniform, hyaline, (4–)5–7(–10) × 2–3 μm. Conidia 1-celled, fusiform elliptical, elliptical, smooth-walled, hyaline, (3–)4–5(–6) × 1.5–2 μm. Sexual morph: Specimens found on adult moths (Lepidoptera), on the underside of dicotyledonous leaves, thin whitish mycelium covering the host body. Perithecia superficial, clustered along the thin whitish synnemata or solitary, ovoid, light yellow (20B), strong orange (26A—26B), (480–)527–729(– 760) × (180–)203–287(–300) μm. Asci cylindrical, hyaline, (223–)299–446(–470) × 3–5 μm. Asci-caps umbonate, hyaline, 2.5–3.5 × 3–3.5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, (3.5– )5–9(–11.5) × 0.5–1 μm.

Colony characteristics: Colony obverse on OA compact, cottony, occasionally wrinkled, raised to the agar surface, entire margin, attaining a diam of 42–43 mm in 20 d, white, produced strong greenish yellow (151A—151B) pigment. Sporulation starts at about 14 d after inoculation. Conidiophores mononematous, cylindrical, smooth-walled, hyaline. Phialides solitary, cylindrical, smooth-walled, hyaline, 10–30 × 1–2 μm. Conidia 1-celled, fusiform to elliptical, obovoid, smooth-walled, hyaline, (4–)5–9(–14) × (1.5–)2–3(–5) μm. Colony reverse on OA brilliant greenish yellow (151D). Colony obverse on PDA closely appressed to the agar surface, cottony, entire margin, attaining a diam of 38–42 mm in 20 d, white, produced brilliant greenish yellow (151D) pigment. Colony reverse on PDA dark greenish yellow (152D) in the middle, brilliant greenish yellow (151D). Conidiophores mononematous, cylindrical, smooth-walled, hyaline. Phialides solitary, cylindrical, smooth-walled, hyaline, (5–)11–23(–32) × 1–1.5 μm. Conidia 1-celled, obovoid, smooth-walled, hyaline, (2.5–)4–9(–14) × 1–3 μm.

Additional specimens examined: Thailand, Ranong Province, Kapoe District, Khlong Nakha Wildlife Sanctuary, Khlong Nakha nature trail, 9°27’33”N, 98°30’15”E, on adult moth (Lepidoptera), 11 Jan. 2006, B. Thongnuch, K. Tasanathai, L.N. Yen, L.T. Huyen, P. Srikitikulchai, S. Mongkolsamrit & W. Chaygate (BBH17548:BCC20119); Narathiwat Province, Waeng District, Hala Bala Wildlife Sanctuary, Headquarters official house nature trail, 5°48’28”N, 101°50’42”E, on adult moth (Lepidoptera), 24 Aug. 2011, A. Khonsanit (BBH31282:BCC49306); Narathiwat Province, Waeng District, Hala Bala Wildlife Sanctuary, Headquarters official house nature trail, 5°48’28”N, 101°50’42”E, on adult moth (Pyralidae), 8 Apr. 2012, A. Khonsanit, K. Tasanathai & W. Noisripoom (BBH32436:BCC52398); Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Mo Sing To nature trail, 14°26’23”N, 101°22’20”E, on adult moth (Lepidoptera), 12 Dec. 2012, A. Khonsanit, D. Thanakitpipattana, K. Tasanathai, R. Somnuk, S. Mongkolsamrit & W. Noisripoom (BBH33237:BCC58484); Narathiwat Province, Waeng District, Hala Bala Wildlife Sanctuary, Headquarters official house nature trail, 5°48’28”N, 101°50’42”E, on adult moth (Pyralis sp.: Pyralidae) 13 Oct. 2013, A. Abdulrohman, A. Khonsanit & N. Wiriyathanawudhiwong (BBH37641:BCC68341, BCC68342); Yala Province, Bannang Sata District, Bang Lang National Park, Bang Lang Dam nature trail, 6°11’56”N, 101°10’24”E, on adult moth (Lepidoptera), 30 Aug. 2014, A. Abdulrohman (MY09888:BCC74583).

Notes: Akanthomyces laosensis is a recent established species by Wang et al. (2024) found on adult moths (Noctuidae) on the underside of a dicotyledonous leaf in Oudomxay Province, Laos. Descriptions and illustrations were only from the asexual morph. Furthermore, phylogenetically, Ak. laosensis is closely related to Ak. phariformis (Fig. 1A). In our study, collections of Ak. laosensis were found only from southern Thailand, while Ak. phariformis was found only from northern Thailand. The hosts of Ak. laosensis are mostly adult moths in the family Pyralidae, while Ak. phariformis was found on different moth families, such as Erebidae and Noctuidae. Morphologically, Ak. laosensis differs from other related species by producing superficial perithecia laterally in the middle of the filiform synnemata (Fig. 6A–D), produces conidia on PDA and OA 14 d after inoculation (Fig. 6K–T). Moreover, Ak. laosensis produces a yellow pigment diffusing into the agar media (Fig. 6K, L, P, Q).

Akanthomyces noctuidarum Aini et al., MycoKeys 71: 9. 2020. Fig. 7.

Asexual morph: Synnemata scattered, arising from different parts of the host’s body, solitary, occasionally curved. Two types of synnemata were produced on adult moths: Long synnema arising from the thorax of the hosts, filiform, simple, solitary, occasionally curved, white, up to 30 mm long. Conidiophores densely compacted into a layer along the synnemata, cylindrical, mononematous. Phialides solitary, cylindrical with papillate end, monophialidic or polyphialidic, smooth-walled, hyaline, (4–)6–8(–9) × 2–3 μm. Conidia 1-celled, smooth-walled, hyaline, cylindrical with rounded end, occasionally elliptical and fusiform, (3–)4–5(–6) × 1–2 μm. Short synnemata scattered, arising from different parts of the host’s body and wing veins, solitary, cylindrical with rounded ends, white. Conidiophores densely compacted into a layer along the synnemata, cylindrical, mononematous. Phialides solitary, cylindrical with papillate ends, monophialidic or polyphialidic, smooth-walled, hyaline, (4–)5–9(–12) × 2–4 μm. Conidia 1-celled, smooth-walled, hyaline, cylindrical with rounded end, occasionally elliptical and fusiform, 4–6 × 1–2 μm. Sexual morph: Specimens found on adult moths, on the underside of dicotyledonous leaves. Perithecia superficial, scattered on the host body, crowded at the apex of synnemata, occasionally scattered on the lateral of filiform synnemata, obclavate, strong greenish yellow (151C), brilliant greenish yellow (151D), 730–1 050 × 260–450 μm. Asci cylindrical, hyaline, (280–)368–511(–793) × 4–6 μm. Ascicaps umbonate, hyaline, 3–4 × 4–5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, 5–11 × 1 μm.

Colony characteristics: Colony obverse on OA compact, raised, cottony, entire margin, attaining a diam of 25–31 mm in 20 d, white, brilliant yellow (9C) in the middle. Colony reverse on OA pale yellow green (157B), light yellowish brown (199C) in the middle. Colony obverse on PDA compact, convex, cottony, entire margin, attaining a diam of 31–33 mm in 20 d, white. Colony reverse on PDA pale greenish white (157D), brownish orange (N167A) in the middle. Conidia and reproductive structures are not observed on both media after 30 d of incubation at 25 °C.

Additional specimens examined: Thailand, Phetchaburi Province, Kaeng Krachan District, Kaeng Krachan National Park, 14°54’13”N, 99°38’03”E, on adult moth (Lepidoptera), 8 Oct. 2002, N.L. Hywel-Jones (NHJ12161); Chaiyaphum Province, Khon San District, Phu Khiao Wildlife Sanctuary, Bung pan nature trail, 16°23’10”N, 101°34’12”E, on adult moth (Lepidoptera), 24 May 2006, B. Thongnuch, K. Tasanathai, P. Srikitikulchai, S. Mongkolsamrit & W. Chaygate (BBH17614:BCC21462); Tak Province, Umphang District, Umphang Wildlife Sanctuary, Thi Lo Su Waterfall nature trail, 15°55’36”N, 98°45’11”E, on adult moth (Lepidoptera), 25 Nov. 2010, A. Khonsanit, K. Sansatchanon, K. Tasanathai, P. Srikitikulchai & W. Noisripoom (MY06426:BBH30264); Narathiwat Province, Waeng District, Hala Bala Wildlife Sanctuary, 500 nature trail, 5°48’28”N, 101°50’42”E, on adult moth (Lepidoptera), 24 Nov. 2020, A. Khonsanit & K. Tasanathai (MY12641.1:MY12641.2).

Notes: Akanthomyces noctuidarum was reported producing only one type of synnema arising from the hosts’ body and wing veins, cylindrical with papillate end phialides (5–10 × 2–3 μm), and cylindrical with rounded end conidia (3–6 × 1 μm) (Aini et al. 2020). In our study, we describe and illustrate two types of synnemata: one long synnema and numerous short synnemata (NHJ12161, BBH17614, BBH30264 & MY12641.2), collected from various regions in Thailand. However, the size and shape of phialides and conidia are the same (Table 3). Herewith we also describe and illustrate the sexual morph.

Table 3.

Morphological comparisons between sexual-morph of Akanthomyces species on moths

Species	Perithecia (μm)	Asci (μm)	Asci-caps (μm)	Part-spores (μm)	References
Ak. buriramensis	Superficial, ovoid, 500–750 × 300–400	250–450 × 3.5–4	3 × 4–5	4–14.5 × 0.5–1	This study
Ak. fusiformis	Torrubielloid, superficial, ovoid, 580–640 × 170–210	302–450 × 4–5	4.5–5.5 × 4–5	5–18 × 1	This study
Ak. laosensis	Superficial, ovoid, 480–760 × 180–300	223–470 × 3–5	2.5–3.5 × 3–3.5	3.5–11.5 × 0.5–1	This study
Ak. niveus	Torrubielloid, superficial, ovoid, 400–530 × 180–200	273–568 × 4–5	2.5–3.5 × 4–5	5–22 × 0.5–1	This study
Ak. noctuidarum	Superficial, ovoid, 530–1000 × 290–425	170–550 × 2–4	3 × 4–5	5.5–14.5 × 0.8–1	Aini et al. (2020), This study
Ak. phariformis	Superficial, ovoid, 360–550 × 200–260	400–470 × 5	−	3–11 × 0.5–1	This study
Ak. pyralidarum	Superficial, ovoid to obpyriform, 290–650 × 150–340	170–360 × 2–4	−	5–12 × 1	Aini et al. (2020)
Ak. pyralidarum		170–360 × 2–4	−	5–12 × 1	Aini et al. (2020)
Ak. pseudonoctuidarum	Superficial, ovoid, 550–750 × 220–440	263–418 × 3–5	3–4 × 4–5	4–20 × 0.5–1	This study
Ak. tuberculatus	Superficial, narrowly ovoid, dark brown, 420–900 × 180–370	300–600 × 4–5	4 μm thick	2–6 × 0.5–1	Mains (1950)

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Akanthomyces pseudonoctuidarum H. Yu bis & Y. Wang, MycoKeys 101: 124. 2024. Fig. 8.

Asexual morph: Synnemata scattered, arising from different parts of the hosts’ body, simple, occasionally curved. Two types of synnemata were observed on adult moths: Long synnemata arising from the thorax and abdomen of the hosts, filiform, simple, occasionally curved, white, (163D). Conidiophores densely compacted into a layer along the synnemata, cylindrical, mononematous. Phialides solitary, monophialidic or polyphialidic, ampulliform, obclavate, obpyriform, lageniform, smooth-walled, hyaline, (6–)8–11(–13) × 2–4 μm. Conidia 1-celled, obovoid, smooth-walled, hyaline, (3–)4–5(–6) × 1.5–2 μm. Short synnemata scattered, arising different parts of the host body, filiform, simple, solitary, occasionally curved, white, light yellow (163D), pale yellow (158A), 1–30 mm long. Conidiophores densely compacted into a layer along the synnemata, cylindrical, mononematous. Phialides solitary, monophialidic or polyphialidic, ampulliform, obclavate, obpyriform, lageniform, smooth-walled, hyaline, (4–)7–11(–17) × 2–4 μm. Conidia 1-celled, obovoid, smooth-walled, hyaline, 3–5 × 1.5–2.5 μm. Sexual morph: Specimens found on adult moths (Arctiidae, Erebidae, Lymantriidae), on the stem of living plants and underside of dicotyledonous leaves. Perithecia superficial, scattered on the host body, crowded at the apex of synnemata, occasionally scattered on the lateral of filiform synnemata, ovoid, strong orange (N163B), brownish orange (165B), (550–)604–737(–750) × (220–)266–388(–440) μm. Asci cylindrical, hyaline, (263–)311–388(–418) × 3–5 μm. Ascicaps umbonate, hyaline, 3–4 × 4–5 μm. Ascospores filiform, multi-septate, breaking into part-spores, hyaline. Part-spores cylindrical, hyaline, (4–)7–15(–20) × 0.5–1 μm.

Colony characteristics: Colony obverse on OA compact, umbonate, cottony, entire margin, attaining a diam of 29–32 mm in 20 d, white, occasionally pale greenish yellow (2D) in the middle. Colony reverse on OA pale yellow (8D). Colony obverse on PDA compact, umbonate, cottony, entire margin, attaining a diam of 31–34 mm in 20 d, white with light greenish yellow (3D). Colony reverse on PDA pale greenish yellow (2D).

Additional specimens examined: Thailand, Kamphaeng Phet Province, Klong Lan District, Klong Lan National Park, Khlong Lan Waterfall nature trail, 16°07’49”N, 99°16’35”E, on adult moth, 24 Sep. 2008, A. Khonsanit, K. Tasanathai, P. Srikitikulchai, R. Ridkaew & W. Chaygate (BBH24658:BCC32598, BCC32599); Songkhla Province, Hat Yai District, Ton Nga Chang Wildlife Sanctuary, Lamnao Phrai nature trail, 6°56’59”N, 100°14’02”E, on adult moth, 10 Aug. 2008, A. Khonsanit, B. Thongnuch, K. Tasanathai, P. Srikitikulchai, S. Mongkolsamrit & W. Chaygate (BBH25301:BCC32161); Nakhon Si Thammarat Province, Nopphitam District, Khao Luang National Park, Krung Ching Waterfall nature trail, 8°43’06”N, 99°41’29”E, on adult moth, 17 Feb. 2009, K. Tasanathai, P. Srikitikulchai, R. Promharn, S. Mongkolsamrit & T. Chohmee (BBH26442:BCC35979, BCC35980); Phetchabun Province, Nam Nao District, Nam Nao National Park, Headquarters nature trail, 16°44’25”N, 101°34’24”E, on adult moth, 19 Feb. 2009, K. Tasanathai, P. Srikitikulchai, S. Mongkolsamrit & T. Chohmee (BBH26449:BCC36300, BCC36301); Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Mo Sing To nature trail, 14°26’20”N, 101°22’19”E, on adult moth, 19 Jul. 2009, K. Tasanathai, P. Srikitikulchai, R. Ridkaew, S. Mongkolsamrit & T. Chohmee (BBH26350:BCC37665:BCC37666); Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Mo Sing To nature trail, 14°26’20”N, 101°22’19”E, on adult moth, 19 Jul. 2009, K. Tasanathai, P. Srikitikulchai, R. Ridkaew, S. Mongkolsamrit & T. Chohmee (BBH26350:BCC37665:BCC37666); ibid., on adult moth, 14 Aug. 2009. K. Tasanathai, P. Srikitikulchai, R. Ridkaew, S. Mongkolsamrit & T. Chohmee (BBH26530:BCC37916); ibid., on adult moth, 19 Sep. 2009, K. Tasanathai, P. Srikitikulchai, R. Ridkaew, S. Mongkolsamrit & T. Chohmee (BBH26583:BCC39698); ibid., on adult moth (Lymantriidae), 7 Jul. 2011, A. Khonsanit, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH30740:BCC48709); Chiang Mai Province, Chiang Dao District, Ban Huathung community forest nature trail, 19°25’07”N, 98°57’21”E, on adult moth (Arctiidae), 16 Jul. 2011, A. Khonsanit, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai & S. Mongkolsamrit (BBH26350:BCC37665, BCC37666); ibid., on adult moth, 31 Aug. 2011, A. Khonsanit, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH31342:BCC49336); Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Chao Ying nature trail, 14°26’20”N, 101°22’19”E, on adult moth (Lymantriidae), 9 Aug. 2012, A. Khonsanit, K. Tasanathai, K. Sansatchanon, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH35668:BCC54481, BCC54889); Chiang Rai Province, Phan District, Doi Luang National Park, Pu Kaeng Waterfall nature trail, 19°27’02”N, 99°43’00”E, on Erebus macrops (Erebidae), 12 Nov. 2013, C. Suriyachadkun (BBH38012:BCC69011); Nakhon Ratchasima Province, Pak Chong District, Khao Yai National Park, Kong Kaeo Waterfall nature trail, 14°26’21”N, 101°22’25”E, on adult moth (Lymantriidae), 28 Nov. 2013, A. Khonsanit, D. Thanakitpipattana, P. Srikitikulchai, S. Mongkolsamrit & W. Noisripoom (BBH37920:BCC69056); Chiang Mai Province, Chiang Dao District, Ban Huathung community forest nature trail, 19°25’07”N, 98°57’21”E, on adult moth (Arctiidae), 29 Oct. 2014, A. Khonsanit, D.Thanakitpipattana, K. Tasanathai, P. Srikitikulchai, S. Wongkanoun & W. Noisripoom, (BBH40206:BCC76478); ibid., on adult moth (Arctiidae), 30 Oct. 2014, A. Khonsanit, D.Thanakitpipattana, K. Tasanathai, P. Srikitikulchai, S. Wongkanoun & W. Noisripoom, (BBH40318:BCC75746, BCC76483); Nakhon Si Thammarat Province, Nopphitam District, Khao Luang National Park, Krung Ching Waterfall nature trail, 8°43’06”N, 99°41’29”E, on adult moth, 26 Jan. 2016, D.Thanakitpipattana, K. Tasanathai, P. Srikitikulchai, R. Promharn, S. Mongkolsamrit, S. Wongkanoun & W. Noisripoom (BBH41238:BCC80356).

Notes: Akanthomyces pseudonoctuidarum is a recently described species by Wang et al. (2024) found on adult moths (Noctuidae) on the underside of leaves from Chiang Mai Province, Thailand. However, they reported only from the asexual morph. In our study, we describe and illustrate the asexual and sexual morph from the same specimen, and the presence of two types of synnemata: a long synnema and short synnemata. We found additional moth families Arctiidae, Erebidae and Lymantriidae as their hosts. Phylogenetically, Ak. pseudonoctuidarum is closely related to Ak. buriramensis (Fig. 1A). Morphologically, Ak. pseudonoctuidarum differs from other related species by producing superficial perithecia at the basal part of the long filiform synnemata (Fig. 8C) and superficial perithecia at the tip of short synnemata (Fig. 8D).

Keys to species of Akanthomyces on moth

Based on asexual morph characters

1a. No synnema, mononematous .............................................................................................................................................................. Ak. fusiformis
1b. Produce two types of synnemata .............................................................................................................................................................................. 2
1c. Produces one type of synnemata .............................................................................................................................................................................. 3
2a. Phialides cylindrical with papillate end, 4–12 × 2–4 μm, conidia cylindrical with round end, ellipsoid, 4–6 × 1–2 μm .................. Ak. noctuidarum
2b. Phialides cylindrical with papillate end, ampulliform, obclavate, obpyriform, lageniform, 4–17 × 2–4 μm, ................................................................ conidia ellipsoidal to long oval, obovoid, 2.6–6.4 × 1.5–2.5 μm ........................................................................................... A. pseudonoctuidarum
2c. Phialides cylindrical to ellipsoidal with papillate end, 5–10 × 1.8–3 μm, conidia fusoid, 1–3.2 × 1–2 μm ..................................... Ak. tortricidarum
3a. Phialides ɥ 10 μm long subcylindrical, narrowly ellipsoidal, 6–16 × 2.5–4 μm, conidia ellipsoid, obovoid, 3–6 × 2–3 μm ................... Ak. aculeatus
3b. Phialides ≤ 10 μm long .............................................................................................................................................................................................. 4
4a. Conidia ellipsoid, fusiform, obovoid, 3–5 × 1–2 μm, found only in Buri Ram Province, Thailand ................................................... Ak. buriramensis
4b. Conidia fusiform elliptical, elliptical, 3–6 × 1.5–2 μm, produced yellow pigment into agar media, found only in ........................................................ Southern Thailand .................................................................................................................................................................................. Ak. laosensis
4c. Conidia cylindrical, narrowly fusiform, 4.5–6 × 1.2–1.5 μm, found in Ghana ................................................................................... Ak. tuberculatus

Based on sexual morph characters

1a. Produces torrubielloid perithecia .............................................................................................................................................................................. 2
1b. Produces superficial perithecia on the synnemata ................................................................................................................................................... 3
2a. Asci-caps cone-shaped, 4.5–5.5 × 4–5 μm ........................................................................................................................................... Ak. fusiformis
2b. Asci-caps umbonate, 2.5–3.5 × 4–5 μm ..................................................................................................................................................... Ak. niveus
3a. Produces only crowded perithecia at the apex of synnemata ........................................................................................................... Ak. phariformis
3b. Produces only superficial perithecia in the middle of synnemata ............................................................................................................................. 4
3c. Produces crowded perithecia at the apex and superficial perithecia in the middle of synnemata ........................................................................... 5
4a. Perithecia crowded in the middle of short filiform synnemata ....................................................................................................... Ak. buriramensis
4b. Perithecia clustered in the middle of long filiform synnemata ............................................................................................................... Ak. laosensis
5a. Perithecia ≥ 750 μm long ........................................................................................................................................................................................... 6
5b. Perithecia ≤ 750 μm long ........................................................................................................................................................................................... 7
6a. Perithecia ovoid, 530–1000 × 290–425 μm. Part-spore cylindrical 5.5–14.5 × 0.8–1 μm ............................................................... Ak. noctuidarum
6b. Perithecia narrowly ovoid, 420–900 × 180–370 μm. Part-spores cylindrical 2–6 × 0.5–1 μm .......................................................... Ak. tuberculatus
7a. Perithecia ovoid to obpyriform, 290–650 × 150–340 μm. Part-spores cylindrical 5–12 × 1 μm ...................................................... Ak. pyralidarum
7b. Perithecia ovoid, 550–750 × 220–440 μm. Part-spores cylindrical 4–20 × 0.5–1 μm ........................................................... Ak. pseudonoctuidarum

Residual species of Akanthomyces

The following species of Akanthomyces s. l. could not be confidently assigned in the new classification because they were not sampled as part of our phylogenetic analyses. These species are provisionally retained within Akanthomyces s. l. until further molecular phylogenetic studies are made to classify them in a phylogenetic system, or when we find the sexual morph connection for the species of Akanthomyces s. l.

Akanthomyces angustisporus Mains, Mycologia 42: 573. 1950.

Akanthomyces aranearum (Petch) Mains, Mycologia 42: 574. 1950.

Basionym: Hymenostilbe aranearum Petch, Trans. Brit. Mycol. Soc. 16: 221. 1932.

Akanthomyces clavata (Mains) K.T. Hodge, In: White et al., Clavicipitalean Fungi: Evolutionary Biology, Chemistry, Biocontrol, and Cultural Impacts (New York): 87. 2003.

Basionym: Insecticola clavata Mains, Mycologia 42: 577. 1950.

Akanthomyces fragilis (Petch) (Mains) K.T. Hodge, In: White et al., Clavicipitalean Fungi: Evolutionary Biology, Chemistry, Biocontrol, and Cultural Impacts (New York): 87: 2003.

Basionym: Hymenostilbe fragilis Petch Trans. Brit. Mycol. Soc. 21: 56. 1937.

Akanthomyces gracilis Samson & H.C. Evans, Acta Bot. Neerl. 23: 29. 1974.

Akanthomyces johnsonii (Massee) Vincent et al., Mycologia 80: 685. 1988.

Basionym: Pistillaria johnsonii Massee, Bull. Misc. Inf., Kew: 165. 1901.

Novel genera and new combinations

Arachnidicola Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 848891.

Etymology: Refers to the hosts of this genus that is colonizes — spiders (Araneae).

Typus: Arachnidicola sulphurea (Mongkol. et al.) Khons., Thanakitp. & Luangsa-ard

Diagnosis: Isaria-like or lecanicillium-like. Synnemata numerous, covered by dense white to cream, or yellowish white mycelia bearing numerous conidiophores with powdery conidia. Conidiophores loosely grouped together, arising from different parts of the spider, verticillate with phialides in whorls of two to five. Phialides hyaline, cylindrical to ellipsoidal basal portion, tapering into a thin, distinct neck, somewhat inflated base. Conidia in chains, hyaline, cylindrical to ellipsoidal, fusiform to ellipsoidal (Fig. 9B). Sexual morph: Stromata consisting of dense hyphae, completely covering the host. Perithecia torrubiellalike, scattered on the abdominal part of the host, dark orange, deep yellow, superficial, narrowly ovoid. Asci hyaline, cylindrical, 8-spored, apex and bottom of asci thicker than the middle part. Ascospores hyaline, cylindrical, filiform dissociating into part-spores.

Fig. 9 — Types of conidiogenous cells and conidia of known and new genera. A. *Akanthomyces aculeatus* on host after Seifert et. *al.* (2011), plate 84C. B. *Arachnidicola sulphurea* on culture media adapted from fig. 3 in Mongkolsamrit et al. (2018). C. *Corniculantispora psalliotae* on culture media adapted from fig. 12 in Zare & Gams (2001). D. *Corpulentispora magnispora* on culture media adapted from fig. 30 in Zhang *et al*. (2020). E. *Kanoksria zaquensis* on culture media adapted from fig. 3 in Wang *et al*. (2023). F. *Lecanicillium lecanii* on culture media adapted from fig. 3 in Zare & Gams (2001). G. *Zarea fungicola* on culture media adapted from fig. 3 in Zare & Gams (2008). H. *Zouia cauligalbarum* on culture media adapted from fig. 3 in Zhou *et al.* (2018). Line drawings by P. Wongpitakchai (B–E, G–H).

Habitat: On spiders (Araneae).

Distribution: Thailand and China.

Arachnidicola araneicola (W.H. Chen et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 849875.

Basionym: Akanthomyces araneicola W.H. Chen et al., Phytotaxa 409: 228. 2019.

Arachnidicola araneogena (Z.Q. Liang et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 849877.

Basionym: Akanthomyces araneogenus Z.Q. Liang et al., Phytotaxa 379: 69. 2018.

Arachnidicola bashanensis (W.H. Chen et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852005.

Basionym: Akanthomyces bashanensis W.H. Chen et al., MycoKeys 98: 305. 2023.

Arachnidicola beibeiensis (W.H. Chen et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852006.

Basionym: Akanthomyces beibeiensis W.H. Chen et al., MycoKeys 98: 307. 2023.

Arachnidicola kanyawimiae (Mongkol. et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 849878.

Basionym: Akanthomyces kanyawimiae Mongkol. et al., Mycologia 110: 237. 2018.

Arachnidicola kunmingensis (H. Yu bis et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852008.

Basionym: Akanthomyces kunmingensis Hong Yu bis et al., MycoKeys 101: 121. 2024.

Arachnidicola subaraneicola (H. Yu bis et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852009.

Basionym: Akanthomyces subaraneicola H. Yu bis et al., MycoKeys 101: 126. 2024.

Arachnidicola sulphurea (Mongkol. et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 849883.

Basionym. Akanthomyces sulphureus Mongkol. et al., Mycologia 110: 237. 2018.

Arachnidicola thailandica (Mongkol. et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 849879.

Basionym: Akanthomyces thailandicus Mongkol. et al., Mycologia 110: 240. 2018.

Arachnidicola tiankengensis (W.H. Chen et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852012.

Basionym: Akanthomyces tiankengensis W.H. Chen et al., Microbiol. Spectr. 10: 6. 2022.

Arachnidicola waltergamsii (Mongkol. et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 852013.

Basionym: Akanthomyces waltergamsii Mongkol et al., Mycologia 110: 241. 2018.

Key to species of Arachnidicola on spiders

1a. Produces lecanicillium-like asexual morph ................................................................................................................................................................ 2
1b. Produces isaria-like asexual morph ........................................................................................................................................................................... 3
2a. Conidia cylindrical to ellipsoid, (3–5 × 1–2 μm), found in Thailand ..................................................................................................... Ar. thailandica
2b. Conidia obovoid, sub-globose, ellipsoid, (2–3 × 1–2 μm), found in China ...................................................................................... Ar. tiankengensis
3a. Phialides cylindrical, inflated base, tapering into a thin neck, ≤ 12 μm long ............................................................................................................. 4
3b. Phialides cylindrical, inflated base, tapering into a thin neck, ≥ 12 μm long ............................................................................................................. 7
4a. Conidia ellipsoid, found in Thailand ...................................................................................................................................................... Ar. sulphurea
4b. Conidia fusiform to ellipsoid, found in China ............................................................................................................................................................ 5
4c. Conidia sub-globose, ellipsoid, obovoid .................................................................................................................................................................... 6
5a. Conidia fusiform to ellipsoid, 1.7–2.6 × 1.6–1.8 μm ......................................................................................................................... Ar. bashanensis
5b. Conidia fusiform to ellipsoid, 2.0–3.3 × 2.0–2.6 μm ........................................................................................................................... Ar. beibeiensis
6a. Spider host covered with dense white to cream mycelia, found in Thailand ................................................................................. Ar. waltergamsii
6b. Spider host covered with dense white to cream mycelia with powdery conidia, found in Thailand .............................................. Ar. kanyawimiae
7a. Conidia ellipsoid, obovoid (2–5 × 1–2 μm), found in China ..................................................................................................................Ar. araneicola
7b. Conidia ellipsoid, globose (2–3 × 1–2 μm), found in China ............................................................................................................. Ar. araneogenum
7c. Conidia ellipsoid, to long oval (1.9–3.5 × 1.1–1.8 μm), found in China .......................................................................................... Ar. kunmingensis
7d. Conidia ellipsoidal to long oval (3.0–5.4 × 1.8–3.4 μm), found in China .........................................................................................Ar. subaraneicola

Corniculantispora Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 851978.

Etymology: From the Latin “corniculantis” means crescent-shaped, refers to the shape of the macroconidia produced in this genus.

Typus: Corniculantispora psalliotae (Treschew) Khons., Thanakitp. & Luangsa-ard

Diagnosis: Conidiogenous cells aculeate, aphanophialides, rather long, inflated, verticillate, arising from undifferentiated prostrate conidiophores, first flask-shaped, tapering into thread-like neck, but soon collapsing and becoming reduced to very fine denticles with solitary conidia still remaining at apex, solitary or more often 3 or 6 in whorls. Conidia dimorphic, can be macro- and microconidia formed from phialides, typically solitary or in small clusters at right angles to phialide tip, commonly falcate with pointed ends, subsequent ones oval to ellipsoidal. Macroconidia crescent-shaped, slightly curved, usually with sharply pointed ends, 1-celled. Microconidia formed subsequently, ellipsoidal to fusoid with round ends, oval to ellipsoidal, sub globose, nearly straight to slightly curved. Octahedral crystals present or absent (Fig. 9C). Sexual morph: Unknown.

Habitat: The genus is found on a wide range of hosts including Araneae, cicadellid on leaf of Theobroma cacao, collembola, eggs of Meloidogyne, greenhouse whitefly (Trialeurodes vaporariorum), mites, mosquito larvae, nymphs of Ixodes (tick), silk moth (Bombyx mori) and associated with Agaricus bisporus and A. bitorquis, Puccinia coronata, Sphaerotheca fuliginea, S. pannosa. It was also reported from leaf litter and soil.

Distribution: Bulgaria, China, Cuba, Germany, Ghana, Netherlands, India, Indonesia, Israel, Sri Lanka, South Africa, Spain, UK, USA.

Corniculantispora aranearum (Petch) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851984.

Basionym: Acremonium aranearum Petch, Trans. Brit. Mycol. Soc. 16: 242. 1932.

Synonyms: Aphanocladium aranearum (Petch) W. Gams, Cephalosporium-artige Schimmelpilze: 198. 1971. Aphanocladium aranearum var. sinense C.D. Chen, Acta Mycol. Sin. 3: 96. 1984.

?Acremonium fimicola Massee & E.S. Salmon, Ann. Bot. 16: 79. 1902.

?Sporotrichum roseolum Oudem. & Beij., Ned. Kruidk. Arch. 3, 2: 910. 1903.

Lecanicillium aphanocladii Zare & W. Gams, Nova Hedwig. 73: 27. 2001.

Corniculantispora dimorpha (J.D. Chen). Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851985.

Basionym: Aphanocladium dimorphum J.D. Chen, Acta Mycol. Sin. 4: 230. 1985.

Synonym: Lecanicillium dimorphum (J.D. Chen) Zare & W. Gams, Nova Hedwig. 73: 24. 2001.

Corniculantispora psalliotae (Treschew) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851987.

Basionym: Verticillium psalliotae Treschew, Dan. Bot. Arkiv. 11(1): 7. 1941.

Synonym: Lecanicillium psalliotae (Treschew) Zare & W. Gams, Nova Hedwig. 73: 21. 2001.

Corpulentispora Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 851994.

Etymology: Latin “Corpulentus” means large or larger, refers to its large conidia.

Typus: Corpulentispora magnispora (Z.F. Zhang & L. Cai) Khons., Thanakitp. & Luangsa-ard

Diagnosis: Mycelium hyaline, septate, smooth. Conidiophores arising from aerial hyphae, erect, smooth, hyaline. Phialides arising from aerial hyphae solitary, or in whorls of 2–5 at the apex of conidiophores, straight or slightly curved, tapering to the apex, smooth, hyaline. Conidia rare, unicellular, smooth, hyaline, variable in size. Macroconidia long fusiform or falcate. microconidia much fewer than macroconidia (Fig. 9D). Sexual morph: Unknown.

Habitat: Soil.

Distribution: China.

Corpulentispora magnispora (Z.F. Zhang & L. Cai) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851995.

Basionym: Lecanicillium magnisporum Z.F. Zhang & L. Cai, Fungal Divers. 106: 75. 2020.

Synonym: Gamszarella magnispora (Z.F. Zhang & L. Cai) Crous, Persoonia 51: 390. 2023.

Kanoksria Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 851982.

Etymology: Kanoksria is named in honour of Miss Kanoksri Tasanathai, who is working on invertebrate pathogenic fungi research in BIOTEC for more than 20 years.

Typus: Kanoksria zaquensis (Y.H. Wang et al.) Khons., Thanakitp. & Luangsa-ard

Diagnosis: The vegetative hyphae delicate, hyaline, smooth-walled. Phialides occurring directly on the aerial hyphae, simple. Conidia produced on PDA, one-celled, smooth-walled, hyaline, adhering in heads at the apex of the phialides, long-ellipsoidal to almost cylindrical (Fig. 9E). Sexual morph: Unknown.

Habitat: Stroma and the sclerotium of O. sinensis on the ground.

Distribution: China.

Kanoksria zaquensis (Y.H. Wang et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851983.

Basionym: Akanthomyces zaquensis Y.H. Wang et al., Phytotaxa 579: 203. 2023.

Lecanicillium (Zimm) Zare & Gams, Nova Hedwig. 72: 50. 2001.

Refers to members of the clade B of our phylogenetic analyses consisting of the type of the genus Lecanicillium (L. lecanii) and other Lecanicillium species. We propose that the genus Lecanicillium should be resurrected to accommodate species in this clade.

Typus: Lecanicillium lecanii (Zimm.) Zare & W. Gams

Diagnosis: Conidiophores arising from aerial hyphae, prostrate and occasionally from the subtending hyphae, erect short bearing one or two whorls phialides. Phialides discrete, aculeate, tapering to a narrow tip, in which collarette and periclinal wall thickening are hardly visible, verticillate or solitary. Conidia adhering in slimy heads or fascicles, sometimes forming chains, short to long ellipsoidal to falcate with point ends (Fig. 9F, Zare & Gams 2001). Sexual morph: White to cream thin mycelium covering on scale insect, slightly tufted, pulverulent. Perithecia irregularly scattered to crowded, superficial or slightly embedded, ovoid, reddish brown to dark chestnut brown. Asci cylindrical, thickened at the apex. Ascospores filiform, multiseptate (Mains 1949).

Habitat: On scale insects [e.g. Pulvinaria floccifera, Coccus viridis (Coccidae)] on the underside of leaves of forest plant, ladybug (Coleoptera), spider (Araneae), lepidopteran pupa and adult moths (Lepidoptera), weevil (Coleoptera) and soil.

Distribution: China, Canada, Colombia, Dominican Republic, Germany, Indonesia, Iran, Jamica, Kenya, Mexico, New Zealand, Peru, South Africa, Sri Lanka, Turkey, UK, and USA.

Lecanicillium attenuatum Zare & W. Gams, Nova Hedwig. 73: 19. 2001.

Synonym: Akanthomyces attenuatus (Zare & W. Gams) Spatafora et al., IMA Fungus 8: 342. 2017.

Lecanicillium araneogenum Z.Q. Liang et al., Phytotaxa 379: 69. 2018.

Synonym: Akanthomyces neoaraneogenus (Z.Q. Liang et al.), Shrestha et al., Mycol. Progr. 18: 986. 2019.

Lecanicillium araneosum (W.H. Chen et al.) Khons. et al., comb. nov. MycoBank MB 848900.

Basionym: Akanthomyces araneosus W.H. Chen et al., Microbiol. Spectr. 10: 6. 2022.

Lecanicillium lecanii (Zimm.) Zare & Gams, Nova Hedwig. 72: 51. 2001.

Basionym: Cephalosporium lecanii Zimm., Teysmannia 9: 241. 1899.

Synonyms: Verticillium lecanii (Zimm.) Viégas, Revista Inst. Café São Paulo 14: 754. 1939.

Akanthomyces lecanii (Zimm.) Spatafora et al., IMA Fungus 8: 343. 2017.

Torrubiella confragosa Mains, Mycologia 41: 305. 1949.

Cordyceps confragosa (Mains) G.H. Sung et al., Stud. Mycol. 57: 49. 2007.

Hirsutella confragosa Mains, Mycologia 41: 305. 1949.

Lecanicillium lepidopterorum (W.H. Chen et al.) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 848901.

Basionym: Akanthomyces lepidopterorum W.H. Chen et al., Phytotaxa 459: 121. 2020.

Lecanicillium longisporum (Petch) Zare & W. Gams, Nova Hedwig. 73: 16. 2001.

Basionym: Cephalosporium longisporum Petch, Trans. Brit. Mycol. Soc. 10: 166. 1925.

Synonyms: Cephalosporium dipterigenum Petch, Naturalist (Hull) 56: 102. 1931.

Akanthomyces dipterigenus (Petch) Spatafora et al., IMA Fungus 8: 343. 2017.

Lecanicillium muscarium (Petch) Zare & W. Gams, Nova Hedwig. 73: 13. 2001.

Basionym: Cephalosporium muscarium Petch, Naturalist (Hull), ser. 3: 102. 1931.

Synonym: Akanthomyces muscarius (Petch) Spatafora et al., IMA Fungus 8: 343. 2017.

Lecanicillium neocoleopterorum (W.H. Chen et al.), Khons., Thanakitp., Mongkols., Kobmoo & Luangsa-ard, comb. nov. MycoBank MB 848903.

Basionym: Akanthomyces neocoleopterorum W.H. Chen et al., Phytotaxa 432: 122. 2020.

Lecanicillium pissodis Kope & I. Leal, Mycotaxon 94: 334. 2006.

Synonym: Akanthomyces pissodis (Kope & I. Leal) W.H. Chen et al., Phytotaxa 459: 120. 2020.

Lecanicillium sabanense Chir.-Salom. et al., Phytotaxa 234: 68. 2015.

Synonym: Akanthomyces sabanensis (Chir.-Salom. et al.) Chir.-Salom. et al., IMA Fungus 8: 343. 2017.

Lecanicillium uredinophilum M.J. Park et al., Mycotaxon 130: 997. 2015.

Synonym: Akanthomyces uredinophilus (M.J. Park et al.) Manfrino & Leclerque, Diversity 118: 5. 2022.

The following Lecanicillium species were placed in the genus based on morphological characteristics. We retain them in Lecanicillium pending future phylogenetic analyses:

Lecanicillium aranearum (Petch) Zare & W. Gams, Nova Hedwig. 73: 30. 2001.

Basionym: Cephalosporium aranearum Petch, Trans. Brit. Mycol. Soc. 16: 226 .1932.

Lecanicillium evansii Zare & W. Gams, Nova Hedwigia Beih. 73: 32. 2001.

Lecanicillium fusisporum (W. Gams) Zare & W. Gams, Nova Hedwig. 73: 34. 2001.

Basionym: Verticillium fusisporum W. Gams, Cephalosporiumartige Schimmelpilze (Stuttgart): 182. 1971.

Lecanicillium nodulosum (Petch) Zare & W. Gams, Nova Hedwig. 73: 18. 2001.

Basionym: Cephalosporium nodulosum Petch, Trans. Brit. Mycol. Soc. 23: 144. 1939.

Lecanicillium tenuipes (Petch) Zare & W. Gams, Nova Hedwig. 73: 29. 2001.

Basionym: Acremonium tenuipes Petch, Trans. Brit. Mycol. Soc. 21: 66. 1937.

Synonyms: Verticillium tenuipes (Petch) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 176. 1971.

Sporotrichum aranearum Cavara, Fung. Long. Exsicc. 5: 240. 1895.

Engyodontium aranearum (Cavara) W. Gams et al., Persoonia 12: 138. 1984.

Note: Since there is already a species called Lecanillium aranearum, Zare & Gams (2001) used the next oldest epithet available in order to avoid homonymy.

Lecanicillium saksenae (Kushwaha) Kurihara & Sukarno, Mycoscience 50: 377. 2009.

Basionym: Verticillium saksenae Kushwaha, Curr. Sci. 49: 948. 1980.

Keys to species of Lecanicillium

Based on asexual morph characters

1a. Found on Araneae (spider) ......................................................................................................................................................................................... 2
1b. Found on Coleoptera ................................................................................................................................................................................................. 4
1c. Found on Lepidoptera .................................................................................................................................................................... L. lepidopterorum
1d. Found on Hemiptera .................................................................................................................................................................................................. 5
1e. Found on a range of hosts ..........................................................................................................................................................................................8
2a. Presence of octahedral crystals ............................................................................................................................................................. L. aranearum
2b. Absence of octahedral crystals ...................................................................................................................................................................................3
3a. Conidia fusiform (3.1–5.0 × 1.0–1.8 μm) ................................................................................................................................................ L. araneosus
3b. Conidia cylindrical (3.2–8.6 × 1.3–1.6 μm) ....................................................................................................................................... L. araneogenum
3c. Conidia ovoid to ellipsoid (3–6.5 × 1–1.5 μm) ........................................................................................................................................... L. tenuipes
4a. Found on rove beetles; conidia fusoid with pointed ends, slightly curved; found in Indonesia .....................................................L. kalimantanense
4b. Found on ladybugs; conidia cylindrical; found in China ............................................................................................................ L. neocoleopterorum
4c. Found on weevil; conidia cylindrical to ovoid; found in Canada ................................................................................................................. L. pissodis
5a. Phialide cylindrical, ≤ 20 μm long (13–19 × 2–4 μm) .............................................................................................................................. L. sabanense
5b. Phialide cylindrical, ≥ 20 μm long .............................................................................................................................................................................. 6
6a. Octahedral crystals absent ............................................................................................................................................................... L. uredinophilum
6b. Octahedral crystals present ....................................................................................................................................................................................... 7
7a. Phialides produced 1−5 per node (15–35 × 1–1.7 μm) ................................................................................................................................................. Conidia ellipsoid to subcylindrical (2–6 × 1–1.8 μm) ............................................................................................................................. L. muscarium
7b. Phialides produced 1−6 per node (11–30 × 1.3–1.8 μm) ............................................................................................................................................. Conidia short ellipsoid to subcylindrical (2.5–4.2 × 1–1.5 μm) ..................................................................................................................... L. lecanii
7c. Phialides produced 5−6 per node (20–40 × 1.2–2.7 μm) .............................................................................................................................................. Conidia ellipsoid to oblong-oval (5–10.5 × 1.5–2.5 μm) ..................................................................................................................... L. longisporum
8a. Octahedral crystals absent ........................................................................................................................................................................ L. saksenae
8b. Octahedral crystals present ....................................................................................................................................................................................... 9
9a. Conidia monomorphic ............................................................................................................................................................................................. 10
9b. Conidia dimorphic ................................................................................................................................................................................................... 11
10a. Conidia fusiform (3–5 × 1.5–2 μm) ...................................................................................................................................................... L. fusisporum
10b. Conidia ovoid (2.5–4.5 × 1.2–1.5 μm) ................................................................................................................................................. L. nodulosum
11a. Macroconidia fusiform, with very narrow and pointed ends (11–18 × 0.8–1.5 μm) ................................................................................................... Microconidia ellipsoid (3–4 × 0.8–1.2 μm) .......................................................................................................................................... L. antillanum
11b. Macroconidia slightly falcate with pointed ends (4.5–7.5 × 0.8–1.2 μm) ................................................................................................................... Microconidia ellipsoid with pointed ends, slightly curved (2–3 × 0.8–1.2 μm) .......................................................................................... L. evansii

Zarea Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 851990.

Etymology: Zarea is named in honour of Dr Rasoul Zare who, together with Dr Walter Gams, contributed immensely to our understanding of the genus Lecanicillium.

Typus: Zarea fungicola (Preuss) Khons., Thanakitp. & Luangsa-ard

Diagnosis: Conidiophores arising typically from submerged hyphae, erect (later sometimes also procumbent), thick-walled, with 2–5 (to over 10 in old colonies) whorls of 3–7 phialides. Phialides arising at an oblique angle from the conidiophore stipe. Conidia adhering in mostly globose, slimy heads, the heads of single phialides often coalescing to form large slimy masses, fusiform, long-ellipsoidal to almost cylindrical, the basal end indistinctly truncate, sometimes curved along the longitudinal axis, asymmetrically biconvex to concave-convex to slightly sickle-shaped, smooth-walled, of very unequal size, with 1–2 or more inconspicuous guttules (Fig. 9G). On mushrooms the ellipsoidal form of conidia may dominate, in vitro the fusiform shape is commonly present. Sporulation abundant. Chlamydospores absent. Octahedral crystals present or absent. Sexual morph: Unknown.

Habitat: The genus is associated with Agaricus bisporus, Agaricus bitorquis, Allopsalliota geesterani, Coltricia perennis, Gomphidius glutinosus, Henningsomyces candidus, Hypholoma capnoides, Laccaria laccata, Marasmiellus ramealis, Russula nigricans, Thelephora terrestris, unidentified agarics, leaf litter and soil.

Distribution: Australia, Belgium, Germany, Finland, France, Iran, Netherlands, Scotland, UK, USA.

Notes: Zare & Gams (2008) revised the Verticillium fungicola species complex and noted the affinity of this complex to Lecanicillium. The authors proposed the transfer of members of the species complex to Lecanicillium but elevated only var. flavidum to species level while retaining var. fungicola and var. aleophilum, citing indistinguishable morphological characters and minimal differences in ITS sequences. In our multi-gene analyses, however, var. fungicola and var. aleophilum could be seen as distinct a species as Z. flavidum hence we propose to elevate these two varieties to species level as members of the genus Zarea (Fig 1B clade 4).

Zarea aleophila (W. Gams & Zaayen) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851992.

Basionym: Verticillium fungicola var. aleophilum W. Gams & Zaayen, Neth. J. Pl. Path. 88: 62. 1982.

Synonym: Lecanicillium fungicola var. aleophilum (W. Gams & Zaayen) W. Gams & Zare, Mycol. Res. 112: 821. 2008.

Zarea flavida (W. Gams & Zaayen) Khons., Thanakitp. & Luangsaard, comb. nov. MycoBank MB 853004.

Basionym: Verticillium fungicola var. flavidum W. Gams & Zaayen, Neth. J. Pl. Path. 88: 64. 1982.

Synonym: Lecanicillium flavidum (W. Gams & Zaayen) W. Gams & Zare, Mycol. Res. 112: 821. 2008.

Zarea fungicola (Preuss) Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851993.

Basionym: Acrostalagmus fungicola Preuss, Linnaea 24: 126. 1851.

Synonyms: Verticillium fungicola var. fungicola (Preuss) Hassebrauk, Phytopath. Z. 9: 514. 1936.

Lecanicillium fungicola var. fungicola (Preuss) Zare & W. Gams, Mycol. Res. 112: 818. 2008.

Cephalosporium costantinii F.E.V. Smith, Trans. Brit. Mycol. Soc. 10: 90. 1924.

Verticillium malthousei Ware, Ann. Bot. 47: 781. 1933.

Zouia Khons., Thanakitp. & Luangsa-ard, gen. nov. MycoBank MB 851980.

Etymology: Zouia is named in honour of Dr Xiao Zou who collected the specimens and described the species Lecanicillium cauligalbarum (2018) on stemborer (Lepidoptera).

Typus: Zouia cauligalbarum (Zou et al.), Khons., Thanakitp. & Luangsaard

Diagnosis: Mycelium hyaline, smooth, septated, branched. Conidiophores usually arising from aerial hyphae, sporulate abundant. Phialides gradually tapering towards the apex, solitary or 2–3 whorls. Conidia cylindric, aseptate. The phialides and conidia on cultures are of similar general shape and size to those found on the host stemborer (Fig. 9H).

Sexual morph: Unknown.

Habitat: Stemborer (Lepidoptera) hidden amongst pieces of wood in humid forests.

Distribution: China.

Zouia cauligalbarum (X. Zou et al.), Khons., Thanakitp. & Luangsa-ard, comb. nov. MycoBank MB 851981.

Basionym: Lecanicillium cauligalbarum X. Zou et al., MycoKeys 43: 65. 2018.

DISCUSSION

In this study, we conducted a comprehensive phylogenetic and morphological examination of entomopathogenic fungi presumptively belonging to the genus Akanthomyces, discovered on moths in Thailand. We examined 15 strains of Akanthomyces isolated from adult moths using both morphological characters and molecular phylogenetic analyses. We also integrated data from prior studies into our analyses. The results of the phylogenetic analyses unveiled that Akanthomyces spp., collected from diverse insect hosts (adult moths, ladybugs, lepidopteran larvae, scale insects, whiteflies, spiders, and weevils), as well as from leaf litter, fungus, and soil, were distinctly segregated into four monophyletic groups (Akanthomyces, Arachnidicola, Lecanicillium, Kanoksria). Each group exhibited variations in host range and the diversity of asexual morphs produced.

The Akanthomyces clade was exclusively associated with moth infections, the Arachnidicola clade was identified solely from spiders, while Lecanicillium demonstrated versatility across various substrates, and Kanoksria was found exclusively on Ophiocordyceps sinensis. Consequently, the type of substrate or the taxonomic group of the host could serve as a valuable criterion for fungal taxonomic classification at the genus level. The study suggests that host specificity to insects provides highly informative characteristics for species identification of invertebrate pathogenic fungi (Johnson 1968, Evans et al. 2011, Kobmoo et al. 2012, 2015, Araújo et al. 2018, Kuephadungphan et al. 2020, Kuephadungphan et al. 2022).

Akanthomyces sensu Kepler et al. (2017) was known to occur on various insects such as adult moths, ladybugs, lepidopteran larvae, scale insects, whiteflies, spiders, and weevils (Mains 1950, Samson & Evans 1974, Samson & Brady 1982, Hsieh et al. 1997, Chen et al. 2018, Aini et al. 2020). However, our analyses indicate that Akanthomyces sensu stricto is limited to moths, presenting an important generic discriminating criterion. In clade C (Fig. 1A), we propose four new species of Akanthomyces on adult moths: Ak. buriramensis, Ak. fusiformis, Ak. niveus, and Ak. phariformis. Additionally, we present and illustrate a sexual morph linked to Ak. laosensis and Ak. pseudonoctuidarum. Two types of synnemata for Ak. noctuidarum and Ak. pseudonoctuidarum are also documented.

It is crucial to note that Ak. tuberculatus OSC 111002 (Kepler et al. 2012) and Ak. tuberculatus HUA 186131 (Kepler et al. 2017) are distinct entities requiring further study, as these two strains lack morphological data for comparison with the type species. The absence of DNA sequence data from the type specimen poses challenges in determining whether query sequences from new samples represent new or previously described taxa. While Aini et al. (2020) demonstrated moth family specificity in Akanthomyces, our study did not reveal such host-specificity. Most reported species of Akanthomyces sensu stricto are from the Old World (OW), with limited data from the New World (NW). Increased sampling from the NW is likely to unveil additional species. Previous studies in entomopathogenic fungi have highlighted differences in the distribution of morphologically similar but distinct species of the same genus between the OW and NW (Araújo et al. 2018).

The Arachnidicola clade (clade A) predominantly comprises Akanthomyces species that mostly exhibit an isaria-like asexual morph, except for Ak. thailandicus and Ak. tiankengensis, which produce a lecanicillium-like asexual morph. Given that species within this clade exclusively infect spiders, we propose the name Arachnidicola, signifying “dwelling in spiders” rather than Akanthomyces. Additionally, we present a key to the species of Arachnidicola. The new taxonomic combinations include Arachnidicola araneicola (= Akanthomyces araneicola), Ar. araneogenum (= Ak. araneogenum), Ar. bashanensis (= Ak. bashanensis), Ar. beibeiensis (= Ak. beibeiensis), Ar. kanyawimae (= Ak. kanyawimae), Ar. kunmingensis (= Ak. kunmingensis), Ar. subaraneicola (= Ak. subaraneicola), Ar. sulphurea (=Ak. sulphureus), Ar. tiankengensis (= Ak. tiankengensis), Ar. thailandica (= Ak. thailandicus), and Ar. waltergamsii (=Ak. waltergamsii). Within Cordycipitaceae, more than 80 fungal species have been reported as spider pathogens (Shrestha et al. 2019). The genera Gibellula, Hevansia, Jenniferia, and Polystromomyces have been identified as obligate parasites of spiders (Kuephadungphan et al. 2020, 2022, Mongkolsamrit et al. 2022). Different Gibellula species have shown specialization to distinct groups of spiders; for instance, G. cebrennini, G. pigmentosinum, and G. scorpioides were found on Cebrenninus cf. magnus, Storenomorpha sp. (Zodariidae), and Portia sp. (Salticidae), respectively (Kuephadungphan et al. 2022). Hevansia minuta was reported by Mongkolsamrit et al. (2022) on Meotipa sp. (Theridiidae), Jenniferia griseocinerea on Diaea cf. dorsata and Diaea sp. (Thomisidae), J. thomisidarum on Diaea cf. dorsata (Thomisidae), and Polystromomyces araneae on an unidentified spider egg sac. A similar level of specialization may also apply to Arachnidicola, necessitating further investigation into the identity of the hosts. It is noteworthy that all Arachnidicola species produce white conidia on mononematous conidiophores.

The Lecanicillium clade (clade B) encompasses Akanthomyces species that primarily produce lecanicillium-like asexual morphs, infecting a diverse host range across major arthropod groups (Insecta: Coleoptera, Diptera, Hemiptera, Lepidoptera, Aranaea), mushrooms, and soil. This clade includes the type species of the genus Lecanicillium (L. lecanii) and core species such as L. attenuatum, L. muscarium, L. pissodis, L. sabanensis, and L. uredinophilum. We propose the resurrection of the genus Lecanicillium to accommodate these species (Akanthomyces lecanii (= Lecanicillium lecanii), Ak. attenuatus (= L. attenuatum), Ak. dipterigenus (= L. longisporum), Ak. muscarius (= L. muscarium), Ak. neoaraneogenus (= L. araneogenum), Ak. pissodis (= L. pissodis), Ak. uredinophilus (= L. uredinophilum), and Ak. sabanense (= L. sabanense)). This resurrection also implies new taxonomic combinations, including L. araneosus (= Ak. araneosus), L. neocoleopterorum (= Ak. neocoleopterorum), and L. lepidopterorum (= Ak. lepidopterorum). As originally described by Gams & Zare (2001), the characteristics of Lecanicillium include yellowish to white colonies, usually fast-growing, with one or two whorls of phialides occurring on erect, short conidiophores. The phialides are discrete, aculeate, tapering to a narrow tip, verticillate, or solitary, and in some taxa, they form a short, swollen base that bears conidia. The conidia, short to long ellipsoidal to falcate with pointed ends, adhere together into slimy heads or fascicles, sometimes forming chains (Gams & Zare 2001). The morphology of the asexual morph of all the species in Clade B aligns with the characteristics ascribed to Lecanicillium. We present a key to Lecanicillium species based on host identity and the typical characters of the asexual morph. The panglobal distribution of Lecanicillium is noteworthy, found on a diverse arthropod host range (Kope et al. 2005). Lecanicillium is also of interest as a biological control agent for insect pests; for example, L. pissodis, isolated from the white pine weevil (Pissodes strobi) in British Columbia, exhibits pathogenicity to weevil populations and has potential for controlling the white pine weevil (Kope et al. 2005). Additionally, L. uredinophilum (= Ak. uredinophilus), isolated from hemipteran insects in America, serves as a biological control agent against agricultural pests (Manfrino et al. 2022) and rust fungi (Uredinales) (Park et al. 2015).

Kanoksria (clade D) stands as a monotypic genus, comprising solely Kanoksria zaquensis, isolated from the stroma and sclerotium of Ophiocordyceps sinensis. This fungus exhibits single, lanceolate, or narrowly lageniform phialides directly from aerial hypha. It marks the tenth genus containing mycoparasitic species within Cordycipitaceae, alongside Corniculantispora, Flavocillium, Lecanicillium, Liangia, Niveomyces, Pseudogibellula, Pseudoniveomyces, Simplicillium, and Zarea.

Basal to Neohyperdermium and Pleurodesmospora lies the newly identified genus Corniculantispora, encompassing three distinct species: Corn. aranearum, Corn. dimorphum and Corn. psalliotae. The designated type species is Corn. psalliotae (formerly L. psalliotae), primarily associated with Agaricus bisporus and A. bitorquis but exhibiting versatility across various hosts, including Arthropoda (cicadellid, mites, nymphs of Ixodes), Nematoda (eggs of Meloidogyne), Fungi (Rhopalomyces), Plantae (Capsicum annuum), dead insects, forest soil, as well as casing soil in mushroom cultures (Dayal & Barron 1970, Balazy et al. 1987, Viaene & Abawi 1998, Hidalgo et al. 2000, Zare & Gams 2001). Corniculantispora showcases two distinct conidial types: (a) crescent-shaped, 1-celled macroconidia, typically with sharply pointed ends, and (b) oval or ellipsoidal microconidia, with Corn. aranearum being an exception, producing obovoid conidia. The presence of oosporein, a toxic fungal secondary metabolite, can be seen as a pigment diffusing in the agar medium. This phenomenon, observed in the cultures of Corn. aranearum and Corn. psalliotae, shows a distinctive pinkish-red or purplish-red appearance of the agar medium. Octahedral crystals are consistently observed in all species. Historically, Corniculantispora dimorphum (= L. dimorphum) posed challenges in morphological differentiation from Corn. psalliotae (= L. psalliotae) and Corn. aranearum (= L. aphanocladii). However, phylogenetic analysis based on ITS rDNA revealed that they are indeed distinct species (Zare & Gams 2001). Notably, the production of octahedral crystals, reminiscent of Corniculantispora, is also observed in certain Lecanicillium species, including L. dipterigenus, L. fusisorum, L. lecanii, L. muscarium, and L. nodulosum.

Zouia represents another monotypic genus positioned in close phylogenetic proximity to Parengyodontium and Blackwellomyces. The designated species, Zouia cauligalbarum, was originally isolated from a stemborer (Lepidoptera). Morphologically, Zo. cauligalbarum shares similarities with Corniculantispora aranearum, L. attenuatum, and L. nodulosum in terms of short conidiogenous cells. Nevertheless, a notable distinction arises in Zo. cauligalbarum through its spore production pattern, reminiscent of Mariannaea, notably lacking the octahedral crystals observed in Corn. aranearum and L. nodulosum.

Predominantly found occurring on mushrooms, the newly classified genus Zarea also exhibits distribution in soil, forest litter, insects, and spiders (Zare & Gams 2008). The type species, Zarea (Acrostalagmus) fungicola, was initially discovered on an unidentified agaric in Germany, establishing the foundation for this genus. Currently, two recognized species inhabit this genus: Za. flavidum and Za. fungicola, with Za. flavidum distinguishing itself through repeated branching of its conidiophores, the presence of octahedral crystals, slower growth, and a lower optimum temperature for development. In contrast, Za. fungicola features a simple conidiophore axis, with octahedral crystals either present or absent. The taxonomic distinction initially proposed by Zare & Gams (2008) introduced two varieties, var. fungicola and var. aleophilum, primarily based on optimal temperature preferences in culture. Although, molecular data, particularly from ITS analyses, clearly indicate a clustering between these varieties, devoid of any morphological disparities, our comprehensive study, employing multiple loci, reinforces the segregation of Za. fungicola and Za. aleophilum varieties, allowing species nomination valid in the current taxonomic framework.

Lecanicillium magnisporum (= Gam. magnispora) was transferred by Crous et al. (2023) to Gamszarella together with Lecanicillium antillanum and Gamszarella buffelskloofina based on molecular phylogenetic analysis. These three species group together in a well-supported clade that also included Neotorrubiella, with Gamszarella magnispora taking a basal position (Fig 1B). Gamszarella antillana, Gam. buffelskloofina and Gam. magnispora produce either single or whorls of 2–4 conidiogenous cells, with phialides in Gam. buffelskloofina terminating in denticles. We propose the new genus Corpulentispora to be erected to transfer the basally positioned Gam. magnispora as Corpulentispora magnispora. The morphological characteristics of Cor. magnispora and Gam. antillana are similar; both species produce two types of fusoid conidia, macroconidia and microconidia, while Gam. buffelskloofina produces only one type of subcylindrical conidia. Nevertheless, Cor. magnispora has larger conidia than Gamszarella. Corpulentispora, positioned in Clade 7, stands as a monotypic genus exclusively characterized by the asexual reproductive capabilities of Corpulentispora magnispora, originating from soil (Zhang et al. 2020). The two other species could be saprobic on dead insects and agarics (Crous et al. 2023)

The taxonomic history within Cordycipitaceae is intricate (Kepler et al. 2017). This study significantly contributes to our comprehension of Akanthomyces and Lecanicillium, shedding light on their relationships within Cordycipitaceae. The members of Akanthomyces and Lecanicillium were identified and reestablished as separate genera through this research. However, certain previously recognized Akanthomyces and Lecanicillium species couldn’t be incorporated into the new classification due to the unavailability of samples for phylogenetic analyses and the absence of DNA sequence data from the type specimens. Therefore, further molecular data are necessary for their classification within the present system.

ACKNOWLEDGEMENTS

We would like to thank the staff of Plant Microbe Interaction Research Team (APMT) including B. Thongnuch, K. Tasanathai and W. Noisripoom for samples collection. We are grateful to S. Kuephadungphan for some beautiful photos of the specimens as well as P. Wongpitakchai from the Department of Microbiology, Kasetsart University for the line drawing of some new genera in Fig. 9. This study was supported by a BIOTEC Mid-Career Research Fellowship, Platform Technology Management Section, National Center for Genetic Engineering and Biotechnology (BIOTEC), Grant Agreement No. P-1950231, the National Science, Research and Innovation Fund, Thailand Science Research and Innovation (TSRI) Grant Agreement No. P-2351531 and the European Commission MSCA Research and Innovation Staff Exchange (RISE) funding on “MYCOBIOMICS”, Grant Agreement No. 101008129.

Conflict of interest: The authors declare that there is no conflict of interest.

Supplementary Material: http://fuse-journal.org/

Fig. S1A.

Phylogenetic tree based on the LSU sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1A.psd^{(3.5MB, psd)}

Fig. S1B.

Phylogenetic tree based on the TEF1 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1B.psd^{(3MB, psd)}

Fig. S1C.

Phylogenetic tree based on the RPB1 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1C.psd^{(3MB, psd)}

Fig. S1D.

Phylogenetic tree based on the RPB2 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1D.psd^{(3MB, psd)}

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Associated Data

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

Supplementary Materials

Fig. S1A.

Phylogenetic tree based on the LSU sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1A.psd^{(3.5MB, psd)}

Fig. S1B.

Phylogenetic tree based on the TEF1 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1B.psd^{(3MB, psd)}

Fig. S1C.

Phylogenetic tree based on the RPB1 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1C.psd^{(3MB, psd)}

Fig. S1D.

Phylogenetic tree based on the RPB2 sequences. Statistical support value (≥ 50 %) is shown at nodes for maximum likelihood.

fuse-2024-14-271-FigS1D.psd^{(3MB, psd)}

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PERMALINK

A phylogenetic assessment of Akanthomyces sensu lato in Cordycipitaceae (Hypocreales, Sordariomycetes): introduction of new genera, and the resurrection of Lecanicillium

A Khonsanit

D Thanakitpipattana

S Mongkolsamrit

N Kobmoo

N Phosrithong

RA Samson

PW Crous

JJ Luangsa-ard

Abstract

INTRODUCTION

MATERIALS AND METHODS

Isolates

Molecular phylogenetic analyses

Morphology

RESULTS

Molecular phylogenetic analyses

Table 1.

Fig. 1A, B.

Taxonomy

Fig. 2.

Fig. 3.

Table 2.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Table 3.

Fig. 8.

Residual species of Akanthomyces

Novel genera and new combinations

Fig. 9.

DISCUSSION

ACKNOWLEDGEMENTS

Supplementary Material: http://fuse-journal.org/

REFERENCES

Associated Data

Supplementary Materials

ACTIONS

PERMALINK

RESOURCES

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