Phallus chiangmaiensis sp. nov. and a Record of P. merulinus in Thailand

Figures & data

Table 1. Taxa used in the phylogenetic analyses and the new taxa in bold.

Species	Strains	GenBank accession numbers
		ITS	LSU	atp6
Mutinus albotruncatus	UFRN Fungos 2025	MF447826	KC128650	KT183490
Phallus atrovolvatus	INPA 240016	MG678531	MG678470	MG678559
Phallus aureolatus	ICN 176962^T	MF372135	MF372127	–
Phallus calongei	AH 31862	–	FJ785522	–
Phallus campanulatus	ICN 176970	MF372138	MF372130	–
Phallus campanulatus	ICN 176971	MF372139	MF372131	–
Phallus chiangmaiensis	BCC 92054^T	MT452882	MT447464	MT454265
Phallus chiangmaiensis	BCC 92055	MT452883	MT447465	MT454263
Phallus cinnabarinus	INPA 255835	KJ764821	MG678471	MG678561
Phallus cinnabarinus	INPA 255836	MG678533	MG678472	MG678562
Phallus coronatus	LE 295238	MG678522	MG678466	MG678554
Phallus costatus	MB 02040	–	DQ218513	–
Phallus denigricans	INPA 272383^T	MG678486	MG678455	MG678541
Phallus denigricans	UFRN Fungos 2805	MG678485	MG678454	–
Phallus dongsun	GDGM 29086	MN307394	MN264676	–
Phallus dongsun	GDGM 75402^T	MN307397	MN264679	–
Phallus echinovolvatus	GDGM 79020	MN523216	–	–
Phallus echinovolvatus	TNS F 34480	MF372137	MF372129	–
Phallus flavocostatus	RE 2004	MG678524	MG678467	MG678556
Phallus fuscoechinovolvatus	GDGM 48589^T	MF039581	MF039585	–
Phallus fuscoechinovolvatus	GDGM 48663	MF039582	MF039586	–
Phallus hadriani	AH 39161	KF481956	–	–
Phallus haitangensis	HKAS 88197^T	KU705383	–	–
Phallus impudicus	CBS 294.53	–	MH868748	–
Phallus impudicus	GDGM 77656	MN307393	MN264675	–
Phallus indusiatus	INPA 264929	MG678498	–	MG678548
Phallus indusiatus	INPA 264931	MG678500	MG678463	MG678550
Phallus lutescens	GDGM 49991	MN131081	MN131077	–
Phallus lutescens	GDGM 72218^T	MN131079	MN131075	–
Phallus luteus	146778	HQ414538	–	–
Phallus luteus	TNS Kasuya B 218	KP222543	KP222545	–
Phallus mengsongensis	HKAS 78342	KF052627	–	–
Phallus mengsongensis	HKAS 78343^T	KF052624	–	–
Phallus merulinus	INPA 240010	MG678530	MG678469	MG678558
Phallus merulinus	BCC 92056	MT466468	MT447463	MT454264
Phallus merulinus	BCC 92057	MT466469	MT447462	MT454266
Phallus multicolor	MEL 2382891	KP012762	–	–
Phallus purpurascens	SINOP 26	MG678488	MG678457	MG678543
Phallus purpurascens	UFRN Fungos 2808 ^T	MG678487	MG678456	MG678542
Phallus ravenelii	CUW s.n.	–	–	DQ218799
Phallus rubrovolvatus	YZS 040	KF939503	–	–
Phallus rugulosus	TNS F 46049	MF372142	MF372134	–
Phallus serratus	HKAS 78340^T	KF052622	–	–
Phallus squamulosus	UFRN Fungos 2806^T	MG678497	–	MG678547
Phallus ultraduplicatus	HMAS 253050^T	KJ591584	KJ591586	–
Phallus ultraduplicatus	HMAS 253051	KJ591585	KJ591587	–

The “T” represents ex-holotype strains. Mutinus albotruncatus was used as an outgroup.

Figure 1. Phylogenetic relationships of Phallus spp. inferred from ITS sequences. Numbers at the significant nodes represent ML bootstrap values/MP/Bayesian posterior probabilities, multiplied by 100; bold lines in the tree represent 100% bootstrap (BSMP, BSML) and 1.00 posterior probability (BPP).

Figure 2. Phylogenetic relationships of Phallus spp. from a combined ITS, LSU, and atp6 analyses. Numbers at the significant nodes represent ML bootstrap values/MP/Bayesian posterior probabilities, multiplied by 100; bold lines in the tree represent 100% bootstrap (BSMP, BSML) and 1.00 posterior probability (BPP).

Figure 3. Phallus chiangmaiensis (BBH 47825, holotype). (a) Mature basidiomata. (b) Reticulate cap. (c) Indusium. (d) Immature basidiomata (egg). (e) Pseudostipe and section of immature basidiomata. Scale bars: a = 50 mm, b–e = 10 mm.

Figure 4. Microscopic features of Phallus chiangmaiensis. (a,b) Cap cells and hyphae. (c) Cells of indusium. (d) Cells of pseudostipe. (e) Crystals in volva hyphae (arrowed). (f,g) Volva hyphae with clamp connections (arrowed). (h,i) Rhizomorph hyphae with clamp connections (arrowed). (j) Basidia with sterigmata and basidiospores. (k) Basidiospores. (l) Colony on PDA (surface and reverse plate). Scale bars: a, c–e = 20 µm, b, f–j = 10 µm, g, k = 5 µm, l = 10 mm.

Figure 5. Line drawing of Phallus chiangmaiensis. (a) Fruiting body. (b) Cells of indusium. (c,d) Cells of pseudostipe. (e) Volva hyphae. (f) Rhizomorph hyphae. (g) Basidia. (h) Basidiospores. Scale bars: a = 50 mm, b–d = 50 μm, e = 10 μm, f = 20 μm, g–h = 5 μm.

Table 2. Synopsis of macro- and micro- characteristics of Phallus chiangmaiensis, P. echinovolvatus, P. fuscoechinovolvatus, P. lutescens, P. luteus, and P. multicolor.

Species name	P. chiangmaiensis	P. echinovolvatus	P. fuscoechinovolvatus	P. lutescens	P. luteus	P. multicolor
Cap shape	Campanulate	Campanulate	Ovoid to slightly conical or campanulate	Subovoid to campanulate	Conical to slightly campanulate	Campanulate
Cap size (H × W, mm)	40 – 50 × 25 – 45	25 – 30 × 25 – 30	22 – 40 × 10 – 22	15 – 20 × 16 – 23	25 – 40 × 26 – 38	25 – 30 × 20
Cap color	Light yellow	Nearly white to yellow	Yellowish white	Yellowish white to pale yellow	Pale yellow to yellowish orange	Lemon yellow
Cap surface characters	Strongly reticulated	Reticulated	Strongly rugose	Reticulated with irregular ridges	Strongly reticulated	Reticulated
Gleba color	moderate olive brown	Olive to dark brown	Olivaceous brown	Olive brown	Olivaceous brown to greenish black	Olive brown
Indusium length (mm)	130 – 160 or ¾ the size of the pseudostipe	70 – 100	¾ the size of the pseudostipe	Expanded to ⅔–⅚ portion of pseudostipe	60 –160	78
Indusium color	White	White	White	White, yellowish-white to pale yellow	Yellow to yellowish orange	Lemon yellow to yellowish orange
Indusium characters (meshes)	Hexagonal or polygonal	Polygonal	Hexagonal or polygonal	Polygonal	Polygonal to round	N/A
Pseudostipe shape	Cylindrical, tapering toward the apex	Cylindrical to fusiform	Cylindrical or fusiform	Cylindrical usually tapered upwards and enlarged downwards	Cylindrical, subfusoid or tapering toward the base	Narrow upward
Pseudostipe size (mm)	158 – 165 × 20 – 25	90 – 150 × 20 – 30	80 – 130 × 15 – 20	70 – 90 × 9 – 23	70 – 220 × 15 – 25	60 – 85 × 25 – 30
Pseudostipe color	White	Nearly white	Snow white to milky white	Snow white to very weak cream white	N/A	Yellowish white
Pseudostipe characters	Spongy, hollow	Hollow	Hollow	Spongy, hollow	Spongy, hollow	Hollow
Volva characters	Non-echinulate	Echinulate	Echinulate	Smooth or lightly rugose	Non-echinulate	Non-echinulate
Basidia	7.0–15.0 × 1.5–3.0 μm, elongated, cylindrical, a bit broader at the center, and hyaline. Sterigmata 4–8 in number	6.0–8.0 × 2.5–3.5 μm, cylindrical or clavate, Sterigmata 4–6 in number	N/A	N/A	N/A	N/A
Spore shape	Cylindrical to broadly ellipsoid	Oval to ellipsoid	Cylindrical to broadly ellipsoid	Cylindrical to long ellipsoid	Broadly ellipsoid to cylindrical	Long-elliptical to nearly cylindrical
Spore size (µm)	3.0 – 4.0 × 1.5 – 2.0	3.0 – 4.0 × 1.3 – 2.0	2.5 – 4.0 × 1.0 – 2.0	3.0 – 4.3 × 1.1 – 1.8	3.0 – 4.0 × 1.5 – 2.0	3.94 – 4.33 × 1.77 – 1.97
Spore color	Greenish white	Light brownish green to olive	Hyaline and very light olivaceous	Hyaline and light olivaceous	Hyaline	Hyaline

Figure 6. Phallus merulinus (BBH 47826). (a,b) Basidiomata. (c,d) Immature basidiomata (eggs) with rhizomorph (arrowed). (e) Indusium and pseudostipe. (f) Volva. Scale bars: a–b = 50 mm, c–f = 20 mm.

Figure 7. Microscopic features of Phallus merulinus. (a–e) Cap cells and hyphae. (f) Cells of indusium. (g) Cells of pseudostipe. (h,i) Volva hyphae. (j,k) Rhizomorph hyphae. (l) Basidiospores. (m) Colony on PDA (surface and reverse plate). Scale bars: a–b, f–g = 20 µm, c–e, h–l = 5 µm, m = 20 mm.

Figure 8. Line drawing of Phallus merulinus. (a) Fruiting body. (b,c) Cap cells and hyphae. (d) Cells of indusium. (e,f). Cells of pseudostipe. (g) Volva hyphae. (h) Rhizomorph hyphae. (i) Basidiospores. Scale bars: a = 20 mm, b, g = 10 μm, c, h, i = 5 μm, d = 20 μm, e–f = 50 μm.