Scleroderma geaster Fr., 1829

Ortiz-Rivero, Javier, Watling, Roy, Guzmán-Dávalos, Laura & Martín, María P., 2021, The many-rooted earthball-Scleroderma geaster and S. polyrhizum revisited, with the description of a new species, Phytotaxa 510 (1), pp. 1-17 : 7-13

Scleroderma geaster Fr. , Systema Mycologicum 3: 46 (1829) [MB#183185] (Figs. 2A–C, 3A)

Lectotype (designated here): Micheli , P. A. (1729) Nova plantarum genera. Gasteromycetes: 213–222. 66: 1 p. 210. Fries SM III refers to plate 99:1 (not 66) and page 210 in Micheli; obviously a printing error by Fries, since the text in Micheli is on page 219. Persoon 1801:156 has it correct.

FIGURE. Basidiomes of Scleroderma species. A –C. Scleroderma geaster (A. MA–Fungi 17587; B. MA–Fungi 28137; C. MA– Fungi 21907). D–F. Scleroderma guzmanii (D. OSC41254, holotype; E. J.A. García Valle 24, IBUG; F. Y. Alquiciras 76, IBUG). G–I. Scleroderma polyrhizum (G. MA–Fungi 22183; H. MA–Fungi 31687; I. MA–Fungi 39352). J–K. Scleroderma australe (J. MEL2061745; K. MEL269386). L. Scleroderma texense (VPIF–0004156). Scale bars = 1 cm.

Epitype (designated here):— GREECE. Crete, 2002, Lasithi plateau, terrestrial, on sandy soil, 15 October 2002, Walkinshaw in Roy Watling collection E 1574 *36 ( E!) [ MBT#391528 ; GenBank accession number ITS MT 270644 ]. Description: —Basidiomes epigeous, globose to subglobose, pyriform, very hard in dry state, (3–)4–7(–13) cm diam. Peridium tough and leathery, 2–4(–6) mm thick, smooth or covered with mycelium to irregularly scaly, scales not well-developed, yellow, bay brown or “Chamois” (colour 250), sessile, stipe absent or with very short pseudostipe at the base, with numerous, short, mycelial cords. Gleba compact and whitish in immature state, pulverulent and burnt umber (colour 701–702) in adult stage. Dehiscence stelliform .

Basidiospores globose to subglobose,(5.8–)7.9–9.2(–10.6)µm diam, dark brown in KOH, including ornamentation, subreticulate to reticulate under LM and SEM (Fig. 3A), with spines and reticulum to 0.7 µm high.

Ecology and distribution:—This species has European distribution, very common in the Mediterranean region in countries such as Greece, Portugal, Spain with a Mediterranean climate or on volcanic islands such as Madeira. It fruits mainly during the autumn months. Silvicolus, in sandy soil under Quercus sp. and Pinus sp. or with Cistus sp. and Juniperus sp. ; also frequent on roadsides, parks or meadows.

FIGURE. Basidiospores of Scleroderma species under SEM.A. Scleroderma geaster (E 157436). B. Scleroderma guzmanii (OSC41254). C. Scleroderma polyrhizum (MA–Fungi 30844). D. Scleroderma australe (MEL2061737). E. Scleroderma texense (VPIF–0004156). F. Scleroderma yunnanense (PERTH8631875). Scale bars = 5 µm.

Additional specimens examined:— PORTUGAL. Estremadura, Apostiça, in Pinus pinaster forest, 11 November 1991, F.D. Calonge (MA–Fungi 31362, GenBank accession number ITS MT270635 ); Madeira, Queimadas, in mixed forest Pinus / Quercus , 11 November 1991, F.D. Calonge (MA–Fungi 55033, GenBank accession number ITS MT270640 ); Madeira, Sao Vicente, 31 February 1988, M. Gutiérrez (MA–Fungi 57138, GenBank accession number ITS MT270641 ); Minho, P. Nac. Peneda-Gerês, 11 August 1987, A. Guerra (MA–Fungi 22186, GenBank accession number ITS MT270624 ); SPAIN, Ávila, La Adrada, 26 October 1986, J. Barroso (MA–Fungi 16496, GenBank accession number ITS MT270617 ); Barcelona, Montseny, oak wood, 14 March 1992, S. Estanyol Duocastella (MA– Fungi 32448, GenBank accession number ITS MT270637 ); Cáceres, Abadía, 26 November 1987, F.D. Calonge (MA– Fungi 21908, GenBank accession number ITS MT270623 ); Cáceres, Jaraiz de la Vera, 26 November 1982, F.D. Calonge (MA–Fungi 21875, GenBank accession number ITS MT270621 ); Cáceres, Parque Nacional de Monfragüe, 7 December 1991, F.D. Calonge (MA–Fungi 30846, GenBank accession number ITS MT270631 ); Ciudad Real, Viso del Marqués, 9 December 1989, F.D. Calonge (MA–Fungi 31688, GenBank accession number ITS MT270636 ); Córdoba, El Cerillo, 14 November 1992, J. Gómez & A. Castro (MA–Fungi 30592, GenBank accession number ITS MT270629 ); Córdoba, Ctra de Villaviciosa, 14 March 1992, A. Castro (MA–Fungi 30895, GenBank accession number ITS MT270633 ); Córdoba, Cuesta del Cambrón, 11 December 1992, J. Gómez & A. Castro (MA–Fungi 30591, GenBank accession number ITS MT270628 ); Córdoba, Villaviciosa, 14 November 1992, J. Gómez & A. Castro (MA–Fungi 30590, GenBank accession number ITS MT270627 ); Huelva, Doñana, Sabinar del Marqués, Juniperus forest-sect. Sabina , 24 February 1977, F.D. Calonge et al. (MA–Fungi 181, GenBank accession number ITS MT270612 ); Huelva, Hinojos, Pinus forest, 20 November 1986, M.A. Cortés, L. Cox, J. Jauregui & F.D. Calonge (MA–Fungi 21873, GenBank accession number ITS MT270620 ); Huelva, Matalascañas, 17 February 1980, B. Valdés (MA–Fungi 345, GenBank accession number ITS MT270613 ); Huelva, Mazagón, Lagunas de Moguer, 18 November 1976, Costa et al. (MA–Fungi 5, GenBank accession number ITS MT270611 ); Lugo, Pantón, Os Peares, Pinus forest, 23 December 1993, R. González & D. Gómez (MA–Fungi 34025, GenBank accession number ITS MT270638 ); Madrid, Canencia, 15 October 1992, F.D. Calonge (MA–Fungi 30847, GenBank accession number ITS MT270632 ); Orense, Barco de Valdeorras, Cistus laurifolius , 14 November 1992, J. Gómez & A. Castro (MA–Fungi 35266, GenBank accession number ITS MT270639 ); Orense, La Rúa, 12 November 1986, F.D. Calonge (MA–Fungi 17587, GenBank accession number ITS MT270619 ); Orense, San Cibrao das las Barbantes, 26 November 1982, F.D. Calonge (MA–Fungi 4804, GenBank accession number ITS MT270615 ); Pontevedra, Tuy, 21 November 1991, V. Ruiz (MA–Fungi 30845, GenBank accession number ITS MT270630 ); Salamanca, Ciudad Rodrigo, 25 October 1987, E. Sánchez-Abarca (MA–Fungi 21907, GenBank accession number ITS MT270622 ); Salamanca, Ciudad Rodrigo, 21 November 1990, E. Sánchez-Abarca (MA–Fungi 28202, GenBank accession number ITS MT270626 ); Salamanca, Herguijuela de la Sierra, Pinus pinaster forest, 27 November 2002, S. Pérez Gorjón & P. García Jiménez (MA–Fungi 84857, GenBank accession number ITS MT2706443); Salamanca, Sepulcro Hilario, Quercus ilex forest, 20 October 1986, F.D. Calonge (MA–Fungi 16493, GenBank accession number ITS MT270616 ); Segovia, Prádena, Prunus sp. , 2 June 1991, J.M. Santos (MA–Fungi 28137, GenBank accession number ITS MT270625 ); Sevilla, Aznalcollar, Pinus forest, 6 May 1972, S. Silvestre (MA–Fungi 1278, GenBank accession number ITS MT270614 ); Teruel, Cendrillas, 23 October 1990, F.D. Calonge (MA–Fungi 31358, GenBank accession number ITS MT270634 ); Teruel, Orihuela del Tremedal, Cistus laurifolius , 15 October 1996, E. Suarez (MA–Fungi 37552, GenBank accession number ITS MT270618 ); Toledo, San Pablo de los Montes, Quercus pirenaica , 19 November 2004, F.D. Calonge, F. Prieto, A. González et al. (MA–Fungi 68588, GenBank accession number ITS MT270642 ).

Remarks:— Scleroderma polyrhizum and S. geaster as used by a range of authorities are different species and are not synonymous as previously thought; although both species have a European distribution. Scleroderma geaster has basidiospore size slightly larger than S. polyrhizum , and the basidiomes have a stipe or pseudostipe not present in S. polyrhizum .

Scleroderma guzmanii Ortiz-Rivero, Watling, Guzmán-Dávalos & M.P. Martín , sp. nov. [MB#835840] (Figs. 2D–F, 3B).

Etymology:—In reference to Gastón Guzmán, who wrote a monograph on the genus Scleroderma .

Holotype:— USA. Arkansas, Montgomery , 3 November 1982, J. Trappe (OSC-41254!, GenBank accession number ITS MT 270648 ).

Diagnosis:—This species is very similar to S. polyrhizum in basidiospore size and ornamentation, but they differ in their ITS nrDNA sequences.

Description:— Basidiomes epigeous, globose to subglobose, <11 cm diam. Peridium tough and leathery, <3–6 mm thick, often forming several lobes which become recurved, smooth or covered with mycelium to irregularly scaly, scales not well-developed, yellow, bay brown or “Chamois” (colour 250). Base with numerous, short mycelial cords but without stipe. Gleba compact when young and pulverulent in adult stage, tobacco brown (colour 692) to cocoa (colour 697). Dehiscence stellate.

Basidiospores dark brown in KOH, globose to subglobose,(5.7–)7.2–8.2(–10.2)µm diam,including ornamentation, subreticulate to reticulate under LM and SEM (Fig. 3B), with spines and a reticulum to 0.5 µm high.

Ecology and distribution:—This species has a distribution in the New World from the United States to Mexico. Basidiomes develop mainly during the autumn months, in sandy soil in natural forest or those replanted with Pinus sp. and Eucalyptus sp.

Additional specimens examined:— MEXICO. Jalisco, Guadalajara, Bosque Los Colomos, Pinus / Eucalyptus forest, 1 November 2015, J.A. García Valle 24 (IBUG; GenBank accession number ITS MT270645 ); Jalisco, Mazamitla, El Terrero, Pinus forest, 9 September 1998, Y. Alquiciras 76 (IBUG; GenBank accession number ITS MT270647 ); Jalisco, Zapopan, Bosque La Primavera, Pinus forest, 10 February 1997, J.A. Pérez de la Rosa s.n. (IBUG; GenBank accession number ITS MT270646 ). USA, Massachusetts, Barnstable, 30 November 1908, H.H. Barlett 1600 (MICH12441) [under S. polyrhizum neotype designed by G. Guzmán].

Remarks:—Morphologically, S. guzmanii has an ornamentation and basidiospore size very similar to S. polyrhizum . This species probably presents a wider distribution across the American continent. Molecular and geographical information is key to delimit and to describe the specimens studied, both indicating as belonging to a new species. The specimens identified as S. polyrhizum by Guzmán-Dávalos & Guzmán (1985) could correspond to S. guzmanii .

Scleroderma polyrhizum (J.F. Gmel.) Pers. , Synopsis methodica fungorum: 156 (1801), MB#414483 (Figs. 2G–I, 3C)

≡ Lycoperdon polyrhizum J.F. Gmel. [as ‘ polyrhizon ’], Systema Naturae 2(2): 1464 (1792), MB#508326

≡ Sclerangium polyrhizum (J.F. Gmel.) Léveillé, Annales des Sciences Naturelles Botanique 9: 130 (1848), MB#206625

Lectotype (designated here): Léveillé, J.H. (1848) Annales des Sciences Naturelles Botanique sér. 3, 9, 130 pl. 7, 1848. This measure seems sensible since Persoon in his description of S. polyrhizum refers to Micheli pl 99:1, the same illustration chosen to lectotypify S. geaster .

Epitype (designated here):— SPAIN. Zaragoza, Litago, mixed forest Pinus / Quercus , 8 October 1997, F.D. Calonge (MA–Fungi 39352!) [MBT#391529, GenBank accession number ITS MT270662 ]

Non-neotype of S. polyrhizum (designated by G. Guzmán):— USA. Massachusetts, Barnstable , 30 November 1908, H. H . Barlett 1600 ( MICH12441 !) .

Description: —Basidiomes (3.5–) 7–10 cm diam, subglobose, pyriform or tuberiform, contracting into a small stemlike base. Peridium tough and leathery, <2–3(–5.5) mm thick, often forming several lobes which become recurved, smooth or covered with mycelium to irregularly scaly, scales not well-developed, clear brown (colour 190) to grenadine yellow (colour 199). Base with numerous prominent, short mycelial cords but without distinct pseudostipe. Gleba compact and globose becoming pulverulent with maturity, burnt umber (colour 701–702). Dehiscence stelliform.

Basidiospores dark brown in KOH, globose to subglobose, (5.7–)7.7–8.5(–10.4) µm diam, ornamentation subreticulate to reticulate under LM and SEM (Fig. 3C), with spines and reticulum <0.6 µm high.

Ecology and distribution:—This species has a European distribution, common in the Mediterranean region, but is also frequently found in areas with a cooler and more humid climate, typical of Atlantic areas. It fruits mainly during the autumn months in sandy soil under Quercus sp. , Pinus sp. or Fagus sp. , also on roadsides, and in parks or meadows.

Additional specimens examined:— SPAIN. Asturias, Oviedo , Pinus forest, 1 September 1988, F . D. Calonge ( MA – Fungi 22183, GenBank accession number ITS MT 270655 ) ; Asturias, Oviedo , 26 September 1991, D. Coello ( MA – Fungi 30844, GenBank accession number ITS MT 270657 ) ; Asturias, Pravia , 17 November 1989, J . Arias ( MA – Fungi 31689, GenBank accession number ITS MT 270659 ) ; Asturias, Valgrande , Fagus forest, 10 November 1989, F . D. Calonge ( MA – Fungi 31687, GenBank accession number ITS MT 270658 ); León, Boñar , Pinus forest, 9 September 1990, J. A . Sánchez ( MA – Fungi 28258, GenBank accession number ITS MT 270656 ) ; Navarra, Elzaburu , Quercus forest, 31 October1987, L . Miguel García Bona ( MA – Fungi 67023, GenBank accession number ITS MT 270663 ); Pontevedra, Bueu , 8 October 1986, C . Lado ( MA – Fungi 16668, GenBank accession number ITS MT 270653 ); Toledo, El Chorro , 11 February 1986, E . Fuertes & N . Marcos ( MA – Fungi 21906, GenBank accession number ITS MT 270654 ); Zamora, Villaldeciervos , 2 May 1988, F . D. Calonge ( MA – Fungi 32214, GenBank accession number ITS MT 270660 ) . USA, Illinois, 10 October 2003, M . Siefken (ILLS-56824, GenBank accession number ITS MT 270661 ) .

Remarks:— Guzmán (1970) neotypified S. polyrhizum with a collection (MICH12441) from Massachusetts ( USA); however, this neotypification requires careful reconsideration because molecular studies indicate marked differences to the European material designated with the same name. The specimen designated as the neotype of S. polyrhizum by Guzmán corresponds to what we are calling S. guzmanii .

For decades Scleroderma geaster and S. polyrhizum have been confused because of their morphological similarity. However, typically the former possesses a very short pseudostipe at the base of the basidiomes with development of short mycelial cords. In contrast S. polyrhizum lacks the pseudostipe although the mycelial cords at the base may fuse together. There is also a slight difference in basidiospore morphology with those of S. geaster generally very slightly larger than S. polyrhizum and slightly more roughened, although both species possess a wide range; see accompanying figure.

TABLE. Specimens of Scleroderma studied in each clade with measurements of basidiospores and ornamentation.

TABLE. (Continued)