Elfin saddle Helvella lacunosa, copyright Len Worthington.

Belongs within: Pezizales.

Saddling the truffles
Published 19 July 2011
The black elfin saddle Helvella lacunosa, photographed by Fred Stevens.

The subject of today’s post is the fungus family Helvellaceae. In the past, the Helvellaceae have been treated as the family including the morels and false morels. False morels and morels are ascomycetes* that produce convoluted fruiting bodies generally supported above ground by a stalk. However, molecular analyses have unanimously indicated the non-monophyly of the morels and false morels relative to the truffles (Percudani et al. 1999), which produce their fruiting bodies underground (while above-ground fungi have their spores generally dispersed by the wind, truffles have spores dispersed by passing through an animal’s digestive system after it eats the truffle). The intermingled relationship between truffles and morels had already been indicated by morphologists based on microscopic features of the spores and asci, and so past members of the Helvellaceae have been dispersed among multiple families. At the same time, genera of truffles have been shown to have a relationship with the Helvellaceae, so of the five genera listed in Helvellaceae in the most recent “Outline of Ascomycota” (Lumbsch & Huhndorf 2007) only two (Helvella and Cidaris) are above-ground fruiters, while the other three (Balsamia, Barssia and Picoa) are truffles. This is ignoring the point that the single known species of Cidaris has not seemingly been identified since its original description (Underwood 1896) and its relationship to Helvella would probably require investigation.

*One of the major groups of fungi, ascomycetes produce spores in an ascus, an elongate structure with spores contained in a row within it.

A stem-less Helvella, H. astieri, photographed by Thomas Læssøe.

Members of the genus Helvella are commonly known as ‘saddle fungi’ or ‘elfin saddles’ due to the appearance of the fruiting bodies in some species. Other species possess a variety of different fruiting morphologies, some cup-like, some irregularly folded and lumpy. Not all Helvella species produce fruiting bodies supported by a stalk: in some, the fruiting body sits on the ground or remains partially submerged (Kimbrough et al. 1996), and it has been suggested that such forms may provide some indication how the truffles evolved from above-ground forms. All Helvella species fruit on soil (i.e. never on rotting wood or other such substrates) and it seems likely that all members of the Helvellaceae form ectomycorrhizal associations with plant roots (Hansen 2006).

How to spot desert truffles… (from here).

The truffle members of the Helvellaceae have a solid gleba (the spore-bearing inner mass) interspersed with veins or pockets of hymenia (the spore-producing tissues) separated by sterile tissue (Kimbrough et al. 1996). Picoa species grow in association with Helianthemum (rockrose) species and are among the ‘desert truffles’ collected in arid parts of the Mediterranean. They are eaten, but are not considered commercially significant due to their small size. Among Helvella species, the white saddle Helvella crispa and black saddle Helvella lacunosa have been described as edible, so long as they are cooked properly.

What the picture above may lead you to… Picoa juniperi, from here.
Systematics of Helvellaceae
<==Helvellaceae [Balsamiaceae]
    |--Balsamia Vittad. 1831 [incl. Pseudobalsamia Fisch. 1907]KC01
    |--Fischerula Mattir. 1928EB03, KC01
    |--Leucangium Quél. 1883KC01
    |--Picoa Vittad. 1831KC01
    |--Cidaris Fr. 1849 nec Leske 1778 (ICZN) nec Swainson 1840 (ICZN)KC01
    |--Underwoodia Peck 1890KC01
    |--Barssia Gilkey 1925 [incl. Phymatomyces Kobayasi 1937]KC01
    |    `--B. oregonensisLK04
    |--Wynnella Boud. 1885 [incl. Midotis Fr. 1828]KC01
    |    `--W. silvicolaPA-W02
    `--Helvella L. 1753 (see below for synonymy) KC01
         |--H. compressaJK06
         |--H. cupularisE99
         |--H. lacunosaSL02
         |--H. macropusE99
         |--H. rivularisE99
         `--H. terrestris (Velen.) Landvik 1999 [=*Pindara terrestris]E99

Helvella L. 1753 [=Costapeda Falck 1923; incl. Acetabula (Fr.) Fuckel 1870, Biverpa (Fr.) Boud. 1907, Boletolichen Juss. 1789, Coelomorum Paulet 1793, Cowlesia Nieuwl. 1916, Cyathipodia Boud. 1907, Fuckelina Kuntze 1898 non Sacc. 1875, Geomorium Speg. 1922, Globopilea Beauseign. 1926, Leptopodia Boud. 1885, Macropodia Fuckel 1869 non Leach 1814 (ICZN), Macroscyphus Gray 1821, Paxina Kuntze 1891, Phaeomacropus Henn. 1899, Phleboscyphus Clem. 1903, Pindara Velen. 1934, Tubipeda Falck 1923]KC01

*Type species of generic name indicated


[E99] Eriksson, O. E. (ed.) 1999. Notes on ascomycete systematics. Nos 2440–2755. Myconet 2: 1–41.

[EB03] Eriksson, O. E., H. O. Barah, R. S. Currah, K. Hansen, C. P. Kurtzman, G. Rambold & T. Laessøe (eds.) 2003. Outline of Ascomycota—2003. Myconet 9: 1–89.

Hansen, K. 2006. Systematics of the Pezizomycetes—the operculate discomycetes. Mycologia 98 (6): 1029–1040.

[JK06] James, T. Y., F. Kauff, C. L. Schoch, P. B. Matheny, V. Hofstetter, C. J. Cox, G. Celio, C. Gueidan, E. Fraker, J. Miadlikowska, H. T. Lumbsch, A. Rauhut, V. Reeb, A. E. Arnold, A. Amtoft, J. E. Stajich, K. Hosaka, G.-H. Sung, D. Johnson, B. O’Rourke, M. Crockett, M. Binder, J. M. Curtis, J. C. Slot, Z. Wang, A. W. Wilson, A. Schüßler, J. E. Longcore, K. O’Donnell, S. Mozley-Standridge, D. Porter, P. M. Letcher, M. J. Powell, J. W. Taylor, M. M. White, G. W. Griffith, D. R. Davies, R. A. Humber, J. B. Morton, J. Sugiyama, A. Y. Rossman, J. D. Rogers, D. H. Pfister, D. Hewitt, K. Hansen, S. Hambleton, R. A. Shoemaker, J. Kohlmeyer, B. Volkmann-Kohlmeyer, R. A. Spotts, M. Serdani, P. W. Crous, K. W. Hughes, K. Matsuura, E. Langer, G. Langer, W. A. Untereiner, R. Lücking, B. Büdel, D. M. Geiser, A. Aptroot, P. Diederich, I. Schmitt, M. Schultz, R. Yahr, D. S. Hibbett, F. Lutzoni, D. J. McLaughlin, J. W. Spatafora & R. Vilgalys. 2006. Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature 443: 818–822.

Kimbrough, J. W., L.-T. Li & C.-G. Wu. 1996. Ultrastructural evidence for the placement of the truffle Barssia in the Helvellaceae (Pezizales). Mycologia 88 (1): 38–46.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

Lumbsch, H. T., & S. M. Huhndorf (eds) 2007. Outline of Ascomycota—2007. Myconet 13: 1–58.

[LK04] Lutzoni, F., F. Kauff, C. J. Cox, D. McLaughlin, G. Celio, B. Dentinger, M. Padamsee, D. Hibbett, T. Y. James, E. Baloch, M. Grube, V. Reeb, V. Hofstetter, C. Schoch, A. E. Arnold, J. Miadlikowska, J. Spatafora, D. Johnson, S. Hambleton, M. Crockett, R. Shoemaker, G.-H. Sung, R. Lücking, T. Lumbsch, K. O’Donnell, M. Binder, P. Diederich, D. Ertz, C. Gueidan, K. Hansen, R. C. Harris, K. Hosaka, Y.-W. Lim, B. Matheny, H. Nishida, D. Pfister, J. Rogers, A. Rossman, I. Schmitt, H. Sipman, J. Stone, J. Sugiyama, R. Yahr & R. Vilgalys. 2004. Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. American Journal of Botany 91 (10): 1446–1480.

[PA-W02] Pang, K.-L., M. A. Abdel-Wahab, S. Sivichai, H. M. El-Sharouney & E. B. G. Jones. 2002. Jahnulales (Dothideomycetes, Ascomycota): a new order of lignicolous freshwater ascomycetes. Mycological Research 106 (9): 1031–1042.

Percudani, R., A. Trevisi, A. Zambonelli & S. Ottonello. 1999. Molecular phylogeny of truffles (Pezizales: Terfeziaceae, Tuberaceae) derived from nuclear rDNA sequence analysis. Molecular Phylogenetics and Evolution 13 (1): 169–180.

[SL02] Schweigkofler, W., K. Lopandic, O. Molnár & H. Prillinger. 2002. Analysis of phylogenetic relationships among Ascomycota with yeast phases using ribosomal DNA sequences and cell wall sugars. Organisms Diversity & Evolution 2: 1–17.

Underwood, L. M. 1896. On the distribution of the North American Helvellales. Minnesota Botanical Studies Bulletin 9 (8): 483–500.

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