(Ascomycetes)—IV. Morchellaceae, Helvellaceae, Rhizinaceae

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(Ascomycetes)—IV. Morchellaceae, Helvellaceae, Rhizinaceae PERSOONIA Published by the Rijksherbarium, Leiden Volume Part 9, 1, pp. 1-38 (1976) Ultrastructure of the ascospore wall in Pezizales (Ascomycetes)—IV. Morchellaceae, Helvellaceae, Rhizinaceae, Thelebolaceae, and Sarcoscyphaceae. General discussion Emily Merkus Rijks herbarium, Leiden* Plates (With 1-12, two Tables, and one Text-figure) The of wall and ornamentation of development layers ascospores, plus studied in specialized plasmic structures, are with the electron microscope members of the Morchellaceae, Helvellaceae, Rhizinaceae, Thelebolaceae, and Sarcoscyphaceae; additional results in the Pyronemataceae are given. The of former studies findings are compared with those (Merkus, 1973, 1974, 1975). A general discussion on the origin and development of the different the series. A based structures concludes classification upon species representing typical developments of all the species of the four studies is made. INTRODUCTION Following former publications on the subject (Merkus, 1973, 1974, 1975) this series electron microscopy study completes the of papers on the ultrastructure of the wall in Pezizales. The includes the results in ascospore present paper species belonging to the Morchellaceae, Helvellaceae, Rhizinaceae, Thelebolaceae, and additional results Sarcoscyphaceae. It also gives in the Pyronemataceae sensu Eckblad, including those in Pyronema omphalodes, Coprobia granulata, Geopyxis carbonaria, Mycolachnea hemisphaerica, and Octospora musci-muralis. In general, Eckblad's classification of the taxa (1968) has been followed. Thecotheus ornamented and spec, (an eight-spored, species) Iodophanus carneus are provisionally classifiedin the Eckblad in the Thelebolaceae; e.g. placed Iodophanus Pyronemataceae. According to Eckblad a member ofthe Otideaceae, Geopyxis carbonaria, is transferred Eckblad. provisionally to the Pyronemataceae sensu concludes with discussion the The paper a general on subject. * Present address: National Institute of Public Health, Bilthoven. I Persoonia Vol. Part 1976 2 9, i, REVIEW OF EARLIER WORK Only a few of the species studied by Le Gal (1947) could be examined under the electron microscope. Le Gal's light microscopy of these species is briefly reviewed. Some of the names she used (see footnotes) have been changed in accordance with the 'International Code of Botanical Nomenclature'. 1 Discina Rhizina undulata Fr. per Pers. and the related perlata (Fr. per Fr.) Fr. ornamentation that is formed between the wall and develop a simple spore primary its covering layers. The ornamentation consists of callose and pectine and is of sporal origin. The primary wall is covered by an "assise sous-perisporique" and a before "pelliculemembranaire". The "assise sous-perisporique" is formed the ornamen- tation develops. The "pellicule membranaire"is termed "coque interperisporique", as it is formed at the same time as the ornamentationand is penetrated by the sub- stance ofornamentation; the "coque interperisporique" and the substance of the ornamentation both grow into one, the 'coque interperisporique' also consisting of callose and pectine. During the development of the ornamentation a "perispore" is the outside of the this remains. present on ascospores; "perispore" Helvetia atra Holmskj. Fr. 2 Helvetia elastica Bull, St.-Am. 3 Helvetia per , per , 4 5 acetabulum (L. St.-Am.) Quel. Helvetia macropus (Pers. S. F. Gray) P. Karst. per , per and some related species, all members of the Helvelleae of Le Gal (1947: 284), which in Helvetia smooth are now placed L. per St.-Am., normally develop ascospores. But in few the show ornamentation the a cases species an on ascospores, consisting of of connected isolated Le Gal this coarse folds or or warts. According to ornamenta- tion is false and substance in "couche formed by a situated a sous-episporique", which deforms the epispore locally. materialofthe ofthe studied Gal Unfortunately, no species Sarcoscyphaceae by Le available. coccinea S. F. Lamb, was Instead Sarcoscypha (Scop, per Gray) and Desma- zierella acicola Lib., which also belong to this important taxon, could be studied. Le Gal's descriptions of the ornamentationpatterns ofthe species involved resemble those of Rhizina undulata and Discina perlata. The "perispore" is permanent in all which contain its species; the ornamentation, does not callose and pectine, penetrates covering layers in Cookeina sulcipes (Berk.) O. K., Cookeina tricholoma (Mont.) O. K. and Cookeina insititia (Berk. & Curt.) O. K., and remains within these layers in Aurophora dochmia (Berk. & Curt, apud Berk.) Rifai 6 domingensis Berk., , Phillipsia Plectania Nannf. Korf 7 and Plectania Rifai8 campylospora (Berk.) apud , platensis (Speg.) . 1 Karst. Rhizina inflata Schaeff. per P. 2 Leptopodia atra (Fr. ex Konig) Boud. 3 Leptopodia elastica (Bull.) Boud. 4 Helvetia sulcata Afz. 5 Macropodia macropus (Pers.) Fckl. 6 Phillipsia dochmia (Berk. & Curt, apud Berk.) Seaver. 7 Sarcosoma sarasini (P. Henn.) Boud. 8 Umula platensis Speg. Merkus: Ultrastructure of ascospore wall in Pezizales—IV 3 MATERIALS AND METHODS Fresh material was collected in the Netherlands, Belgium, Turkey, Papua New and in Guinea, Indonesia, Kenya, Uganda, Canada; some species were provided culture the Centraalbureau The Netherlands. pure by voor Schimmelcultures, Baarn, details about their The following list gives more the specimens and origins: coemansii Boud. of — ‘Ascophanus’ (a species Coprotus) van Brummelen, culture on cow dung, 21.VI. 1973 (L); — Piepenbroek, on cow dung, near Welsum, Olst, Overijsel, The Netherlands; Ascozonus woolhopensis (Berk. & Br.) Boud. — van Brummelen, several cultures isolated from rat dung, on oatmeal agar enriched with horse dung Boud. — decoction, Ekeren, Belgium (L) ; Coprobia granulata (Bull, per Merat) van Brummelen Hollandse 3562, culture on cow dung, Rading, Maartensdijk, Utrecht, The Netherlands — culture (L) ; van Brummelen, on cow dung, Bathmen, Overijsel, Lib. — The Netherlands, 6. VIII. 1973 (L); Desmazierella acicola J. Daams, on fallen needles of Pinus, "Spanderswoud", Hilversum, North Holland, The Netherlands, 11. V. 1973; Geopyxis carbonaria (Alb. & Schw. per Pers.) Sacc. — van Brummelen & Piepenbroek 4082, on burnt soil, 't Joppe, Gorssel, Gelderland, The Netherlands, 6. VIII. 1973 (L); - Piepenbroek 783, on burnt soil, 't Joppe, Gorssel, Gelderland, The esculenta Fr. — & Netherlands (L); Gyromitra (Pers. per Krombh.) Tjallingii rubbish in a The Nether- van Brummelen, among garden, Wageningen, Gelderland, lands, 2. IV. 1974 (L); Helvella crispa Scop, per Fr. — van Brummelen 4073, under North The — Pinus, Bergen, Holland, Netherlands, 3. XI. 1973 (L); van Brummelen 4506, under deciduous trees beside a road, Diepenveen, Overijsel, The Netherlands, VII. musci-muralis Graddon — 15. 1974 (L); Octospora Piepenbroek 720, among mosses on an old wallin a garden, between Deventer and Twello, Voorst, Gelderland, The Korf Kimbr. Netherlands, 12.XI. 1973 (L); Iodophanus carneus (Pers. per Pers.) apud & Korf— van Brummelen, culture on sheep dung, Elspeet, Gelderland, The Nether- lands, 7. XI. 1972 (L) ; — van Brummelen, culture, Vogelenzang, North Holland, The Netherlands — culture with (L) ; Prof. Necat, on oatmeal agar enriched horse dung Lasiobolus — decoction, Kaskaloglu, Bornova, Izmir, Turkey; monascus Kimbr. van Brummelen several cultures isolated from oatmeal 3563, porcupine dung, on agar enriched with horse dung decoction, Mt. Suckling, Papua New Guinea, 6.IX.1972 Lasiobolus Sacc. — (L); pilosus (Fr.) van Brummelen, culture on sheep dung, Elspeet, The — Brummelen Gelderland, Netherlands, 7.XI.1972 (L); van 4077, culture on horse dung, Overveen, Bloemendaal, North Holland, The Netherlands, 14.V. 1973 hartebeest (L)J- van Brummelen, culture on ( Alcelaphus caama) dung, zoological The Netherlands Morchella esculenta garden, Amsterdam, North Holland, (L) ; L. — C. Bas South per St.-Am. 6003, on sandy soil, Meyendel, Wassenaar, Holland, The — C. under Netherlands, 16.IV.1974 (L); Bas, on sandy soil-peaty soil, Populus and Crataegus, Voorschoten, South Holland, The Netherlands, 22. IV.1975 (L); Persoonia Part 4 Vol. g, I, 1976 S. F. Maire — Mycolachnea hemisphaerica (Wigg. per Gray) Piepenbroek j6g, on sandy soil, Epserbos, Epse, Gelderland, The Netherlands, 21.VII. 1974 (L); Pyronema omphalodes (Bull.) per St.-Am.) Fckl. — van Brummelen 4.081, on burnt soil, 't Joppe, The 6.VIII. — CBS Gorssel, Gelderland, Netherlands, 1973 (L); 373.62, several cul- tures cornmeal Rhizina undulata Fr. Pers. — Brummelen on agar (L); per van 4076, on burnt soil, 't Joppe, Gorssel, Gelderland, The Netherlands, 6.VIII.1973 (L); — Piepenbroek 761, on burnt soil, "de Poll" between Wilp and Voorst, Gelderland, The Netherlands, 20.VII.1974 (L); Sarcoscypha coccinea (Scop, per S. F. Gray) Lamb. — of unknown (probably Central European) origin (comm. Mr. G. D. Swanenburg de Thecotheus ■— culture rhinoceros P. Veye), 8.III.1974 (L); spec. on dung, Pentjang, W. Java, Indonesia, 28.X.1972 (L); Thelebolus crustaceus (Fckl.) — CBS 713.69, isolated enriched culture from carnivore dung, on oatmeal agar with horse dung decoction, R. F. Cain & al., Mt. Speke, Ruvenzori Mts, Uganda, 23.VII.1969 (L); Tode Steudel — isolated from Thelebolus stercoreus per CBS 717.69, culture deer oatmeal enriched with horse D. Simcoe dung, on agar dung decoction, Malloch, Co., Ontario, Canada, 13.V. 1968 (L). collections As not all field contained ripening apothecia some specimen were kept moist in conditioned growth chambers, in alternating light and darkness; Lasiobolus
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