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Cercophora rubrotuberculata sp. nov., a new pyrenomycete from the Great Smoky Mountains National Park.

     Section for Biodiversity, Illinois Natural History Survey, 1816 S. Oak Street, Champaign, Illinois 61820-6970

George K. Mugambi

     Botany Department, The Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496, and University of Illinois at Chicago, Department of Biological Sciences, Chicago, Illinois 60607-7060

Sabine M. Huhndorf

     Botany Department, The Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605-2496

	ABSTRACT

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An interesting wood-inhabiting pyrenomycete was discovered while collecting for the All Taxa Biodiversity Inventory currently being conducted in the Great Smoky Mountains National Park. This species is unique in possessing superficial ascomata with reddish tubercles and ascospores that develop an apical swollen brown cell and a long, basal hyaline cell. Because these ascospore characters fit the traditional morphological circumscription of Cercophora, this species is described as a new species within this genus.

Key words: Ascomycota, Lasiosphaeriaceae, Sordariales, southern Appalachians, systematics, temperate forests

	INTRODUCTION

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A unique member of the Sordariales (Sordariomycetes, Pezizomycotina, Ascomycota) was found on decaying wood during a weeklong "bioblitz," a taxon-discovery effort conducted as part of an All Taxa Biodiversity Inventory (ATBI) currently taking place in the Great Smoky Mountains National Park (GSMNP) (Miller 2006). This species is unique in possessing superficial, obpyriform ascomata with large, reddish tubercles, cylindrical asci with subapical globules and ascospores that develop an apical swollen cell and a basal cylindrical cell. Most genera within the Sordariales are distinguished by ascospore shape (Lundqvist 1972, Miller and Huhndorf 2005) and ascospore morphology suggests this taxon should be placed in the genus Cercophora in the Lasiosphaeriaceae (Lundqvist 1972). Although recent molecular studies have shown that many genera in the Sordariales, including Cercophora, are polyphyletic (Miller and Huhndorf 2005), additional study is required before generic relationships within the order can be resolved and revised circumscriptions of genera can be proposed. Therefore this species is placed within Cercophora based on morphological characters.

	MATERIALS AND METHODS

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Morphological characterization.— – Ascomata were squash-mounted in water and images of micromorphological structures were captured with a QImaging QColor 3 digital camera mounted on either a Leica MZ7.5 dissecting microscope with a Schott KL1500 fiber optics light source or an Olympus BX51 compound microscope using differential interference or phase contrast microscopy. Images were processed with Adobe Photoshop 7.0 (Adobe Systems Inc., Mountain View, California). A minimum of 30 measurements was taken for all morphological structures with NIH Image 1.63 (National Institute of Health, Bethesda, Maryland). Mean and standard deviation (shown in brackets) were calculated for ascospores. Ascomata were embedded in Tissue-Tek^® O.C.T. embedding compound and sectioned with a Leica SM2000R freezing microtome at ca. 30 µm thick. Color terms are taken from Kornerup & Wanscher (1978). No attempts were made to culture this species.

	RESULTS

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Cercophora rubrotuberculata A.N. Mill., Mugambi & Huhndorf, sp. nov. FIGS. 1–11

View larger version (108K): [in this window] [in a new window]   FIGS. 1–11. Cercophora rubrotuberculata (all from Holotype). 1–3. Ascomata on substrate. 4. Longitudinal section through ascoma. 5. Longitudinal section through ascomal wall showing reddish crystalline exudate covering tubercle. 6. Longitudinal section through ascomal wall. 7, 8. Asci. 9. Ascus apex. 10. Ascospore producing phialides. 11. Paraphyses. Bars: 1 = 500 µm; 2–4 = 100 µm; 5–11 = 10 µm.

Etymology.. Refers to the reddish tuberculate surface of the ascomata.

Ascomata ovoidea vel obpyriformia, papillata, 300–550 µm diam, 450–900 µm alta, numerosa, dispersa vel gregaria, superficialia; superficie tuberculata subcollo, tubercula aurantiaca deinde rubella; collo conico, interdum sulcato, nigro; subiculum sparsum, 1.5–3.0 µm crassum, pilis brunneis. Paries ascomatis in sectione longitudinali bistratosus. Paraphyses filiformes, septatae. Asci cylindrici, 190–270 x 18–24 µm, longe-stipitati, stipus 50–95 x 3.5–6.5 µm, unitunicatus, cum annulo apicali et globulo sub-apicali, octospori. Ascosporae cylindricae, 54–64.5 x 4.5–6 µm, hyalinae, aseptatae; utrinque caudatis gelatinosis, 16–19 µm; deinde bicellularis, cellula superior inflata, ovoidea, 16–22.5 x 7.5–9.5 µm, cellula basis 34.5–45.5 x 4.5–5 µm, hyalina; demum ascosporae triseptatae usque ad 7-septatae.

Ascomata ovoid to obpyriform, papillate, occasionally collapsing laterally when dried, 300–550 µm diam, 450–900 µm high, numerous, scattered to gregarious, superficial; surface tuberculate below neck, tubercles large, at first orange red (8A8), becoming reddish (9B8) with age; neck conical, sometimes sulcate, black; subiculum sparse, hairs brown, 1.5–3.0 µm wide, thin-walled, septate. Ascomatal wall of textura angularis in surface view; in longitudinal section 2-layered, composed of pseudoparenchymatous cells, inner layer 15–30 µm thick, composed of 3–5 layers of elongate to flattened, brown cells, outer layer 30–80 µm thick, composed of 5–8 layers of polygonal, brown cells; tubercles 50–90 µm wide x 60–75 µm tall, composed of polygonal, brownish cells, apex encrusted with a reddish exudate that does not dissolve in water. Ascomatal apex with periphyses. Centrum hyaline. Paraphyses filiform, 3.0–7.0 µm wide, hyaline, numerous, septate, unbranched, persistent. Asci cylindrical, 190–270 x 18–24 µm, long-stipitate, stipe 50–95 x 3.5–6.5 µm, numerous, unitunicate, thin-walled, apex truncate; ring narrow, shallow, refractive; subapical globule large, 4.5–6.5 µm diam, smooth; with 8 biseriate to triseriate ascospores. Ascospores cylindrical, ends rounded, 54–64.5 x 4.5–6 µm (58.5 ± 2.5 x 5 ± 0.5), slightly sigmoid or geniculate, hyaline, aseptate; with bipolar appendages, 16–19 µm, gelatinous, lash-like; becoming differentiated into a swollen head and pedicel, transversely uniseptate; head ovoid, 16–22.5 x 7.5–9.5 µm, subacute to rounded at the apex, truncate at the base, hyaline to pale brown; pedicel 34.5–45.5 x 4.5–5 µm, hyaline; ascospore 3–7-septate after liberation from the ascus, occasionally producing phialides directly from the ascospore.

Habitat.. Scattered to gregarious on dead, decorticated wood in mixed coniferous-deciduous forest.

Distribution.. Known only from type locality.

	DISCUSSION

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The genus Cercophora Fuckel, which was established in 1870, was synonymized under Sordaria Ces. & De Not. (Fuckel 1873) and seldom used until it was reintroduced by Lundqvist (1972). The genus is recognized by large, membranous to coriaceous ascomata and hyaline, cylindrical ascospores, which develop an apical, swollen brown cell and a long, basal hyaline cell. Cercophora currently contains more than 50 species of lignicolous and coprophilous taxa that occur throughout temperate and tropical regions. Although the genus recently has been shown to be polyphyletic (Miller and Huhndorf 2005), additional work is necessary before Cercophora can be segregated into several distinct, well supported mono-phyletic groups. Thus new species, which fit the traditional morphological circumscription of Cercophora, should be placed within the genus until a thorough revision can be completed.

Cercophora rubrotuberculata is distinguished in having ascomata with reddish tubercles and ascospores that develop an apical swollen cell and a basal cylindrical cell. Both ascomatal characters (i.e. reddish color and tuberculate surface) are uncommon in Cercophora. Only two other species in the genus possess ascomata with reddish coloration, Cercophora spirillospora (Penz. & Sacc.) N. Lundq. and Cercophora citrina (Petch) N. Lundq. The ascomata of the type specimen of C. spirillospora presently are yellowish-brown, however they originally were described as being reddish when fresh (Penzig and Saccardo 1897). Cercophora spirillospora is easily distinguished from C. rubrotuberculata in the type of substrate (bamboo culms vs. wood), ascomatal surface (tomentose vs. tuberculate), centrum color (yellowish vs. hyaline) and ascospore size (37–45 vs. 54–64.5 µm). Cercophora citrina occurs on elephant dung and differs in possessing smooth ascomata with a reddish tomentum, yellow centrum contents and longer ascospores (72–89 µm).

Species with tuberculate or warty ascomata also are uncommon in Cercophora. Cercophora ambigua (Sacc.) R. Hilber and C. arenicola R. Hilber are described as having tuberculate ascomata (Hilber and Hilber 1979). While these two species are similar, they can be distinguished from C. rubrotuberculata in having dark brown to black, sparsely setose ascomata. The ascomata in Cercophora atropurpurea A.N. Mill. and Huhndorf also are described as being warty, primarily around the neck (Miller and Huhndorf 2001), but this species possesses dark purple ascomata and shorter ascospores (33–51 vs. 54–64.5 µm).

Although C. rubrotuberculata was not grown in culture, it most likely possesses a Phialophora-like anamorph because phialides were observed germinating directly from the ascospores (FIG. 10). This characteristic has been described in C. ambigua (Hilber and Hilber 1979) and more recently in Lasiosphaeria ovina (Pers. : Fr.) Ces. & de Not. and L. sorbina (Nyl.) P. Karst. (Miller and Huhndorf 2004). Several species of Cercophora and Lasiosphaeria are known to possess Phialophora-like anamorphs (Gams 1973, Gams and Holubová-Jechová 1976, Gams 2000, Miller and Huhndorf 2001, 2004).

	ACKNOWLEDGMENTS

 
This species was discovered during fieldwork supported by a Discover Life in America grant (DLIA2005-11) to ANM, while the remaining research was supported by a National Science Foundation grant (DEB-0515558) to ANM. The authors thank J.L. Crane for his comments on a draft of this manuscript.

	FOOTNOTES

 
Accepted for publication March 27, 2007.

¹ Corresponding author. E-mail: amiller{at}inhs.uiuc.edu

	LITERATURE CITED

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
Fuckel L. 1873. Symbolae mycologicae. Beitrage zur Kenntniss der rheinischen Pilze. Zweiter Nachtrag. Jahrb Nass Ver Naturk 27–28:1–99.

Gams W. 1973. Phialides with solitary conidia? Persoonia 7:161–169.

———. 2000. Phialophora and some similar morphologically little-differentiated anamorphs of divergent ascomycetes. Stud Mycol 45:187–199.

———, Holubová-Jechová V. 1976. Chloridium and some other dematiaceous hyphomycetes growing on decaying wood. Stud Mycol 13:1–99.[Medline]

Hilber R, Hilber O. 1979. Einige anmerkungen zu der gattung Cercophora Fuckel (Lasiosphaeriaceae). Z Mykol 45:209–233.

Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3rd ed. London: Eyre Methuen. 252 p.

Lundqvist N. 1972. Nordic Sordariaceae s. lat. Symbol Botanic Upsalienses 20:1–374.

Miller AN, Huhndorf SM. 2001. Neotropical Ascomycetes 10. New and interesting Cercophora species. Sydowia 53(2):211–226.

———, ———. 2004. Using phylogenetic species recognition to delimit species boundaries within Lasiosphaeria. Mycologia 96(5):1106–1127.[Abstract/Free Full Text]

———, ———. 2005. Multi-gene phylogenies indicate ascomal wall morphology is a better predictor of phylogenetic relationships than ascospore morphology in the Sordariales. Mol Phylo Evol 35:60–75.[CrossRef][Medline]

———. 2006. Pyreno pursuit: a mycoblitz to discover the diversity of pyrenomycetes in the Great Smoky Mountains National Park. Inoculum 57(2):1–3.

Penzig O, Saccardo PA. 1897. Diagnoses fungorum novorum in insula Java collectorum 1. Malpighia 11:387–411, 491–530.

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