Antarctic Science 18 (3), 291–301 (2006) © Antarctic Science Ltd Printed in the UK DOI: 10.1017/S0954102006000344 Lichenicolous fungi described from Antarctica and the subAntarctic islands by Carroll W. Dodge (1895–1988) DAVID L. HAWKSWORTH1* and TERESA ITURRIAGA2 1 Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, Plaza Ramón y Cajal, Ciudad Universitaria, ES-28040 Madrid, Spain 2 Departamento Biología de Organismos, Universidad Simón Bolívar, Apartado 89000 Sartenejas, Baruta, Edo. Miranda, Venezuela *Address for correspondence: MycoNova, The Yellow House, Calle Aguila 12, Colonia La Maliciosa, Mataelpino, ES-28492 Madrid, Spain myconova@terra.es Abstract: The identities of the 12 lichenicolous fungi described as new to science from the Antarctic and sub-Antarctic islands by Carroll W. Dodge between 1938 and 1968 are reassessed on the basis of the original publications, and where extant the original collections and slides in the Farlow Herbarium and Reference Library (Harvard University, Cambridge, MA, USA). Typifications are discussed and lectotypes and neotypes designated as necessary. Only one species is accepted in its original placement (Phacopsis usneae) and one had already been transferred to an appropriate genus (Didymella cladoniae to Zwackhiomyces). Six prove to be synonyms of previously described species, and four are combined into other genera here: Arthonia parmeliae (syn. Diplonaevia parmeliae), Carbonea antarctica (syn. Alectoria antarctica), Sphaerellothecium buelliae (syn. Orbicula buelliae), and Weddelomyces gasparriniae (syn. Phaeospora gasparriniae) combs. nov. The generic name Thelidiola is also confirmed as a synonym of Muellerella. In addition, as a result of earlier misinterpretations of Dodge’s taxa, two new species are described: Endococcus matzeri, and Zwachkiomyces diederichii spp. nov. Intralichen lichenum is also reported from the Antarctic for the first time. Descriptions and illustrations are included where appropriate. Received 4 October 2005, accepted 30 January 2006 Key words: Ascomycota, discomycetes, hyphomycetes, lichens, pyrenomycetes, taxonomy Introduction published extensively on those from little-explored regions, especially Africa, Antarctica, and South America until 1973, aged 78 years. When he started to become interested in lichens around 1912–15, there were only two active lichenologists in the whole of the USA, and they were based in the southern states. In consequence, he worked in isolation, and relied largely on literature accounts for his interpretation of previously published species. Although he always produced lengthy descriptions including detailed microscopical observations, his concepts of genera and species were often idiosyncratic and re-evaluations of the taxa he described as new are essential. The first author corresponded with Dodge from 1967–74, mainly with respect to alectorioid lichens, and was privileged to receive various collections and slides from him on loan during a period when he had ceased to communicate with many lichenologists; this provided valuable insights into his ways of working. For many years his collections were deteriorating in a garage at his home, prior to their transfer to the Farlow Herbarium and Reference Library (Harvard University, Cambridge, MA, USA; FH) in 1985. A staggering 75 000 collections were accessioned, including those of non-lichenized fungi, and after laborious conservation work they started to become available for study in the early 1990s. Detailed biographical information, and a full list of Dodge’s publications, is provided by Fungi obligately occurring on lichens as parasites, saprobes or commensals represent a major ecological niche for fungi. There are over 1500 currently known lichenicolous species, most of which belong to genera only known from lichen hosts (Lawrey & Diederich 2003). In view of the predominance of lichens in the Antarctic (Øvstedal & Smith 2001), numerous lichenicolous fungi are to be expected growing on them, but while there are scattered reports, this ecological niche has not yet been examined in depth. A reappraisal of the already documented biota is a prerequisite for further research, and this paper seeks to address the identities of the lichenicolous taxa described as new to science from the Antarctic and sub-Antarctic islands by Carroll W. Dodge (1895–1988) over the period 1938–68. Dodge was an enigmatic Vermont figure who made major well-respected contributions to medical mycology (Dodge 1936), and hypogeous fungi (e.g. Dodge & Zeller 1934). He held positions at Harvard University (1921–31) and the Missouri Botanical Garden (1931–63), and as a Research Professor in the University of Vermont (1963 on) after his retirement; he also taught courses on medical mycology in different South American countries from 1934–60. However, he also had an absorbing interest in lichens which went back to his undergraduate days, and these became his main preoccupation after his retirement in 1963. He 291 http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 202.75.16.18 292 D.L. HAWKSWORTH & T. ITURRIAGA Fig. 1. Carbonea antarctica. Reproduction of the published illustration by Dodge & Baker 1938: pl. 51, figs 209–216. Fig. 209, habit sketch, x 6; fig. 210, detail of thallus ends with apothecium, x 35; fig. 211, section of apothecium, x 55; fig. 212, development of ascus through maturity, x 1100; fig. 213, paraphysis, x 1100; fig. 214, mature spores, x 1100; fig. 215a, hypha from medulla, x 1100; fig. 215b, detail of cortex, x 1100; and fig. 216, crosssection of thallus, x 177. Rudolph (1990). With respect to the lichens he described as new from Antarctica, their identities have now mostly been resolved based on a re-examination of the original collections (Hale 1987, Castello & Nimis 1995, Øvstedal & Smith 2001). However, with his broad mycological background, it was perhaps not surprising that he was also interested in the fungi which grow on lichens. At the same time, he also described as novel “lichens” some lichens which were actually infected by a lichenicolous fungus. Hawksworth (in Pegler et al. 1980) endeavoured to make some reassessments of his Antarctic lichenicolous species, but at that time Dodge’s material was not available for study. This contribution provides a fresh evaluation of all the lichenicolous fungi described from Antarctica and the subAntarctic islands by Dodge, whether as lichenicolous fungi or erroneously as lichens, based on the original descriptions and illustrations and where extant the original collections and microscope slides now preserved in FH. For more general information on the biology, ecology, and systematics of lichenicolous fungi, see Lawrey & Diederich (2003) and Hawksworth (2003, 2004). Systematics (1) Alectoria antarctica C.W. Dodge & G.E. Baker, Ann. Mo. Bot. Gdn 25: 599 (1938). Type: Marie Byrd Land, Edsel Ford Range, Mt Rea-Cooper, on Pseudephebe minuscula, 1934, P. Siple, F.A. Wade, S. Corey & O.D. Stancliff R-1 (FH – slide with gummed label, lichenicolous fungus element – lectotypus hic designatus). ≡ Carbonea antarctica (C.W. Dodge & G.E. Baker) D. Hawksw. & Iturr., comb. nov. (Fig. 1) http://journals.cambridge.org Downloaded: 11 Dec 2013 Lamb (1964, p. 28) reports that he was sent a slide of this taxon by Dodge, along with two other “authentic specimens” from the same locality, but that only the slide had ascomata and the other specimens were typical of the species now called Pseudephebe minuscula. He considered the ascomata either to be abnormal ones of the lichen or of a “parasitic or parasymbiotic Ascomycete (perhaps a species of Nesolechia)” and while listing the name as a synonym of the lichen, as had Murrray (1963), no formal lectotypification was made. Dodge also sent the first author a single slide (perhaps the same one) on loan in 1967, from which it was evident that the ascomata represented a lichenicolous fungus. Three slides are now present in FH, one with a gummed label giving abbreviated type locality information and labelled “Rea-1”, and the two others with “type” (also marked “Mt Rea. 7”) and “type 1” (also marked “Mt Rea-1”) written on the glass, all in Dodge’s hand. The slide labelled “type 1” only has microtome sections of the thallus and none of the ascomata. In the absence of a specimen bearing ascomata from which the slides could have been made, we consequently designate the lichenicolous element on the slide with the gummed label as lectotype for the name here as that has the best preparations showing the exciple structure and paraphyses as well as the “Rea-1” annotation. We selected the lichenicolous fungus as lectotype rather than the host lichen as it was the characters of the fungus that Dodge regarded as distinctive in lacking a thalline exciple (“ampithecium”) to the extent that “it might be considered the type of a new genus and family in the lecideine series of families” (Dodge & Baker 1938, p. 598). As pointed out by Lamb, the ascomata definitely appear to be lecideine and have a well-developed dark brown exciple, without any trace of algae to suggest the thalline exciple typical of Pseudephebe was present. This is supported by the published illustrations of the apothecia and IP address: 202.75.16.18 293 LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE the section (Fig. 1, figs 210 & 211). Our studies of the slides show the exciple to be carbonaceous and composed of compacted angular thick-walled pseudoparenchymatous cells 6–10 x 4–8 μm diameter, with walls 2.5–3 μm thick and a small lumen, forming a textura angularis to textura globulosa. The asci are lecanoralean, 38–35 x 7.5–11 μm, and the ascospores are hyaline, narrowly ellipsoid, somewhat curved in lateral view, with one end frequently broader than the other, and 6–8 x 3–4.5 μm – much narrower and more ellipsoid than in P. minuscula where they measure 6.7–9 x 7.7–8 μm (Brodo & Hawksworth 1977 p. 141). Further, the paraphyses are repeatedly anastomosed and with lumina only 1–1.5 μm thick. These features all indicate that the ascomata are not deformed ones of the host lichen, but of a lichenicolous fungus that is most appropriately placed in Carbonea (Triebel 1989), a genus including both lichen-forming and lichenicolous species; a formal transfer to that genus is therefore made here. Carbonea antarctica differs from the two other recognized Antarctic species (Øvstedal & Smith 2001) and others described in the genus in the sizes of the ascospores, as well as being the first to be recognized growing on a macrolichen. These authors also mention, but do not illustrate or describe in detail an unnamed “Lecanora sp. D” on Pseudephebe minuscula which we suspect might be the same species; although placement in that genus suggests a thalline exciple including algal cells was present, and the asci and ascospores were given as c. 20 x 8 μm and c. 8 x 6 μm, respectively. We also noted in the sections mounted on the lectotype slide hyphae immersed in the hymenium which evidently represent the hyphomycete Intralichen lichenum (Diederich) D. Hawksw. & M.S. Cole (Hawksworth & Cole 2002) which appears to be the first report of this species from Antarctica. (2) Didymella cladoniae C.W. Dodge, BANZARE 1929–31, series B, 7: 262 (1948). Type: Macquarie Island, north end, Station 81, on “Cladonia mawsonii” [C. cervicornis subsp. mawsonii], BANZARE B540-1 (FH- holotype [two slides]). ≡ Zwackhiomyces cladoniae (C.W. Dodge) Diederich, in Aptroot et al., Bibltheca Lich. 64: 210 (1997). (Fig. 2) Fig. 2. Zwackhiomyces cladoniae (FH - holotype). a. mature ascospores, b. maturing ascus and c. anastomosing paraphysoids. Bar = 10 μm. http://journals.cambridge.org Downloaded: 11 Dec 2013 Ascomata perithecioid, black, 300–425 μm diameter; walls dark brown, pseudoparenchymatous, textura angularis, individidual cells 4.5–19 x 4.5–15.5 μm, cell walls c. 2 μm thick. Hamathecium persistent, composed of abundant branched and anastomosting interascal filaments, width rather uneven but mainly 0.5–1 μm thick. Asci bitunicate in structure, subcylindrical, 70–80 x 8–10 μm, 6(–?8) spored. Ascospores almost uniseriately arranged, slightly overlapping, broadly fusiform, apices generally somewhat pointed, 1-septate, hyaline, coarsely verrucose, the warts in a gelatinous sheath, 12–19 x 3–5 μm. Two slides were located in FH which have abbreviated locality details and the collection number “B540-1”, but not the specimen from which these were made. The two slides are collectively treated as the holotype of Dodge’s name here as they were clearly made from and so are part of the original collection. As the slides had only squash mounts, the above description is not as full as would have been the case with sections or the original specimen. The specimen is indicated as not seen by Grube & Hafellner (1990), and nor was it located by Aptroot et al. (1997) when the species was transferred to Zwackhiomyces. Aptroot et al. examined a fungus on Cladonia yunnana collected on Mount Wilhelm in Papua New Guinea which they considered to be conspecific with Dodge’s taxon on the basis of the original description. However, the perithecia in their material were 75–125(–150) μm diameter, but could reach 170 μm in water, and the asci were shorter (45–63 μm tall); most other measurements they report were very close to those published by Dodge. Sadly, in this case Diederich IP address: 202.75.16.18 294 D.L. HAWKSWORTH & T. ITURRIAGA (in Aptroot et al. 1997) was misled as Dodge´s published measurements of the ascospores, given as “10–11 x 4– 5 μm”, uncharacteristically proved to be inaccurate (see above). The Papua New Guinea material, with substantially shorter asci and smaller ascospores evidently represents a separate species, for which the name Zwackhiomyces diederichii is introduced here. Zwackhiomyces diederichii D. Hawksw. & Iturr., sp. nov. and wider, about 15 x 4 μm” and the range given by Hawksworth (in Pegler et al. 1980) is (12–)13–15(–16) x (4–)4.5–6(–6.5) μm. The species is evidently frequent on lobate and subfruticose species of Caloplaca species and also on Xanthoria elegans in the Antarctic region and on the subantarctic islands (Pegler et al. 1980). (4) Didymosphaeria macquariensis C.W. Dodge, BANZARE 1929–31, series B, 7: 263 (1948). Ascomata semi-erumpescentia, 75–125(–150) μm in diametro. Asci cylindrici, 45–63 x 6.5–7.5 μm, 6–8 spori. Hamathecium ex hyphis ramosis et anastomosantibus, ca 1.5 μm latis. Ascosporae ellipsoideae vel soleiformes, cum apicibus attentuatis, verruculosae, 10–12.5 x 3.5–4 μm. Type: Macquarie Island, Featherbed Flat, on “Pyrenodesmia inclinans”, BANZARE B533-7 (holotype – not located). Typus: Papua New Guinea, Simbu, Mount Wilhelm, Pindaunde Valley, near the hut, c. 3600 m, on Cladonia yunnana, 1992, A. Aptroot 23663 (herb. Diederich – holotypus; herb. Aptroot – isotypus). This species was stated to differ from D. kuttlingeriae in the “larger, thicker-walled perithecia, more numerous and broader asci and slightly larger ascospores” (Dodge 1948, p. 264). However, the perithecium size was not given in the description, and the wall thickness can reach the c. 20 μm cited near the ostiole in P. rugulosarium. Further, the ascospores were given as 15–16 x 4–5 μm which is within the range of the species (see under D. kuttlingeriae above). However, the ascus size was given as “c. 35 x 15 μm” which must either be an error or based on a very immature ascus; were this correct, it is inconceivable that it could accommodate the eight spores cited. On the basis of the information available in the absence of any original material, we see no reason not to treat the name as a synonym of P. rugulosarium as suggested by Hawksworth (in Pegler et al. 1980). For a fuller description and illustrations of the asci and ascospores, see Aptroot et al. (1997, 210–211). (3) Didymosphaeria kuttlingeriae C.W. Dodge, BANZARE 1929–31, series B, 7: 263 (1948). Type: Crozet Archipelago, Possession Island, American Bay, on “Kuttlingeria crozetica” [? Xanthoria elegans], BANZARE B20-22 (holotype – not located). = Polycoccum rugulosarium (Linds.) D. Hawksw., in Pegler et al., Kew Bull. 35: 513 (1980). Also mentioned in the original account from “Blastenia johnstonii” in the type locality (B20-19), and on “B. keroplasta var. athallina” from Kerguelen, Mount Wyville Thompson (B246-22). None of the three specimens could be located in FH, but the very full description agrees in almost all details with that of Polycoccum rugulosarium. For example, the ascospores were given as “dark brown, 2-celled, long ellipsoid to oblong, upper cell slightly longer = Polycoccum rugulosarium (Linds.) D. Hawksw., in Pegler et al., Kew Bull. 35: 513 (1980). (5) Diplonaevia parmeliae C.W. Dodge & G.E. Baker, Ann. Mo. Bot. Gdn 25: 661 (1938). Type: Marie Byrd Land, Edsel Ford Range, Skua Gull Peak, on “Parmelia variolosa” [? Physcia caesia], P. Siple & S. Corey 72W-3 (holotype not located). = Arthonia parmeliae (C.W. Dodge & G.E. Baker) D. Fig. 3. Arthonia parmeliae. Reproduction of the published illustration by Dodge & Baker 1938: pl. 62, figs 396–400. Fig. 396a, detail of hyphae of host, x 1100; fig. 396b, detail of hyphae of parasite, x 1100; fig. 397, paraphysis, x 1100; fig. 398, mature spores, x 1100; fig. 399, section of apothecium, the host surface indicated by dotted lines, x 385; and fig. 400, development of ascus, x 1100. http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 202.75.16.18 295 LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE Hawksw. & Iturr., comb. nov. (Fig. 3) Castello & Nimis (1995) stated that they did not locate the type of the name Parmelia variolosa, a species name introduced by Dodge & Baker (1938), but indicated that it was Physcia caesia on the basis of a specimen they said was an isotype numbered “Siple et al. R-3”. However, no “R-3” is mentioned in the original account, even amongst several paratypes, so the true status of the specimen they studied is unclear. The holotype was stated to be Siple et al. R-1, and we were able to examine two packets with this number in FH (Marie Byrd Land, Mt Rea-Cooper, 77º07'S 145º30'W, on coarse-grained leucogranite, 1934, P. Siple, F.A. Wade, S. Corey & O.D. Stancliff R-1); we consider the packet with an inner envelope name in ink in Dodge’s hand to be the holotype and the other an isotype. An annotation slip indicates that the holotype was examined by Mason E. Hale in 1965, and intriguingly both the holotype and isotype packets have annotation slips by Castello & Nimis from 1994. We confirm that both specimens represent Physcia caesia. We found no lichenicolous fungus in the packets of “R-1”. Diplonaevia parmeliae was described as forming black convex emarginate ascomata up to 150 μm diameter, and having interascal filaments that were straight with slightly enlarged and darkened tips, asci 21–24 x 8–10, and hyaline “apparently 2-celled” ascospores somewhat pointed at one end and 7–8 x 2–2.5 μm. This species is illustrated by Dodge & Baker (1938, pl. 62, figs 396–400), whose drawing is reproduced here as Fig. 3, and shows a vertical section with a dark hypothecium and a thick carbonaceous epithecium arching over the hymenium; in addition, ramifying brown hyphae are seen to penetrate rhizoid-like into the host tissue, and the ascospores figured are narrowly ellipsoid, somewhat narrowed towards the apices, and definitely 1-septate. Asci in different developmental stages, a paraphysis, and hyphae of the fungus and host lichen are also figured. We could find no report of an identical fungus on Physcia caesia, but the details supplied suggest that the species may well belong to Arthonia, although the paraphyses figured are not as anastomosed as would be expected for that genus. There is a species A. epiphyscia Nyl. 1875 which is wellknown on Physcia species, including P. caesia, but that has larger ascospores (10–14 x (3–)3.5–5 μm; Purvis et al. 1992). We also considered a possible placement in Scutula, which would accord more closely with the figured paraphyses, but the dense epithecial tissue figured over the hymenium mitigates against that possibility. In the absence of the holotype specimen, one option would be to designate the published illustration as lectotype but we refrain from doing that here as the specimen may yet be located in FH or fresh material suitable for neotypification may be discovered. A formal transfer to Arthonia is made here, even in the absence of any material, as the fungus should be readily recognizable if recollected and it can evidently be separated Fig. 4. Sphaerellothecium buelliae (FH - lectotype). a. developing ascomata on the surface of the lichen host showing the superficial hyphae, and b. a more mature ascoma. Bars = 10 μm. http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 202.75.16.18 296 D.L. HAWKSWORTH & T. ITURRIAGA from A. epiphyscia by the much shorter and narrower ascospores. (6) Orbicula buelliae C.W. Dodge, BANZARE 1929–31, series B, 7: 261 (1948). Type: Queen Mary Land, Possession Nunatak, c. 66º45'S 98º30'E, on “Buellia muscicola”, 15 Dec. 1912, C.T. Harrisson A.A.E. 12 (FH – element with spherical ascomata on a superficial mycelium, lectotypus hic designatus). ≡ Endococcus buelliae (C.W. Dodge) Matzer, Cryptogamie, Mycol. 14: 14 (1993). ≡ Sphaerellothecium buelliae (C.W. Dodge) D. Hawksw. & Iturr., comb. nov. (Fig. 4) Ascomata superficial, arising on a weft of superficial hyphae on the surface of the host lichen, globose, dark brown to black, (20–)30–55 μm diameter, ostiole not apparent; exciple of 2–3 layers of pseudoparenchymatous cells. Superficial hyphae ramifying over the surface, brown to dark brown, septate, often swollen between the septa, 3–4.5 μm thick, walls 0.5–1 μm thick. Asci and ascospores not seen. The original collection includes two lichenicolous fungi, one described by Dodge with the ascomata occurring on a network of superficial hyphae (this is the only one present on the only slide under this name), and the other with erumpent ascomata not associated with superficial hyphae of which slides were made by Matzer (also preserved in FH) and led him to make the new combination into Endococcus (Matzer 1993). The ascomata and ascospores of the Endococcus were reported by Matzer as 100(–200) μm and c. 18–13 x 6–8 μm respectively, quite different from the measurements of Dodge who gave the ascomata as up to 50 μm diameter and the ascospores “about 6 x 4 μ”. We can only presume that Matzer never studied Dodge’s original slide, and find it inconceivable that Dodge would have been so far out with his microscopic measurements in this description; our experience with other species in the same paper is that his measurement were generally in the same range as those we made. The fungus with small spherical ascomata on the superficial mycelium is clearly the species described by Dodge and that has to be selected as the lectotype element as it is the part that agrees with the protologue. We consider this species to be best accommodated in Sphaerellothecium and therefore make the new combination here. We did not see any mature asci or ascospores, and it seems likely Dodge did not either as he describes them as “hyaline becoming brownish” and does not mention the presence of a septum; we would expect the ascospores to be brown and 1-septate at maturity. Most species in the genus have a narrow host range, none is currently recognized as http://journals.cambridge.org Downloaded: 11 Dec 2013 Fig. 5. Endococcus matzeri (FH – holotype). a. maturing ascus, and b. ascospores. Bar = 10 μm. occurring on Buellia, and the ascospore sizes given by Dodge are smaller than those published for other accepted species in the genus. We therefore make the transfer here, recognizing that a full description will have to await the collection of further material. The second fungus, as Matzer (1993) points out, appears distinct from the known Endococcus species and so we describe this as new below. The host name Dodge used for the host lichen, Buellia muscicola, was a species described by Dodge & Baker (1938) from King Edward VII Land, but this name is not mentioned by Lamb (1968), Castello & Nimis (1995), nor Øvstedal & Smith (2001). We can confirm that it represents a Buellia as apothecia belonging to that genus are present in Harrisson’s collection; the thallus is creamy to whitish, areolate, and the ascospores are 1-septate and 12–15.5 x 7–9 μm, rather short for many of the Antarctic species; the material is so fragmentary that it was impossible to pursue the identity of the host further, but it did not key out clearly in Øvstedal & Smith (2001). Endococcus matzeri D. Hawksw. & Iturr., sp. nov. (Figs 5 & 6) Similis Endococco propinquo, sed ascosporis ornamentatis, asperulatis, 9.5–11.5(–12) x 5–6.5 μm. Typus: Queen Mary Land, Possession Nunatak, c. 66º45'S 98º30'E, on “Buellia muscicola”, 15 Dec. 1912, C.T. Harrisson A.A.E. 12 (FH – element with ellipsoid erumpent ascomata not on a superficial mycelium, holotypus). Ascomata perithecioid, subglobose, ostiolate, immersed at IP address: 202.75.16.18 297 LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE (7.5–)8.5–12.5(–13) x 5–7(–7.5) μm by Triebel (1989), and we did not find any ascospores as wide as the 8 μm he reported. The length:breadth ratios of the ascospores are also in the same range, to judge from numerous collections documented in the folders of notes and drawings on the genus (including the neotype collection in K) of the first author. The key difference is the ascospore ornamentation seen only at high magnifications with quality research microscopes, and most strikingly visible in maturing ascospores where the full wall pigmentation has not developed. This is something not reported before, and, taken with the different host genus, is our reason for recognizing this species here. Previous reports of E. propinquus from Buellia species need to be reassessed to determine if they also have the asperulate ascospores of E. matzeri. (7) Phaeospora gasparriniae C.W. Dodge, BANZARE 1929–31, series B, 7: 262 (1948). Fig. 6. Endococcus matzeri (FH – holotype). a. detail of exciple, the central cavity to the right, b. mature ascus with eight ascospores, and c. immature ascospore showing the ornamented walls. Bar = 10 μm. first but becoming erumpent, black, 140–185 μm diameter; exciple 23–31 μm thick, composed of 8–10 layers of thickwalled angular radially compressed and pseudoparenchymatous cells, textura angularis, individual cells 5–10 x 2–4 μm; lined internally by 3–4 layers of colourless pseudoparenchymatous cells, individually 4–6 x 2.5–4 μm; ostiole not papillate, gaping, 37–75 μm. Hamathecium absent at maturity. Asci bitunicate in structure, broadly clavate, thick-walled, c. 40 x 14 μm, 8-spored. Ascospores irregularly overlapping within the asci, broadly ellipsoid, rounded at the ends, 1-septate, not or only slightly constricted at the septum, cells ± equal in size, appearing ± smooth at lower magnifcations, but faintly but regularly asperulately ornamented at 1500x, without a distinct gelatinous sheath, 9.5–11.5(–12) x 5–6.5 μm, length:breadth ratio 1.5–1.6–2.2. This species is very close to Endococcus propinquus (Körb.) D. Hawksw. 1979 which occurs mainly on lichens in the Porpidiaceae (e.g. Amygdalaria, Porpidia) but is also reported from a variety of other crustose lichens, including Buellia buellioides (Triebel 1989). Matzer (1993) stressed the relatively smaller size of the perithecia, ascospore size, and ascospore ornamentation. We found the ascomata to be in the range of E. propinquus, given as (100–)150–200(–400) by Triebel (1989); we suspect that Matzer's belief that they were mainly to 100 μm may have been due to superficial measurements made on the Sphaerellothecium which he did not recognise as also being present. The ascospore range he gave, “c. 8–13 x 6–8 μm”, is also pretty well in the range of E. propinquus, given as http://journals.cambridge.org Downloaded: 11 Dec 2013 Type: Macquarie Island, north end, on “Gasparrinia maquariensis” [? Caloplaca sp.], BANZARE. B-540-16 (holotype not located). ≡ Weddellomyces gasparriniae (C.W. Dodge) D. Hawksw. & Iturr., comb. nov. The original description indicated that this fungus formed spherical black ostiolate perithecia c. 180 μm diameter, which were semi-immersed when mature. It was described as having both periphyses and discrete unbranched slender paraphyses, cylindrical to narrowly fusiform 8-spored asci c. 65 x 9 μm, and brown thick-walled 3-septate ascospores constricted at the septa and measuring c. 22 x 5 μm. The host was presumably a lobate species of Caloplaca. The presence of interascal filaments precludes a placement in Phaeospora, and that they were described as “slender” suggests they were paraphysoids even though no anastomoses were noted. This feature, along with the thick-walled brown 3-septate ascospores constricted at the septa, indicates that the fungus was almost certainly a species of Weddellomyces. Three species of that genus are known to grow on Caloplaca species in Europe, but all have longer and broader ascospores than the fungus Dodge described: (21.5–)24–31(–38) x (9–)9.5–12(–13.5) μm in W. epicallopisma (Wedd.) D. Hawksw. 1986, (28.5–) 30–40(–45) x (10–)11–15 μm in W. erythrocarpae Nav.–Ros. & Roux 1995, and (24.5–)27–32.5(–39) x (10.5–)11–13.5(–14–5) μm in W. protearius Nav.-Ros. & Roux 1997 (Hawksworth 1986, Navarro-Rosinés & Roux 1995, 1997, Calatayud & Navarro-Rosinés 1998). While we hesitated over making a new combination for this fungus without seeing any material, as the probability that this is the correct genus is high, this decision will ensure that the name is taken into consideration when additional species of the genus on Caloplaca are discovered IP address: 202.75.16.18 298 D.L. HAWKSWORTH & T. ITURRIAGA Fig. 7. Phacopsis usneae (FH – neotype). a. apothecia clustered on a branch of the host lichen, b. paraphyses apices showing granules and the apical caps, c. asci in different stages of development, and d. mature ascospores. Bars = 0.5 mm (a), and 10 μm (b–d). and material suitable for neotypification is found. (8) Phacopsis usneae C.W. Dodge, BANZARE 1929–31, series B, 7: 264 (1948). Type: Kerguelen, Mount Wyville Thompson, 1000–1500 ft, on Usnea trachycarpa, BANZARE B246-21 (holotype not located); Melchior Archipelago, Lystad Island, north face, alt. 20–30 ft, on U. antarctica [as “U. granulifera”], 14 March 1941, McCoy & P.A. Siple 331 (FH – neotypus hic designatus [specimen and slide]). (Fig. 7) Ascomata apothecioid, erumpent at first but soon becoming superficial, arising singly or more usually aggregated into dense mounded clumps, black, the clumps occupying and extending around the host branches and to 2–3 mm in length; individual ascomata strongly convex, circular from above, 0.25–0.75 mm diameter, emarginate, in vertical section 60–90(–100) μm tall; hypothecium dark brown, composed of thick-walled, intertwined hyphae forming angular pseudoparenchymatous tissue in parts, cells mainly 7–10 μm diameter and the hyphae 5–7 μm thick, together forming a compact cushion-like layer varying in thickness; thecium colourless, 45–65 μm tall; epithecium welldeveloped, 12–20 μm tall, dark blue-grey, composed of a mucilaginous gel surrounding and interspersed with the paraphyses tips, binding them together, homogenous but granular on the paraphyses tips. Hamathecium of densely packed paraphyses, repeatedly septate, often somewhat swollen between the septa, not or more commonly sparsely http://journals.cambridge.org Downloaded: 11 Dec 2013 branched and sometimes anastomosed, thick-walled, mainly 3–4 μm thick, but becoming gradually swollen, elongate–clavate, and to 9.5–10 μm wide at the tips where the cell walls can be 3–4 μm thick, the tips with dark bluegrey granules and then capped with a dark blue-grey mucilaginous sheath that can extend 5–6 μm above the tips. Asci broadly clavate, Lecanora-type, with a strongly thickened apex that can occupy to a third of the ascus length when mature, 40–47 x 14–25 μm, 8-spored. Ascospores irregularly overlapping in the asci, ellipsoid, rounded at the ends, simple, smooth-walled, without an evident sheath, 9.5–10.5 x 5.5–6.5 μm. No chemical reagent tests were carried out in view of the limited amount of material available on the neotype specimen. The distinctive blue-grey pigmented mucilage was evident in water mounts and in lactofuschsin. Our measurements and observations agree closely with those of Dodge except that he gave the ascospores as “10–12 x 4 μ”, but this may be due to our finding few mature ascospores; he also described them as “fusiform”, but the apices are not at all pointed. Although the holotype could not be located, McCoy & Siple 331 is annotated as this species by Dodge, and is represented by a slide. Although collected prior to the description of the fungus, it was not mentioned in the original account and Dodge’s annotations on the packet and slide are undated; it is therefore selected as a neotype rather than a lectotype here. There are also two microscope slides labelled as “Phacopsis usneae?” from different localities on other Usnea species: on “Usnea aurantiaca-atra” from Refuge Island (Bryant 32) and “Usnea siplei” from George VI Sound (Eklund 860). In addition there are two collections not named to species: as “Phacopsis” with “U. fasciata” from Lambda Island (3 March 1941, G. Frazier, Bailey & P.A. Siple 383a-1), and as “Nesolechia” (a name regarded as a synonym of Phacopsis by Triebel et al. 1995) on “U. frigida” from Eights Coast (Jones Mountains, Lichen Point, 73º32'S 94º00'W, ? Jan. 1961, G.F. Webers 61-1-1B-B). This name was evidently overlooked in the monograph of the genus by Triebel et al. (1995), who described one species on Usnea as new, P. falcispora Triebel & Rambold 1995 on U. maculata from South Africa. That species is clearly distinct from P. usneae and was figured as having crescent-shaped ascospores, often attenuated at the apices, and measuring (11.5–)13.5–15(–16) x 5–5–5(–6) μm (var. falcispora) or (12–)14–16(–19) x 5–5.5(–6) μm (var. sipmanii). (9) Thelidiola eklundii C.W. Dodge, Nova Hedwigia 15: 300 (1968). Type: Antarctic Peninsula, Nunatak, 3 miles south-east of Mount Wilcox, north wall of McClary glacier, on “Catillaria eklundii” [? Buellia sp.], 68º00'S 67º47'W, IP address: 202.75.16.18 299 LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE C.Eklund B-6-A [56] pro parte (FH – holotype). = Muellerella pygmaea (Körb.) D. Hawksw., Bot. Notiser 132: 289 (1979). This is the type species of Thelidiola, a new generic name introduced by Dodge (1968, p. 299) for this species and Thelidium caloplacae (see below). Two packets were located in FH under this name, one with slightly fuller locality information, but both with the collector´s number “B-6-A”. However, only one has the US Antarctic Service Expedition number 56, which was cited by Dodge and so that is considered the holotype here. We suspect the second collection was split from the first, perhaps as both also have the species Catillaria eklundii Dodge 1968, described in the same publication, and that this was done to enable a part to be filed in the herbarium separately under the Catillaria name. Castello & Nimis (1995, p. 89) give the correct “B-6A” reference in the entry under C. eklundii, which they considered “by an educated guess, could be attributed to a Buellia”, but under Thelidiola eklundii refer again to two packets having both taxa, but give the reference as “Siple et al. DW-4”; this number actually relates to the holotype of T. caloplacae and not of T. eklundii; we therefore interpret this as a simple lapsus for “B-6-A” which contains a Muellerella, the ascospores of which “suggest M. lichenicola”. However, the holotype specimen also has an annotation label by Vagn Alstrup dated 23 March 1995 identifying the specimen as M. pygmaea. The perithecia arise almost superficially and are rather large, reaching 260 μm diameter, and the ascospores are dark brown, broadly ovoid, 1-septate, and the size range we measured was 7–9(–10) x 3.5–4.5 μm. These observations and measurements essentially agree with those of Dodge, who stated that the perithecia were sessile, to 270 μm diam, and the ascospores 10 x 4 μm. All these features conform to M. pygmaea rather than M. lichenicola, more particularly M. pygmaea var. pygmaea as understood by Triebel (1989). In M. lichenicola the perithecia are generally immersed, 50–100 μm diameter, and the ascospores 5–7(–8) x 2–3 (–4) μm (Hawksworth 1979). (10) Thelidium caloplacae C.W. Dodge & G.E. Baker, Ann. Mo. Bot. Gdn 25: 525 (1938). Type: Marie Byrd Land, Edsel Ford Range, Mt Donald Woodward [as “Mt Rea-Cooper”; see below], on Xanthoria elegans [as “Gasparrinia siplei and Kuttlingeria rufa”], 1934, P. Siple, F.A. Wade, S. Corey & O.D. Stancliff DW-4 (FH – holotype). In addition to the specimen designated as type, Dodge & Baker (1938, p. 526) also cited a collection from Skua Gull Peak, F. Siple & S. Corey 72W-6. Specimens with the numbers DW-4 and 72W-6 are both present in FH and Castello & Nimis (1995, p. 88) considered these were “probably” Muellerella pygmaea. There is, however, some uncertainty over the typification as the locality name on packet DW-4 is given as “Mt Rea-Cooper”; this is presumably why Castello & Nimis state “type material not located” even though they had studied DW4. Interestingly, according to the route map published by Siple (1938, p. 475) Donald Woodward Mount is very close to but separated from Mount Rea and Mount Cooper (which are separate peaks in the same mountain structure) by the Ames Glacier. However, Siple (1938, p. 487) mentions this species as present on Mount Donald Woodward, but not on Mount Rea-Cooper (Siple 1938, p. 492–493). At the former site the lichens are reported as growing on a schist, while at the latter on a coarse-grained light pink granite. The specimen does not appear to be on a granite, but rather a homogenous schist, and the substrate indicated in the protologue is schist not granite; we therefore suggest that the locality name on the packet is an error of which Siple was aware and so he cited the species only from the one Mount Donald Woodward site. Dodge (1973, p. 14) also does not mention Mount Rea Cooper as a locality for this species. The species was originally described as “parasitic on the thalli of Gasparrinia siplei and Kuttlingeria rufa” in Dodge & Baker (1938), while the packets “DW-4” and “72W-6” in FH are labelled “on sterile Blasteniaceae”. The host in DW4 is actually a fragment of Xanthoria elegans, of which the name K. rufa is a synonym according to Castello & Nimis (1995). The last authors indicate that Gasparrinia siplei (as K. siplei) is Caloplaca lucens but we did not note this on the material we studied. Santesson (1960, p. 508) was the first to suggest the species was probably conspecific with M. lichenicola, as noted by Hawksworth (in Pegler et al. 1980, p. 522). The perithecia were immersed, erumpent only in the uppermost part, and the largest we measured was 100 μm diameter and the ascospores in DW-4 were 6–7.5 x 2–3(–4.5) μm, all within the reported range for M. lichenicola (e.g. Hawksworth 1979, Triebel 1989). (11) Thelidium minutum C.W. Dodge, Nova Hedwigia 15: 301 (1968). ≡ Thelidiola caloplacae (C.W. Dodge & G.E. Baker) Dodge, Nova Hedwigia 15: 299 (1968). Type: Victoria Land, Walcott Glacial Area, south-east of Lake Penny, on Rhizoplaca melanophthalma [as “Lecanora lavae”], 22 February 1964, K.P. Rennell [E. Schofield AA135] (FH –holotype). = Muellerella lichenicola (Sommerf.) D. Hawksw., Bot. Notiser 132: 289 (1979). = Muellerella pygmaea (Körb.) D. Hawksw., Bot. Notiser 132: 289 (1979). http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 202.75.16.18 300 D.L. HAWKSWORTH & T. ITURRIAGA According to Castello & Nimis (1995, p. 89) this was a Muellerella, “perhaps M. pygmaea (Körb.) D. Hawksw.” However, a later annotation slip by Vagn Alstrup (23 March 1995) identifies this is Muellerella lichenicola growing on Rhizoplaca melanophthalma. The perithecia measure 120–250 μm diameter and the ascospores are broadly ellipsoid, dark brown and 2-celled when mature, and measure 6–8(–9) x 4.5 μm, which matches well with the concept of M. pygmaea s. str. (Hawksworth 1979, Triebel 1989). The thallus consists of bullate areoles and is clearly that of a Rhizoplaca and not at all like Lecanora flotowiana which has no evident superficial thallus and of which the name L. leave is a synonym (Øvstedal & Smith 2001, p. 206). (12) Thelidium parvum C.W. Dodge & G.E. Baker, Ann. Mo. Bot. Gdn 25: 526 (1938). Type: Marie Byrd Land, Edsal Ford Range, Skua Gull Peak, growing over “Parmelia variolosa” [? Physcia caesia], P. Siple & S. Corey 72W-4 (holotype not located). = Stigmidium pumilum (Lett.) Matzer & Hafellner, Bibltheca Lich. 37: 115 (1990). The biological status of this species is obscure as the fungus was described as forming hyphae which surrounded “miscellaneous algae principally Scytonema and Pleurococcus” with the whole “growing over the thallus of Parmelia variolosa”. The host lichen named as P. variolosa is Physcia caesia (see under Diplonaevia parmeliae above). No specimen of Thelidum parvum was located by Castello & Nimis (1995, p. 73), and nor did we find one, but in this case an illustration was provided by Dodge & Baker (1938, pl. 62, figs 393–395). This shows a section of a perithecium with a pseudoparenchymatous cell wall, some suggestion of interascal tissue; maturing asci (probably the contents without the outer wall as a dimple appears at the apex), two broadly soleiform ascospores, and hyphae ensheathing the two kinds of algae. The perithecia were described as to 50 x 55 μm, with no paraphyses but “gelatinous filmy strands among the asci”; the asci as elongate and somewhat clavate, 28–31 x 10–12 μm, and 8-spored; and the ascospores as 1-septate, hyaline, not constricted at the septum, with one cell larger and blunt at the end and the other smaller and more pointed, and 7.5–9 x 3–3.5 μm. The maximum perithecium size given is much too small for the lichen-forming genus Thelidium, the size range for perithecia in the genus being 0.08–6.5 mm (Purvis et al. 1992). Critical to any placement is the nature of any interascal tissues, but it seems unlikely that “gelatinous filmy strands” were really some form of paraphyses as such structures were carefully documented in many other species treated in the same work. It seems most probable that Dodge & Baker were dealing with a lichenicolous species of Stigmidium, although they do not state if the perithecia were http://journals.cambridge.org Downloaded: 11 Dec 2013 immersed or superficial, and perhaps some other hyphal fungi and cyanobacteria/algae. Indeed, the dimensions of the ascomata and asci are in the ranges reported in S. pumilum by Matzer & Hafellner (1990). Stigmidium pumilum is confined to Physcia caesia and the closely allied P. dubia and known from Chile and New Zealand as well as the Canary Islands and Europe (Matzer & Hafellner 1990, Wedin 1994). However, the ascospores in S. pumilum measure (8–)9–13(–16) x 3–7 μm, embracing those reported for T. parvum, but also becoming longer and broader. Bearing in mind the coincidences as to dimensions and possible host, however, we suggest that the species belongs here and that Dodge & Baker simply measured rather few and still maturing ascospores. Acknowledgements We are indebted to Donald H. Pfister, Scott LaGreca, and especially Genevieve Lewis-Gentry, for diligently and repeatedly searching for specimens and slides in the Dodge collections at the Farlow Herbarium and Reference Library (Harvard University) for us to study and to be sure elusive collections were really no longer present. We are also indebted to Josef Hafellner for his perceptive and helpful comments on our original submission. This work was undertaken while D.L.H. was supported by the Programa Ramón y Cajal of the Ministry of Science and Technology of Spain, and T.I. was on sabbatical in the Departamento de Biología Vegetal II of the Facultad be Farmacia in the Universidad Complutense de Madrid funded by the Ministry of Education and Culture of Spain. References APTROOT, A., DIEDERICH, P., SÉRUSIAUX, E. & SIPMAN, H.J.M. 1997. Lichens and lichenicolous fungi from New Guinea. Bibliotheca Lichenologica, 64, 1–220. BRODO, I.M. & HAWKSWORTH, D.L. 1977. Alectoria and allied genera in North America. Opera Botanica, 42, 1–164. CALATAYUD, V. & NAVARRO-ROSINÉS, P. 1998. Weddellomyces xanthoparmeliae sp. nov. and additions to the chorology of other species of the genus. Mycotaxon, 69, 503–514. CASTELLO, M. & NIMIS, P.L. 1995. A critical revision of Antarctic lichens described by C. W. Dodge. Bibliotheca Lichenologica, 57, 71–92. DODGE, C.W. 1936. Medical mycology. London: Henry Kimpton, 900 pp. DODGE, C.W. 1948. Lichens and lichen parasites. British and New Zealand Antarctic Research Expedition 1929–1931, Reports, Series B (Zoology and Botany), 7, 1–276. DODGE, C.W. 1968. Lichenological notes on the flora of the Antarctic continent and the sub-Antarctic islands. VII and VIII. Nova Hedwigia, 15, 285–332. DODGE, C.W. 1973. Lichen flora of the Antarctic Continent and adjacent islands. Cannan, NH: Phoenix Publishing, 399 pp. DODGE, C.W. & BAKER, G.E. 1938. [Second Byrd Antarctic Expedition – Botany] II. Lichens and lichen parasites. Annals of the Missouri Botanical Garden, 25, 515–727. DODGE, C.W. & ZELLER, S.M. 1934. Hymenogaster and related genera. Annals of the Missouri Botanical Garden, 21, 625–705. IP address: 202.75.16.18 301 LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE GRUBE, M. & HAFELLNER, J. 1990. Studien an flechtenbewohnenden Pilzen der Sammelgattung Didymella (Ascomycetes, Dothideales). Nova Hedwigia, 51, 283–360. HALE, M.E. 1987. Epilithic lichens in the Beacon sandstone formation, Victoria Land, Antarctica. Lichenologist, 19, 269–287. HAWKSWORTH, D.L. 1979. Studies in the genus Endococcus (Ascomycotina, Dothideales). Botaniska Notiser, 132, 283–290. HAWKSWORTH, D.L. 1986. Notes on British lichenicolous fungi: V. Notes from the Royal Botanic Garden Edinburgh, 43, 497–519. HAWKSWORTH, D.L. 2003. The lichenicolous fungi of Great Britain and Ireland: an overview and annotated checklist. Lichenologist, 35, 191–232. HAWKSWORTH, D.L. 2004. Fungi living on lichens: a source of unexplored diversity. British Wildlife, 15, 192–199. HAWKSWORTH, D.L. & COLE, M.S. 2002. Intralichen, a new genus for lichenicolous “Bispora” and “Trimmatostroma” species. Fungal Diversity, 11, 87–97. LAMB, I.M. 1964. Antarctic lichens I. The genera Usnea, Ramalina, Himantormia, Alectoria, Cornicularia. British Antarctic Survey Scientific Reports, No. 38, 1–34. LAMB, I.M. 1968. Antarctic lichens II. The genera Buellia and Rinodina. British Antarctic Survey Scientific Reports, No. 61, 1–129. LAWREY, J.D. & DIEDERICH, P. 2003. Lichenicolous fungi: interactions, evolution, and biodiversity. Bryologist, 106, 80–120. MATZER, M. 1993. Beitrag zur Kenntnis der Ascomycetengattungen Globosphaeria, Roselliniopsis und Synaptospora. Cryptogamie, Mycologie, 14, 11–19. MATZER, M. & HAFELLNER, J. 1990. Eine Revision der lichenicolen Arten der Sammelgattung Rosellinia (Ascomycetes). Bibliotheca Lichenologica, 37, 1–138. http://journals.cambridge.org Downloaded: 11 Dec 2013 MURRAY, J. 1963. Lichens from Cape Hallett area, Antarctica. Transactions of the Royal Society of New Zealand, 2(5), 59–72. NAVARRO-ROSINÉS, P. & ROUX, C. 1995. Le genre Weddellomyces (Dothideales, Dacampiaceae) en Catalogne et en Provence. Mycotaxon, 53, 161–187. NAVARRO-ROSINÉS, P. & ROUX, C. 1997. Weddellomyces protearius sp. nov. et Lichenochora xanthoriae, champignons lichénicoles non lichénisés parasites de Caloplaca proteus. Mycotaxon, 61, 433–440. ØVSTEDAL, D.O. & SMITH, R.I.L. 2001. Lichens of Antarctica and South Georgia: a guide to their identification and ecology. Cambridge: Cambridge University Press, 411 pp. PEGLER, D.N., SPOONER, B.M. & SMITH, R.I.L. 1980. Higher fungi of Antarctica, the subantarctic zone and Falkland Islands. Kew Bulletin, 35, 499–562. PURVIS, O.W., COPPINS, B.J., HAWKSWORTH, D.L., JAMES, P.W. & MOORE, D.M. 1992. The lichen flora of Great Britain and Ireland. London: Natural History Museum Publications, 710 pp. RUDOLPH, E.D. 1990. Carroll William Dodge, 1895–1988. Mycologia, 82, 160–164. SANTESSON, R. 1960. Lichenicolous fungi from northern Spain. Svensk Botaniska Tidskrift, 39, 499–522. SIPLE, P.A. 1938. The second Byrd Antarctic expedition – Botany I. Ecology and geographical distribution. Annals of the Missouri Botanical Garden, 25, 467–514. TRIEBEL, D. 1989. Lecideicole Ascomyceten: Eine Revision der obligat lichenicolen Ascomyceten auf lecideoiden Flechten. Bibliotheca Lichenologica, 35, 1–278. TRIEBEL, D., RAMBOLD, G. & ELIX, J.A. 1995. A conspectus of the genus Phacopsis (Lecanorales). Bryologist, 98, 71–83. WEDIN, M. 1994. New and noteworthy lichenicolous fungi from southernmost South America. Lichenologist, 26, 301–310. IP address: 202.75.16.18 http://journals.cambridge.org Downloaded: 11 Dec 2013 IP address: 202.75.16.18