J. Hattori Bot. Lab. No. 74: 287-298 (Nov. 1993)

THE GENERA AND PANNOPARMELIA IN

2 ISAO YOSHIMURA1 and JOHN A. Eux

ABSTRACT. Two new of Anzia, A. minor Yoshim. and A. tianjarana Yoshim. et Elix, and two species of Pannoparmelia, P.angustata (Pers. in Gaud.) Zahlbr. and P. wilsonii (Riis.) D. Gall. are recorded from Australia. A key to the species is given and their chemistry, distribution, ecology and described.

INTRODUCTION In a continuation of a worldwide study (by I. Y.) of the genera Anzia and Pannoparmelia (Yoshimura 1987) we undertook a review of the Australian represen­ tatives of these genera. We now report the discovery of two new species of Anzia from the continent, in addition to the well known Pannoparmelia angustata and P. wilsonii (Bratt, Blackman, and Cashin 1976; Galloway 1978).

MATERIAL Material preserved in several herbaria (TNS, US, BM, ANUC, CANB, NICH) including recent collections (mostly by J. A. E.) of the genera Anzia and Pannopar­ me/ia were used for the present study.

M ETHODS Cross- and longitudinal-sections of the material were obtained by means of a freezemicrotome (Reichert-Jung; Frigomobile OM and Freezing-microtome 1206). Sections (20-30µm thick) of thalli were stained with lactophenol cotton-blue. Asci were stained with Iodine solution. Sections were observed using a light-microscope and photographed. Methods for the thin-layer chromatography (TLC) of lichen substances essentially followed those of C. Culberson (1972) and Elix et al. (1987, 1988). Fragments of dried lichen specimens were extracted with acetone. Extracts were spotted on to Merck precoated glass plates (Kiesel Gel 60, no. 5721; 100 X200mm) and chromatograms were developed for 20min. at 25 °C in various solvent systems; i.e. benzene, dioxane, acetic acid (90: 25 : 4, v/v/v); hexane, ether, formic acid (5 : 4 : 1, v/v/v); toluene, acetic acid (85: 15, v/v). High-performance liquid chromatography (HPLC) was carried out with a JASCO IC-800 liquid chromatograph at 25 °C. Conditions were as follows: column, JASCO

1 Kochi Gakuen College, 292 Asahitenjinchi, Kochi, 780, Japan. 2 Department of Chemistry, The Faculties, Australian National University, , ACT 0200, Australia. 288 J. Hattori Bot. Lab. No. 74 I 9 9 3

Finepack SIL C18- 10P 4.6mm X 150mm; solvent system, methanol-water-phosphoric acid (80: 20: 0.9, v/v/v); detector, JASCO UV spectrophotometric detector (250 nm), and Shimazu photodiodearray spectrophotometer (range; 200-400nm, minimum width; 2 nm). Identification of lichen substances by TLC and HPLC was made by comparison with the standard lichen substances isolated by Dr. S. Huneck (protocetraric acid, hypoprotocetraric acid, chloroatranorin, atranorin, usnic acid, divaricatic acid, sekika­ ic acid, and lobaric acid), and Dr. K. Takahashi (sekikaic acid and divaricatic acid). Samples of several species of Anzia were used for comparisons with the substance indicated in parenthesis: A. japonica (anziaic acid), A. gregoriana (anziaic acid), and A. opuntiella ( divaricatic acid). Authentic 4-0-methylhypoprotocetraric acid was isolated from Xanthoparmelia notata (Kurok.) Hale (Cresp, Elix et al. 1972); anziaic acid, subnotatic acid, and nordivaricatic acid were identified by comparison with synthetic material (Djura & Sargent 1977; Elix 1974; Elix & Tearne 1977; Elix & Lajide 1984).

RESULTS AND DISCUSSIONS Chemical Substances A diagram of some of the TLC chromatograms is presented in Figure 1, and the UV spectra of six substances are presented in Figures 2- 8. Using HPLC the fo ll owing three substances can be clearly distinguished from one another and identified by their UV spectra and their retention times: protocetraric acid (Fig. 2), hypoprotocetraric acid (Fig. 3), and 4-0-methylhypoprotocetraric acid (Fig. 4). Atranorin (Fig. 5) can be clearly distinguished from chloroatranorin (Fig. 6), although it cannot be separated in currently used solvents (TA = toluene: acetic acid, 85: 15, v/v; HEF = hexane, ether, formic acid, 5: 4 : 1, v/v/v). An unidentified substance (AT2) is the main chemical substance present in one of the chemical races of Anzia tianjarana. A T2 can be distinguished from anziaic acid ( C + red, UV spectrum: Fig. 8) by its color reaction (C-, KC + red), UV spectrum (Fig. 7), the retention time (Rt. 15 .59), and TLC. The main substance (Rt. 12.58; A max 215, 271, 302; A min 244, 298) present in Pannoparme/ia angustata and P. wilsonii was identified as divaricatic acid, while.nordivaricatic acid (Rt. 10.10; A max 215, 271, 304; Amin 242, 294) was identified by TLC and HPLC comparison with authentic material. Although all specimens of P. angustata and P. wilsonii were characteristicall y yellow­ green to dull green in colour, occasionally the usnic acid concentration was so low that it could not be detected chromatographically.

THE SPECIES Anzia minor Yoshim., sp. nov. (Figures 9- 12) Thallus laciniatus, laciniis plus minus dichotome divisis, strato medullari axem chondroideum praedito, subtus spongiostrato moniliforme constricto. Species cum thallo ut inAnziajamesii sed ad hac specie thallo saxicolo, parviore, lobis angustioribus, exisidiatis, et acido 4-0-methylhypoprotocetrarico vel acido protocetrarico continente differt. I. YOSHIMURA & J. A. Eux: The lichen genera Anzia and Pannoparmelia in Australia 289

TA(85: J5) Rf.

0.9

0.8 atranorin + chloroatranorin

0.7

0.6

0.5 4-0-methylhypvprotocetraric acid •. ::.:: :.:;, isonotatic acid 0.4 - subnotatic acid 0.3 hypoprotocetraric acid CJ CU> ® 0.2 ., "' . . 0.1 4-0-demethylnotatic acid 8 ~ protocetraric acid

0.0 "O <..> V") «: 0 "O <..> ~ '() .c: 00'° ..... «: «: c '°.._ V")°' ("'\ .... .c: 0 °''° <..> v 0 00 .c: <..> c .._ .._ °' t'd °' .._ .... c .... ·~ <:::> 0 c s: ~ .... "':1l . i: .§ .:::; .~ v C:l .s 0. 0 (') -~ ::::: ::: ::: ::: .9 c .... c 0 0. 0 :1l .... ;>., .... --.:: ~ ~ :a "' --.:: 0. ..c:: <..> -;; CD Fig. 1. TLC chromatograms of Anzia minor and comparisons with some authentic substances and some other species of Anzia. Solvents: TA = toluene, acetic acid (85 : 15, v/v).

Type. Australia. . Sandstone escarpment on the road 8 km east of Nerriga, elevation ea. 750m, Dec. 1, 1965, Syo Kurokawa 6405 (TNS-holotype, ANUC, NICH, US­ isotypes). Thall us foliose, saxicolous, loosely attached, orbicular to spreading, 2-6 cm wide. Lobes sublinear, more or less dichotomously branched, 0.4--0.5 (--0.9) mm wide, margins entire, apices rounded. Upper surface pale grey to bluish grey, becoming brown in the herbarium, smooth, matt or shining, occasionally distinctly maculate, often with tangential cracks in older lobes, lacking soredia and isidia. Medulla white. Lower surface white, shining, smooth, occasionally visible between the spongiostratum cushions and at the lobe tips. Spongiostratum moniliform, forming globose to ellipsoid 290 J. Hattori Bot. Lab. No. 74 I 9 9 3

750 protocetraric acid hypoprotocetraric acid 4-0-mdbylhypoprotocetraric acid m

"'80 •oo ff: 4. 91 (• u ) lfl ·l. 46( .. n) 2 3 tT : ll.!6(1in) 4 636 2ZI "' atranorin "' cbloroatranorin unidentified substance (A T2) anziaic acid

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tT : IS. St<1in) 7 Figs. 2- 8. UV spectra of 7 chemical substances found in Anzia minor and A. tianjarana. Fig. 2. Protocetraric acid. Fig. 3. Hypoprotocetraric acid. Fig. 4. 4- 0-Methylhypoprotocetraric acid. Fig. 5. Atranorin. Fig. 6. Chloroatranorin. Fig. 7. Unidentified substance (AT2). Fig. 8. Anziaic acid. cushions, up to 1.0 mm thick, brown-black to black. Rhizines scattered, simple, up to 1.2 mm long, with a tuft at the apex, born singly at the borders of the spongiostratum cushions. Upper cortex ea. 65 µm thick, arranged more or less vertical to the surface of the lobes (palisade); a chondroidal axis of parallel, longitudinally arranged hyphae present in the middle of the lobes, axis 220 X 350µm, originating from the apical meristematic portion of the lobes; medulla up to 50µm thick, composed of loosely interwoven hyphae, 7-IOµm thick; hyphae of spongiostratum derived from the apical meristematic portion of the lobes, 12- 17 µm wide; hyphae of spongiostratum pale brown to black-brown, composed of one row of lumina; rhizines derived from the apical meristematic portion of the lobes, connected to the chondoidal band. Apothecia common, 1.5- 2.0mm diam., pedicellate, weakly convex to more or less flat; disc dark brown to black; thalline exciple concolorous with the thallus and smooth on the upper part; cortex prosoplectenchymatous, ea. 200µm thick; lumina of cortical cells lµm wide, often branched to form net-like spongiostratum; hymenium ea. lOOµm high; paraphyses branched to form nets, septate; apices swollen; asci 55 X 20µm, multi­ spored; spores curved, worm-shaped, 12 X 2- 3 µm. Pycnidia globose, immersed in the I. YosJ-llM URA & J. A. Eux: The lichen genera Anzia and Pannoparmelia in Australia 291

Figs. 9- 12. Anzia minor Yoshim. Fig: 9. Thallus, dorsal view. Fig. 10. Thallus, ventral view. Fig. 11. Longitudinal section of the apical portion of a lobe. Fig. 12. Cross-section of a lobe. Figs. 9, 10. Scale = mm. Figs. 11, 12. Scale bar= 10 µm. lobes; ostiole black-brown, punctiform; conidia cylindrical, 4-5 X 0.5 µm. Chemistry. Chemical race 1: Cortex K + yellow; medulla K - , C - , KC - , P - . Containing atranorin, chloroatranorin, 4-0-methylhypoprotocetraric acid (major) and traces of isonotatic acid, subnotatic acid, hypoprotocetraric acid, 4-0-demethylnotatic acid (by TLC and HPLC). Chemical race 2: Cortex K + yellow; medulla K - , C - , KC - , P + orange-red. Containing atranorin, chloroatranorin, protocetraric acid (major). Specimens Examined: Chemical race 1. AUSTRALLA. New South Wales. On sandstone rocks in open Eucalyptus woodland, , 8 km north-east of Nerriga, 35 °07 'S, 150°08 'E, 750 m, J. A. Elix 5081, 31. X. 1978, J. A. Elix 9159, 18. X. 1981 , J. A. Elix 10171, I. V. 1982, J. A. Elix 11386 and J. Johnston, 6. X. 1983 (ANUC). Chemical race 2. AusTRALLA. New South Wales. On sandstone rocks in open Eucalyptus woodland, Teudts Hill, 5 km south-west of Bundanoon, 680m, J. A. Elix 8969, 17. IX. 1980 292 J. Hattori Bot. Lab. No. 74 I 9 9 3

(ANUC). This new species is distinguished by the small, narrow lobes (less than 1 mm and usually less than 0.5mm wide), the presence of the lower cortex visible between the spongiostratum cushions and the presence of the characteristic medullary depsidones, 4 -0 -methylhypoprotocetraric acid or protocetraric acid. The presence of two different chemical races in the populations of A. minor is noteworthy and may yet indicate that two distinct species are involved. However, the major depsidones present in the two races are biogenetically related, and in the absence of a wide range of specimens, the observed morphological variation of the two chemotypes is not sufficient to warrant specific taxonomic recognition at this time. The type specimen belongs to chemical race I. To date A. minor has only been found in hinterland areas of southern New South Wales growing on outcrops of Hawkesbury sandstone. It occurs on relatively sheltered rock crevices in open Eucalyptus woodland, often growing together with Cladia aggregata (Sw.) Nyl., C. infiata (F. Wils.) D. Gall., C. sullivanii (Milll. Arg.) Martin, Parmelia signifera Nyl. and some Frullania sp.

Anzia tianjarana Yoshim. et Elix, sp. nov. (Figures 13- 16) Thallus laciniatus, laciniis irregulariter divisis, strata medullari axem chondroide­ um praedito, subtus spongiostrato perpetuo formato. Species cum thallo ut in Anzia gregoriana sed ad hac specie thallo parviore, lobis angustioribus et spongiostratus solidibus differt. Type. Australia. New South Wales. On sandstone rocks in open Eucalyptus woodland, Morton National Park, above the Tianjara Falls, 33 km north-north-west of Ulladulla, 35 °05 'S, 150°19 ' E, 510m, J. A. Elix 384b, 13. IX. 1974 (CBG-holotype). Thall us foliose, saxicolous, loosely attached, orbicular, 2- 4 cm wide. Lobes linear, subdichotomously branched, 0.4--0.8 (- 1.0) mm wide, l.0-2.5mm between branches. Upper surface pale grey-white to olive grey, becoming brown in the herbarium, smooth, matt or shining, margins without a white rim, black spongiostratum often projecting beyond the margins, often with tangential cracks in older lobes, lacking soredia and isidia. Medulla white. A chondroidal axis of parallel, longitudinally arranged hyphae present in the middle or the lower medullary area of the lobes, originating from the apical meristematic portion of the lobes. Lower surface covered by continuous spongi­ ostratum; spongiostratum up to 1.0 mm thick, brown-black to black, solid (not hollow), uniformly attached to lower medullary hyphae or chondroidal axis. Rhizines scattered, simple, black, up to 1.2 mm long, connected to the chondoidal axis. Apothecia not seen. Pycnidia saucer-shaped, immersed in the thallus, ostiole black-brown, punctiform; conidia rod-shaped, 5- 6 X 0.5 µm. Chemistry. Chemical race I. Cortex K + weak yellow or K -; medulla K - , C + rose, KC + red, P - . Containing atranorin, chloroatranorin, and anziaic acid. Chemical race 2. Cortex K + weak yellow or K - ; medulla K - , KC + rose, P - . Containing atranorin, chloroatranorin, and A T2 (unidentified substance by TLC and HPLC). Specimens examined of the chemical race 2. AUSTRALIA. Same locality as the I. YOSHIMU RA & J. A. Eux: The lichen genera Anzia and Pannoparmelia in Australia 293

Figs. 13- 16. Anzia tianjarana Yoshim. & Elix Fig. 13. Thallus, dorsal view. Fig. 14. Thallus, ventral view. Fig. 15. Longitudinal section of the apical portion of a lobe. Fig. 16. Cross-section of a lobe. Figs. 13, 14. Scale = mm. Figs. 15, 16. Scale = IOµm. type specimen. J. A. Elix 384a, 13. IX. 1974, CBG. The diagnostic characters of the species include: the thallus composed of the irregularly and repeatedly branched and narrow laciniate (less than 1 mm wide), the presence of the chondroidal band in the thallus (Figs. 7, 8); the continuous spongio­ stratum covering the lower surface (Fig. 14); and the presence of AT2 (an unidentified substance, KC + red) or anziaic acid. The new species has a chondroidal band in the thallus and it belongs to section Nervosae. The thallus of A. tianjarana is composed of irregularly branched laciniae, while the other members of section Nervosae have typical dichotomously branched laciniae. The thallus size of A. tianjarana is as small as that of A. minor; however, A. tianjarana can be distinguished from A. minor by its continuous spongiostratum on lower surface of the laciniate thallus (Fig. 14), while the spongiostratum of A. minor is intermixed with rhizines (Fig. 10). Anzia tianjarana has two chemical races. One contains anziaic acid, while the other 294 J. Hattori Bot. Lab. No. 74 I 9 9 3 contains the unidentified substance (AT2). The type specimen belongs to chemical race I. The unidentified substance (A T2) may be related to anziaic acid but differs from anziaic acid, perlatolic acid ( 4-0-methylanziaic acid) and planaic acid. The UV spectrum of A T2 (Fig. 7) is closely resembles anziaic acid (Fig. 8) and perlatolic acid (A max 215, 271, 304; A min 244, 296, Rt. 40.67) but is quite different from that of planaic acid (A max 210, 279, A min 273; Rt: 12.42). Anzia tianjarana appears to be related to A. gregoriana Mull. Arg. from New Guinea, but the latter species differs in its larger thalli (6- 8 cm diam.), broader lobes (0. 7-1.5 mm wide) and more particularly by the hollow masses of spongiostratum (solid in A. tianjarana). A. tianjarana appears to be a very rare species and to date is known only from the type locality, a hinterland area of southern New South Wales, where it grows on outcrops of Hawkesbury sandstone. It occurs in open Eucalyptus woodland on relatively sheltered rock faces close to water, growing together with Cladia aggregata (Sw.) Nyl., C. inflata (F. Wils.) D. Gall., Hypogymnia subphysodes (Krempelh.) Filson, Parme/ia signifera Nyl. and Xanthoparmelia streimannii (Elix & Armstr.) Elix and Johnst.

Pannoparmelia angustata (Pers. in Gaud.) Zahlbr., Symb. Sin. 3 Lich: 195 ( 1930). Parmelia angustata Pers. in Gaud., Voy. Uranie Bot: 195 (1826). Type: Australia (Nova Hollandiae ), Port Jackson, ad arborem truncos, leg. Gaudichaud? (P-holotype, not seen; US-isotype!) . = Anzia angustata (Pers. in Gaud.) Mi.ill . Arg., Flora 72: 507 (1889). For further synonymy see Galloway 1978, p. 264. Thall us foliose, corticolous, orbicular or spreading, 6- 10 cm wide. Lobes linear or moniliform constricted, 0.5-2.5 mm wide, moderately to densely dichotomously to subdichotomously branched. Upper surface yellow-green to green or yellow-grey, matt, smooth, often slightly wrinkled in older parts, lacking soredia and isidia. Medulla white. Lower cortex white, smooth, shining, occasionally visible between the spongio­ stratum cushions and at the lobe apices. Spongiostratum moniliform, forming hemi­ spherical cushions, up to 1 mm thick, brown to black, often projecting beyond the margins of the lobes. Rhizines frequent, brown to black, simple, up to 1 mm long, often tufted at the apex, developing at the margins of the spongiostratum. Apothecia common, 0.8- 3.5(- 10) mm diam., lamina!, stipitate, cupuliform at first but expanding horizontally, becoming flattened and lacerate with age, disc light brown to brown; thalline exciple concolorous with the thallus, smooth at first but becoming wrinkled, warty, and developing dark spongiostatum around the base of older fruits; asci claviform, 8-spored, 30-34 X 14- 16µm; spores colourless, globose to broad ellipsoid, 6.0-6.5 X 5.0-6.0µm, with a thick epispore. Pycnidia common, solitary, lamina! or submarginal, globose to barrel shaped; ostiole black, punctiform; conidia cylindrical to slightly fusiform, 5.0-6.5 X lµm. Chemistry. Cortex K - ; medulla K - , C - , KC - , P - . Containing usnic acid (major), atranorin (minor), divaricatic acid (major) and nordivaricatic acid (trace) (by TLC and HPLC). I. YOSHIMURA & 1. A. Eux: The lichen genera Anzia and Pannoparmelia in Australia 295

Representative Specimens Examined. AUSTRALIA. New South Wales. On Callitris endlicheri, 15 km east of Cooma along the Numeralla road, 910m, J. A. Elix 1486, 19. 1. 1976 (ANUC); on Callitrisco/umellaris in Callitris woodland, Warranderry Range, Warranderry State Forest, 23 km north of Grenfell, 33 ° 4 I'S, 148° !2'E, 380m, J. A. Elix 17727 and H. Streimann, 17. X. 1984 (ANUC); on Ca/litris, Tinderry Mountains, east of Michelago, south of Canberra, on knoll along the main ridge leading from Tea Tree Creek to south summit of The Twins, 1470m, W. A. Weber and D. McVean s.n., (Lichenes Exsiccati, COLO, no. 281), 22. XII. 1967 (BM, CANB, NICH). Australian Capital Territory. On dead Pinus radiata, along Paddys River near Murrays Corner, SSOm, J. A. Elix 435, 17. IX. 1974 (ANUC); on Callitris endlicheri, I km north of Casuarina Sands, 550 m, J. A. Elix 925, 13. VI. 1975 (ANUC); on Callitris endlicheri, Molonglo Gorge Reserve, 14 km east of Canberra, 35 ° 16'S, 149 ° l 6'E, 650 m, J. A. Elix 11750 and J. Johnston (Lichenes Australasici Exsiccati no. SI), 31. V. 1984 (ANUC, BM, CBG, MEL, TNS). Victoria. Beechworth (BM, herb. Stirton). Tasmania. On Atherosperma moschatum in damp Atherosperma forest, Lake Dobson National Park, old Fonton Track, G. C. Bratt 471, 18. VIII. 1963 (BM); in badly burned forest, Lake St. Clair National Park, trail from Derwent Bridge to lakes below Mt. Hugel, W. A. Weber and D. McVean L-49890, 24. II. 1968 (NICH). Pannoparme/ia angustata is a widespread species known from south-eastern Aus­ tralia (New South Wales, Australian Capital Territory, Victoria, Tasmania), New Zealand and South America (Bratt, Blackman & Cashin 1976; Calvelo and Adler 1992; Galloway 1978). In Tasmania P. angustata is a common epiphyte on Nothofagus cunninghamii in the wetter, western regions, while in south-eastern Australia it is more common in the drier, open sclerophyll forests west of the , particularly on Callitris sp. It is a rather variable species showing differences in morphology which appear to correlate with changes in microclimate, and colour variations dependent on the degree of exposure to sunlight. Shade forms tend to be pale grayish-green rather than distinctly yellow-green, reflecting a lower concentration of usnic acid in the upper cortex. Common associates include Flavoparmelia rutidota (J. D . Hook. & Taylor) Hale, Hypogymnia billardieri (Mont.) R. Fils., Parmelia pseudot­ enuirima Gyeln. and Usnea scabrida Taylor.

Pannoparmelia wilsonii (Ras.) D. Gall., N. Z. J. Bot. 16: 267 (1978).(Figures 17- 20) Anzia wi/sonii Riis., Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo 20(3): 2 (1944). Type: Australia. New South Wales, Eastern Creek, ad lignum vestustum, F. R. M. Wilson, 1892 (H-holotype, BM-isotype). = Parmelia angustata var. isidiella Stirt., Trans. N. Z. Inst. 32: 81 (1900). Type. Australia, New South Wales, Tilba-Tilba, W. Tilden (BM ex herb. Stirton). For further synonymy see Galloway 1978, p. 267 Thallus foliose, corticolous or rarely saxicolous, orbicular or spreading, 3- 8 cm wide. Lobes convex, linear or moniliform constricted, 0.5- 1.0 (- 2.0) mm wide, subdichotomously branched, sometimes becoming densely imbricate towards the centre. Upper surface pale yellow to bright yellow-green, slightly shiny, smooth, Jacking soredia but densely isidiate in the older parts, isidia simple, papillose, then cylindrical, 0.1--0. 5 mm high, fragile, slightly expanded at the apices, becoming coralloid branched in the centre. Medulla white. Lower cortex yellow to pale brown, smooth, matt, occasionally visible between the spongiostratum cushions and at the lobe apices. 296 J. Hattori Bot. Lab. No. 74 I 9 9 3

Figs. 17- 20. Pannoparmelia wilsonii (Ras.) Gallow. Fig. 17. Thallus with apothecia, dorsal view (Elix, 1972). Fig. 18. Cross-section of the hymenium, showing fertile asci (stained by iodine solution). Fig. 19. Fabiform or curved spores. Fig. 20. Fabiform or curved spores in an ascus. Fig. 17. Scale = mm. Figs. 18- 20. Scale bar= IOµm.

Spongiostratum moniliform, forming hemispherical cushions, up to 0. 9 mm thick, brown to black, often projecting beyond the margins of the lobes. Rhizines frequent, brown to black but often pale at base, simple, up to 3.0 mm long, often tufted at the apex, developing at the margins of the spongiostratum. Apothecia rare, 0.8-3.0 mm diam., laminal, sessile to substipitate, cupuliform but becoming flattened and expanding horizontally with age, disc red-brown to brown; thalline exciple concolorous with the thallus, smooth at first but becoming isidiate in older fruits; asci claviform, 8-spored, 50-55 X l5- 20µm; spores colourless, fabiform, 10-15 X 4.0-6.0µm, with a thick epi­ spore. Pycnidia common, solitary, lamina] or submarginal, globose to barrel shaped; ostiole black, punctiform; conidia cylindrical to slightly fusiform, 5.0-6.5 X l µm. Chemistry. Cortex K - ; medulla K -, C -, KC - , P - . Containing usnic acid, atranorin ( ± ), divaricatic acid (major), nordivaricatic acid (trace), unknown ( ± trace) (by TLC and HPLC). Selected Specimens Examined. AUSTRALIA. Queensland. On dead Eucalyptus log in tropical rain forest, Great Dividing Range, Mt. Baldy, 4km south-west of Atherton, l7° 16'S, 145 °23 'E, 1060m, J. A. Elix 17141 and H. Streimann, 2. VII. 1984 (ANUC). New South Wales. On dead wood in dry sclerophyll forest, 50km east of Glen Innes along Highway 38, J. A. Elix 2441, 18. VIII. 1976 (ANUC); on Callitris columellaris in Callitris woodland, Kangarooby State Forest, 20 km south of Gooloog­ ong, 450m, J. A. Elix 8858, 10. IX. 1980 (ANUC); on dead wood in wet sclerophyll forest, Brown Mountain, 14km south-east of Nimmitabel, 36°36'S, 149 °36'E, 920m, J. A. Elix 9672 (Lichenes Australasici Exsiccati no. 11) 18. II. 1982 (ANUC, BM, CBG, MEL, TNS); on dead Eucalyptus logs in moist gully, Estuary, 7.5 km west of , 35 ° 42'S, 150°06'E, 5 m, J. A. Elix 10968, 29. V. 1983 (ANUC); Mt. Wilson, Blue Mountains, 1000 m, S. Kurokawa 5051, 26. X. 1965 (TNS, Yosh.); on bark of Eucalyptus robertsonii in wet sclerophyll forest, Braidwood district, SE facing escarpment of Budawang Range between I. YOSHIMURA & J. A. Eux: The lichen genera Anzia and Pannoparmelia in Australia 297

Majors Creek and Araluen, W. A. Weber and D. McVean s.n. (Lichenes Exsiccati, COLO, no. 311), 18. X. 1967 (BM, CANB). Australian Capital Territory. On Casuarina stricta in dry sclerophyll forest, along the Highway, 2.5 km south of Tharwa Road, 775 m, J. A. Elix 732, 11. IV. 1975 (ANUC). Victoria. On dead wood in dry sclerophyll forest, Strathbogie Range, 21 km east of Euroa along the Merton Road, J. A. Elix 2749, 6. XII. 1976 (ANUC); on dead wood in Eucalyptus forest, Mt. Drummer, 18 km east of Cann River, J. A. Elix 6233, 23. VII. 1979 (ANUC); on dead wood in wet sclerophyll forest, Toolangi State Forest, 24 km south of Yea, J. A. Elix 6233 , 23 . VII. 1979 (ANUC); on sandstone rocks in dry sclerophyll forest, Three Sisters, Three Sisters Track, 23 km north-north-east of Cann River, 37 °23 'S, 149 °06'E, 920 m, J. A. Elix 19547 and H. Streimann, 27. IX. 1985 (ANUC). Tasmania. On exposed granite rocks, Mt. Amos, Freycinet National Park, 42 °09'S, 145 ° 17 'E, 425m, J. A. Elix 5517, 13. I. 1979 (ANUC); on dead wood, Mt. Amos, eastern Tasmania, 42 °09'S, 145 ° 17 'E, 300m, G. C. Bratt & J. A. Cashin 68/1234, 19. IX. 1968 (HO, Yosh.). South Australia. On charred wood in dry sclerophyll forest, Mount Lofty Ranges, 4 km west of Carey Gully, 455 m, J. A. Elix 2824, 21. XII. 1976 (ANUC). Western Australia. On Nuytsia fioribunda in open Eucalyptus woodland, Darling Ranges, Boulder Rock Forest Reserve, 35 km south-east of Perth, J. A. Elix 10524 and L. H. Elix, 21. X. 1982 (ANUC); on Eucalyptus sp. in open forest, Tramway Trail, IQ km north of Pemberton, J. A. Elix 10724 and L. H. El ix, 25 . X. 1982 ( ANUC); on Banksia in mixed scrub, 16km south of Lake Clifton, J. A. Elix 10790 and L. H. Elix, 28 . X. 1982 (ANUC); on dead Eucalyptus in moist forest, Porongurups National Park, 40km north of Albany, along road from Karri Chalet to entrance of Park, A. Henssen 27002 (MB). Pannoparmelia wilsonii is a common species known from southern and eastern Australia (Queensland, New South Wales, Australian Capital Territory, Victoria, Tasmania, South Australia and Western Australia) and New Zealand (Galloway 1978). In Australia P. wilsonii is most common on dead or burnt wood in Eucalyptus woodland, but is also a common epiphyte on a variety of trees including Banksia, Callitris, Casuarina, Eucalyptus and Nuytsia. It occurs rarely on rocks in such forests. Pannoparmelia wilsonii is in fact much more common and widely distributed in Australia than its non-isidiate counterpart, P. angustata. Common associates include Hypogymnia pulverata (Ny!.) Elix, Menegazzia platytrema (Mill!. Arg.) R . Sant., Parmelia tenuirima Ny!., Parmelina conlabrosa (Hale) Elix and Johnst., and P. pseudorelicina (Jatta) Kantvilas and Elix. Most specimens of Pannoparmelia wilsonii lack apothecia; however, some Australi­ an populations rarely have fertile apothecia. According to Galloway ( 1978) spores have not been observed in New Zealand specimens. Many fertile apothecia (Fig. 17) were observed in two specimens (Elix 1972 and Elix 732). Each ascus contains 8 spores (Figs. 18, 20) the same as in P. angustata. As already noted in the original description by Rasanen (1944) the spores of P. wilsonii are fabiform in shape (Fig. 19), although P. angustata has ellipsoid spores. The fabiform spores of P. wilsonii are much broader than the crescent or worm-shaped spores of Anzia species. The general habit of P. wilsonii is similar to that of Anzia japonica (Sect. Anzia) and A. minor (Sect. Nervosae) and the shape of the spores of P. wilsonii seems to be intermediate between that of Pannoparmelia and Anzia. 298 J. Hattori Bot. Lab. No. 74 l 9 9 3

Key to Species of Anzia and Pannoparmelia in Australia I. Upper surface grey (atranorin present), chondroidal axis present m the medulla, asci multispored ...... 2 2. Spongiostratum continuous, medulla KC + rose (unidentified substance) or C + rose (anziaic acid) ...... Anzia tianjarana 2. Spongiostratum discontinuous, medulla KC- (protocetraric or 4-0-methylhypoproto- cetraric acids) ...... Anzia minor I. Upper surface yellow-green (usnic acid present, sometimes only in traces), chondroidal axis absent in the medulla, asci 8-spored ...... 3 3. Upper surface isidiate ...... Pannoparme/ia wilsonii 3. Upper surface non-isidiate ...... Pannoparme/ia angustata

ACKNOWLEDGEMENTS We thank the Australian Research Council for generous financial support of this project and the Ministry of Education, Science and Culture of Japan (Grant-in-Aid for Scientific Research, no. 364244); Dr. G. A. Jenkins (HPLC), Ms. Jen Johnston (TLC), and Mrs. T. Kurokawa (TLC & HPLC) who assisted in the determination of the chemistry of specimens. Thanks are extended to Dr. S. Huneck and Dr. K. Takahashi for gifts of authentic chemical substances, and curators of the several herbaria (BM, US, TNS, CANB, H) for the loan of the material.

LITERATURE CITED Bratt, G . C., A. J. Blackman and J. A. Cashin. (1976). The Anzia in Tasmania. Lichenologist 8: 69- 77. Calvelo, S. and M. Adler. (1992). Pannoparme/ia anzioides: a taxonomic synonym of Panno­ parmelia angustata (, Lichenes). Mycotaxon 43: 487- 498. Cresp, T . M., J . A. Elix, S. Kurokawa and M. V. Sargent. (1972). The structure of two new depsides from the lichen Parmelia notata. Australian Journal of Chemistry 25 : 2167- 2184. Culberson, C. F. (1972). Improved conditions and new data for the identification of lichen products by standardized thin-layer chromatographic method. Journal of Chromatography 72 : 113- 125. Djura, P. and M. V. Sargent. (1977). Depside synthesis IX. Nornotatic acid and notatic acid. Australian Journal of Chemistry 30: 1293- 1304. Elix, J. A. ( 1974). Synthesis of para-olivetol depsides. Australian Journal of Chemistry 27 : 1767- 1779. Elix, J. A., J. Johnston and J. L. Parker. ( 1988). A computer program for the rapid identification of lichen substances. Mycotaxon. 31 : 89-99. Elix, J. A. , J. Johnston and J. L. Parker. (1987). A catalogue of standardized thin layer chromatographic data and biosynthetic relationships for lichen substances. Canberra. Elix, J. A. and P. D. Tearne. ( 1977). Nordivaricatic acid, and new depside from the lichen Heterodea beaug/eholei. Australian Journal of Chemistry 30: 2333- 2335. Galloway, D. J. (1978). Anzia and Pannoparme/ia (Lichenes) in New Zealand. New Zealand Journal of Botany 16: 261- 270. Rasanen, V. (1944). Lichenes novi I. Ann. Bot. Soc. Zoo!. Bot. Fenn. "Vanamo" 20(3): 1- 34. Yoshimura, I. (1987). Taxonomy and speciation of Anzia and Pannoparmelia. In: Progress and problems in lichenology in the eighties. Bibliotheca Lichenologica 25: 185- 195.