J. Hattori Bot. Lab. No. 100: 695- 720 (Aug. 2006)

THE FAMILY IN THE MONTANE REGIONS OF NEW GUINEA

PER M. J0RGENSENI AND H. SIPMAN2

ABSTRACT. The Pannariaceae of the montane regions of New Guinea contain 37 species in nine genera. Ten species are endemic, mostly from the subalpine zone. The following species are newly described: cacuminum, F venusta, tenuis (from Borneo), foli­ icola, P. laceroides, P. montana, P. nitida, P pannarioides, P. piundensis and filicicola, and two new combinations are proposed: lecideola (Zahlbr.) (Java) and Pannaria papuana (Apt­ root & Diederich) (new to Borneo). Pannaria andina P. M. J0rg. & Sipman and Pannaria aff. rubig­ inella P. M. Jmg. & Sipman are reported as new to Papua New Guinea, as well as Parmeliella hawai­ iensis H. Magn.; the former two and Pannaria prolificans Vain. are also new to Borneo and part of a shared element between these disjunct, very high mountains. Parmeliella endomilta (Vain.) Makhija & Adawadkar (syn. novo Parmeliella endoferruginea Aptroot and Parmeliella pannosa var. erythro­ cardia MiiI!. Arg.) is new to Borneo, Indonesia, Madagascar and Mauritius, Parmeliella philippina (Vain.) P. M. Jmg. new to Java and New Guinea, and Parmeliella polyphyllina P. M. J0rg. new to the Philippines. The origin of the Pannariaceae in this region is predominately paleotropical, mainly Aus­ tralasian, but at high altitudes there are boreal, subantarctic and Andean elements, the latter being most intriguing.

INTRODUCTION The lichen flora of New Guinea is poorly known, although it was catalogued by Szata­ la (1956) and more recently by Streimann (1986), and some later publications (especially Aptroot et al. 1997) have attempted to update our knowledge of this lichen-rich island. The Pannariaceae contain several interesting taxa in New Guinea, but have never been revised in toto. The best available treatment from SE Asia is that ofVainio (1921) from the Philip­ pines. The rich collections made by the second author and co-workers, as well as by others, on repeated trips to the mountains of Papua New Guinea (1981- 2005) has made it possible to get a better understanding of the taxa. Although some collecting was also done in the . lowlands, this treatment is restricted to the montane region (from the lower limit of the Nothofagus forest at C. 1500 m alt. (see Paijmans 1976) to the upper mountain grasslands at C. 4000 m alt. or more). Pannariaceae also occur below 1500 m, but mainly taxa of difficult pantropical complexes (e.g. Parmeliella mariana) which are in need of more detailed field studies, especially in other parts of their distribution area. In order to more fully understand the taxa of New Guinea, comparison with material from nearby islands was necessary, and observations and comments on that material are also included when relevant. The following genera are known in the region (those for which recent revisions are

I Botanical Museum, University ofBergen, Allegt 41, N-5007 Bergen, Norway. 2 Botanisches Museum, Konigin-Luise-Str. 6-8, 0-14195 Berlin, Germany. 696 1. Hattori Bot. Lab. No. 100 2 0 0 6 available will not be treated in detail below): 1. Thallus gelatinous, homoiomerous, PD- ...... Kroswia P. M. J0rg. Thallus mostly non-gelatinous; if gelatinous, then heteromerous and PD + ...... 2 2. Thallus foliose; penetrating into the subhymenial parts ...... 3 Thallus squamulose or placodioid: no cyanobacteria in the subhymenial parts ...... 5 3. Thallus with stiff hairs, mostly PD+ orange; apothecia marginal ...... Fee (7 species, see J0fgensen & Sipman 2002a). Thallus with arachnoid hairs or glabrous, always PD - ; apothecia laminal ...... 4 4. Thallus bluish, upper surface arachnoid; asci with I + blue cap ...... Nyl. (2 species, see Galloway & J0rgensen 1987). Thallus brownish, upper surface smooth; asci without I + blue cap ...... Fuscoderma (D. 1. Galloway & P. M. J0fg.) P. M. J0rg. & D. 1. Galloway (I species, see J0fgensen & Sipman 2002b) 5. Apothecia with thalline margin ...... 6 Apothecia without thalline margin ...... 8 6. Thallus always PD-; asci with apical amyloid ring structure ...... 7 Thallus usually PD+ orange; asci without apical amyloid ring structure ...... Pannaria 7. Thallus rosette-forming, resting on prominent prothallus, with bluegreen photobiont; hymenium I + bright blue; predominantly lowland ...... Parmeliella mariana group Thallus irregularly small-squamulose, not resting on a prominent prothallus, with green photo- biont; hymenium I + dirty blue; montane ...... Psoroma 8. Hymenium 1+ persistently bright blue; thallus without lichen substances ...... 9 Hymenium I + blue-green, soon turning red-brown (hemiamyloid); thallus with triterpenoids and fatty acids ...... Fuscopannaria 9. Thallus lobes concentrically striate, pale grey, with dense medulla of horizontal hyphae; asci with amyloid apical sheet ...... Degelia Thallus lobes not concentrically striate, bluish to brownish grey, with loose medulla of irregularly orientated hyphae; asci with amyloid apical ring ...... Parmeliella

THE TAXA

Degelia Arv. & D. 1. Galloway This as originally described by Arvidsson & Galloway (1981) comprised some subtropical to warm-temperate coccocarpioid species previously included in Parmeliella, but later it was enlarged to include more small-squamulose, alpine-subantarctic species (sect. Frigidae P. M . J0rg., J0rgensen 2004b). Aptroot et al. (1997) described two species from alpine New Guinea, but there is a further taxon in the region, previously placed in Parmeliella. The species belong in the small subantarctic element of the lichen biota in the region. All are apparently rare, occurring at high altitudes. The three species can be distin­ guished as follows: I. Thallus sorediate ...... D. sorediata Thallus non-sorediate ...... 2 2. Squamules greyish brown, at least 2 mm diam. ; spores 10- 12.um long; Java ...... D. lecideola Squamules pale grey, to I mm diam.; spores 15- 17 .urn long; New Guinea ...... D. minor No new collections of D. minor or D. sorediata have become available since the de­ tailed descriptions of Aptroot et al. (1997). D. minor is only known from 3600-3700 m on P. M. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 697

Fig. 1. Degelia lecideola , detail ofholotype (W).

Mt. Wilhelm, while D. sorediata is more widespread, also being known from Mt. Kinabalu on Borneo. D. lecideola is only known from Mt. Lawu in Java, from above 3000 m altitude:

Dege/ia lecideola (Zahlbr.) P. M. J0rg. & Sipman comb. novo (Fig. I) Parmeliella lecideola Zahlbr. , Beih. Feddes Rep. 127: 69 (1943). Type: Java, Gunung Lawu, 3240 m, 20 Dec. 1928, F. Ruttner (w, holotype). Zahlbruckner (1943) presents a detailed description, to which can be added that it has the apical apparatus typical of the genus Degelia, with an amyloid apical sheet, rather than the ring-structure found in Parmeliella. This species is larger than D. minor, and darker (greyish brown). It also differs in spore-size.

Fuscopannaria P. M. J0rg. This genus, with an evolutionary centre in Pacific North America and the SE Asian mountains (J0rgensen 2000b, 2002a), is also complex in this region, with several species. It is a temperate Laurasian element. The species can be distinguished as follows: I. Thallus with bluish gymnidia* ...... F. coerulescens Thallus without gymnidia ...... 2 2. Squamules large (to 5 mm long), uniform, widely spread on the prothaltus ...... F. venusta Squamules smaller (to 2 mm long), often white-edged, mostly cushion-forming ...... 3 3. Squamules thick, imbricate, in cushions; spores 15-18 x 9-12 f.1m; on high-alpine soils ...... F cacuminum Squamules thinner, partly ascending; spores 10-12 X 8-1 0 f.1m ; on rotting bark in humid forests ...... F dissecta * gymnidia are isidia without cortex (see J0rgensen & Kashiwadani 2001).

Fuscopannaria cacuminum P. M. J0rg. & Sipman sp. novo (Fig. 2) Fuscopannariae coerulescentis simitis, sed sine gymnidiis, caespitosa; ascosporae magnae, ellipsoideae, 15- 18 X 9-12 mm; terricola et alpina. Type: Papua New Guinea, Simbu Prov. , Mt. Wilhelm, Pindaunde valley, along trail from Lake Piunde to the summit peak, on top of exposed ridge, in short grass vegetation, 4000 m, 13 March 698 1. Hattori Bot. Lab. No. 100 2 0 0 6

Fig. 2. Fuscopannaria cacuminum, holotype (B).

1967, H. Sipman 22055 (B, holotypus). Thallus small-squamulose, individual squamules brownish with paler margins, 1- 2 mm diam., apically indented, imbricate, forming small cushions; in section 250- 3501lm thick with cellular, thick-walled, 30-50llm thick upper cortex, and without lower cortex. Apothecia rare, fiat, blackish brown, irregularly spreading, to 2 mm broad with indistinct thalline margin; ascospores simple, colourless, ovoid, 15- 18X9- 12Jlm, with distinct, smooth exospore with rounded apices. Ch emistry: All reactions negative, thallus containing fatty acids and triterpenoids (TLC). This appears to be the fertile counterpart of F. coerulescens, which is a species of the upper forests, extending into the subalpine scrub. The two are immediately distinguishable since F. cacuminum lacks the bluish gymnidia which are so characteristic of F. coerulescens. The squamules are thicker and crowded, forming caespitose, subfruticose structures. In colour they are darker than in F. coerulescens; the margins, however, appear frosted and develop needle-shaped crystals in the herbarium. The blackish brown apothecia are fiat, irregularly spreading and larger than in that species. There is an ecologically simi­ lar species on high levels in the Himalayas, F. saltuensis P. M. J0rg., which has a quite dif­ ferent, paler brown thallus without any lichen substances and with smaller, convex apothe­ cia. It is interesting to note the adaption of species in this genus to this particular environ­ ment in two different regions of high mountains. Habitat and distribution: This terricolous species occurs in short grass vegetation and in rock crevices. It appears to be endemic, but closely related to other East Asian species of the genus. Specimens examined: PAPUA NEW G UINEA, Simbu Prov., Bognotto Ridge, 4 km SE of Mt. Wil­ helm, 4000m, July 1968, D. McVean 68186 (CANB); Mt. Wilhe1m, along track to the summit, 4200 m, 7 Aug. 1992, P. Diederich (Herb. Diederich). P M. JORGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 699

Fuscopannaria coerulescens P. M. Jl1Irg. This species, previously confused with F leucosticta, has proved to be quite common. It was often confused with the larger, PD+ orange Pannaria conopiea, a rare species in this region (see below), due to the characteristic bluish gymnidia. Description available in J0rgensen (2000b). Habitat and distribution: This widespread species on debris or rotting stems, from c. 2000-3600 m, is an Asian element. Specimens examined (selected): PAPUA NEW GUI NEA, Eastern Highlands Prov., Gahavisuka Prov. Park, above Nagamiza village, N of Goroka, 2000 m, 8 May 1982, H. Streimann (CANB, holotype). Madang Prov., Huon Peninsula, Finistere Range, Yupna valley, Teptep village, trail in NNW and deep valley in N direction, 2500 m, 30 July 1992, P. Diederich 10279, 10680 (Herb. Diederich); Saruwaged Range, Honzeukngon village, S of Derim airstrip in Timbe valley, 1950m, 7 March 1967, H. Sipman 24382 (B). Morobe Prov., Mt. Sarawahat, 4km NE of Lake Owam, 2850m, 1983, T. Koponen 31543 (H). Simbu Prov., Imbuka Ridge, ENE of Lake Piunde, 3600 m, 27 Aug. 1970, R. Hnatiuk (CANB); Pindaunde valley, near hut on the shore of Lake Piunde, 3500 m, 5 Aug. 1992, P. Diederich 10236, 11206 (Herb. Diederich).

Fuscopannaria dissecta P. M. Jl1Irg . This little known species is close to the generic type, F ieucosticta, but has incised squamules resting on a thick blackish hypothallus and clearly smaller, subglobose spores, 10-12X8-\0 f..lm. Description available in Jl1Irgensen (2000b: 250-251 and 2002a: 226-227). Habitat and distribution: Growing on rotting bark of trees, it is new to New Guinea, the southernmost known locality (Jl1Irgensen 2002a: 226) of this SE Asian flora element. Specimens examined: PAPUA NEW GUINEA, Central Prov., Mt. Albert Edward, en route from tent site to summit area, 3000 m, 24 Oct. 1975, S. Kurokawa 9383 (TNS). Eastern Highlands Prov., Daulo Pass, 18km WNW of Goroka, 2550m, 7 Apr. 1982, H. Streimann 18128 (CANB). Simbu Prov.,

Fig. 3. Fuscopannaria venusta, holotype (B). 700 J. Hattori Bot. Lab. No. 100 2 006

300 600 900 1200 1500 1800

Fig. 4. World distribution of Kroswia crystallifera.

Lake Aunde, 6 km SE ofMt. Wilhelm, 3520m, May 1966, D. McVean 66178 (CANB).

Fuscopannaria venusta P. M. J0rg. & Sipman sp. novo (Fig. 3) Thallus squamulosus; squamuli avellanei, integri, ad 5 mm diam., dispersi prothallo atroveneto insidentes; acidis aliphaticis et terpenis continens. Apothecia ignota. Type: Papua New Guinea, Simbu Prov., Mt. Wilhelm, Pindaunde valley, near the hut on the S­ shore of Lake Piunde, subalpine forest remnants, on W-slope of valley, c. 3700 m, 6 Aug. 1992, H. Sipman 35678 (B, holotype). Thallus squamulose, widely spreading (to 5 cm); individual squamules to 5 mm diam., entire, with a tendency to form isidia-like structures, totally brown, resting on a thin bluish­ black prothallus; in section 150- 200 f..lm wide with cellular, thick-walled, 30-40 f..lm thick upper cortex, without lower cortex. Apothecia unknown. Chemistry: All reactions negative, thallus containing fatty acids and terpenoids (running high in TLC). This elegant species has been difficult to place generically since it is sterile and much larger than its relatives, being morphologically rather Pannaria-like. However, the anatomy (small-celled cortex) and the chemistry (fatty acids and terpenoids) clearly indicate Fusco­ pannaria. Its large, sometimes isidiate squamules are unique in the genus, so that the species cannot be confused with any other in the genus. Habitat and distribution: Rare and presumably endemic, only known from the rich type-collection in the subalpine scrub near Lake Piunde, where it overgrows mossy branches. PM. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 70 1

Kroswia P. M. J0rg. This recently described, paleotropical genus (J0rgensen 2002b) has a unique, gelatinous, homoiomerous thallus and is easily mistaken for a sterile (Collemataceae), due to its strongly swelling thallus (in moisture). Only one species is present in New Guinea.

Kroswia crystallifera P. M . J0rg. This species, described in detail by J0rgensen (2002b), somewhat resembles Pannaria globigera Hue from which it is readily distinguished by the gelatinous, non-corticate thal­ lus, which is PD-. Habitat and distribution: The New Guinea specimens were collected in mossy mon­ tane forest at c. 2500m. This is a paleotropical element (Fig. 4). Specimens examined: PAPUA NEW GUINEA, Eastern Highlands Prov., Daulo Pass, 18 km WNW of Goroka, on gentle ridge, 2600 m, 7 Apr. 1982, H. Streiman 18061 (CANB); Mt. Gahavisuka Prov. Park, II km N of Goroka, 2300m, 17 March 1987, H. Sipman 22181b (B). Madang Prov., Huon Peninsula, Finisterre range, Yupna valley, Teptep village, trail in NNW and deep valley in N direc­ tion, 2500 m, 30 July 1992, P. Diederich 10791 (Herb. Diederich); towards Bambu airfield, 2500 m, 30 July 1992, H. Sipman 35262 (B).

Pannaria Del. This predominantly tropical genus appears to be fairly rare in Papua New Guinea, with few, rather locally occurring taxa, mostly at moderate altitudes, and is, with one re­ markable exception discussed below, mainly paleotropical. 1. Thallus with bluish gymnidia* ...... P conoplea Thallus without gymnidia ...... 2 2. Thallus with isidia ...... P prolificans Thallus without isidia ...... 3 3. Thallus with green photobiont and cephalodia ...... 4 Thallus with cyanobiont only ...... 5 4. Thallus dominated by large, spreading lobes with green photobiont; cushion-shaped cephalodia present centrally (not treated further)...... P sphinctrina complex** Thallus dominated by small, brittle, partly ascending squamules with bluegreen photobiont, green photobiont only in the marginal squamules ...... P papuana 5. Thallus gelatinous, swelling much when wet, strongly wrinkled when dry ...... P lurida Thallus not gelatinous, not wrinkled ...... 6 6. Thallus grey-brown, with flat lobes or nearly crustose, resting on a rhizohyphal mat ...... 7 Thallus blue-grey, foliose, with coarse, bundled marginal rhizohyphae ...... 8 7. Thallus with elongated, radiating lobes, resting on a thick cushion of rhizohyphae; ascospores with smooth, gelatinous, tailed exospore, ellipsoid, 16-20 X 9-1 O)lm ...... P andina Thallus squamulose-crustose, on thin layer of rhizohyphae; ascospores warted, subglobose, 12- 15 X 8-9 )lm ...... P aff. rubiginella 8. Lobes narrow, to c. 1 mm broad, apically revolute, PD-; Borneo only ...... P tenuis Lobes broader, to c. 3 mm broad, apical\y flat, PD+ orange; widespread ...... P molkenboeri * Gymnidia are isidia without cortex (Jorgensen & Kashiwadani 2001). ** This complex is still under study; none of the PNG material belongs in P sphinctrina s. str. 702 1. Hattori Bot. Lab. No. 100 2 0 0 6

Pannaria andina P. M. J0rg. & Sipman A recently described species (J0rgensen & Sipman 2004) in the Pannaria rubiginosa complex, characterized by its nearly crustose, small-squamulose thallus resting on a volu­ minose cushion of rhizohyphae, and by spores with a very distinct, gelatinous exospore with apical "tails", which may be of importance for dispersal (Elvebakk & Bjerke 2005: 51 ). Habitat and distribution: This species of the subalpine scrub is most commonly found at c. 3500 m in both New Guinea and Borneo, but occasionally on rocks at higher altitudes (to 4200 m) and sometimes as low as the upper parts of the montane forests. It is an An­ dean element. Specimens examined: MALAYSIA, Sabah, Distr. Ranau, Kinabalu Park, S-slope of Mt. Kinabalu, along summit trail, near Sayat Sayat hut, 3800m, 11 May 1989, H. Sipman 31174 (B). PAPUA NEW GUINEA, Simbu Prov., Mt. Wilhelm, Pindaunde valley, near the hut on the shore of Lake Piunde, 3600 m, 5 Aug. 1992, P. Diederich 10278, 10293, 10303 (Herb. Diederich), H. Sipman 22142 (B); Mt. Wilhelm, by summit track, c. 4000 m, May 1966, D. McVean 66260 (CANB); along track to the summit, 3900-4200 m, 7 Aug. \992, P. Diederich (Herb. Diederich); Mt. Wilhelm, en route from Keglsugl to Kombugomanbuno, 3300 m, 6 Jan. 1974, H. Kashiwadani 11480 (TNS); en route from Kombugomanubo to the Pindaunde Lakes, 3400-3600m, 3 Jan. 1974, H. Kashiwadani 11114-15 (INS). Western Highlands Prov., Kim Magei Mt., 8 km SW of Mt. Hagen, 2500 m, 23 June 1982, H. Streimann 20648 (CANB).

Pannaria conoplea (Ach.) Bory This mainly cold-temperate species is rare in Asia, and is known with certainty from only one locality in Papua New Guinea, although several specimens in herbaria have erro­ neously been named so, due to confusion with the fairly common Fuscbpannaria coerulescens which also has bluish gymnidia, but a smaller, browner thallus containing fatty acids and terpenoids (clearly visible as crystals after some time in the herbarium) in­ stead of pannarin and with a negative PD-reaction. A description of P conoplea is provided by J0rgensen (1978: 21-22). Habitat and distribution: The only New Guinea specimen was found on a shrub in subalpine grassland. It is a northern, temperate element. Specimen examined: PAPUA NEW GUINEA, Simbu Prov. , Mt. Wilhelm, Pindaunde valley, near hut on S-shore of Lake Piunde, 3500 m, 12 March 1987, H. Sipman 21972 (B).

Pannaria lurida (Mont.) Nyl. Syn. novo P sublurida Nyl. (from Java, Nylander 1859). This species is usually easily recognized due its gelatinous nature and strong wrinkles when dry. A description is provided in Flora of Australia 54 (1992: 268). Habitat and distribution: This is a species of moist, fairly light, open lowland forest­ edges, just reaching the lower parts of the zone under treatment. It is a predominantly South Pacific species, but reaches eastern America (J0rgensen 2001a), including Brazil (type of P brasiliensis Zahlbr.). Specimen examined: PAPUA NEW GUINEA, Central Prov. , 2 km N of Woitape Airstrip, 1200- 1350 m, 19 Jan. 1974, H. Kashiwadani 11707 (INS). P. M. JORGENSEN & H. SIPMAN : Lichen family Pannariaceae in the montane regions of New Guinea 703

Fig. 5. Pannaria molkenboeri from Goroka (Herb. Diederich).

Pannaria molkenboeri (Mont.) Hue This is a much misunderstood species, originally described from Java, which has been confused with P iurida, a much thicker, wrinkled species (J0rgensen 2004a). Its variation is poorly understood since the type specimen is small and few other specimens have been available for study. J0rgensen (2004a) realized that the large type specimen of P tjibodensis Zahlbr., also from Java, represented the same species, being characterized by a thin, hardly swelling, neatly divided thallus with PD+ orange medulla and protruding, coarse, black­ ish-blue rhizohyphae. The Goroka material (Fig. 5) is unusually finely divided, but has the basic characters of this as yet poorly understood species, so it is for the time being kept within P molkenboeri. Habitat and distribution: This species of mossy, primary forests is presently only known from widely separated localities in New Guinea at 1350 m (typical of the species) and c. 2500 m altitude. It is an Australasian element. Specimens examined: PAPUA NEW GUIN EA, Central Prov., about 2 km N of Woitape Airstrip, 1200- 1350 m, 19 Jan. 1974, H. Kashiwadani 11658 (TNS). Eastern Highlands Prov., Mt. Gahavisuka Prov. Park, 11 km N of Goroka, along trail to lookout, 2400 m, 3 Aug. 1992 (Herb. Diederich). West­ ern Highlands Prov., Kum Magei Mt. , 8 km SE of Mt. Hagen, 2500 m, 23 June 1982, H. Streiman 20645 (CBG). Morobe Prov., Herzog Mts. , IS km WSW of Lae, 760 m, 13 Jan. 1981 , H. Streimann & T. Umba 10944 (B, CANB). Simbu Prov., Mt. Wilhelm area, Bundi Gap, on road Keglsugl-Bundi, 2800 m, 4 Aug. 1992, H. Sipman 35581 (B).

Pannaria papuana (Aptroot & Diederich) P. M. J0rg. & Sipman comb. novo Basionym: Psoroma papuanum (as "papuana") Aptroot & Diederich, Tropical Bryology 14: 30 (1998). - Type : Papua New Guinea, Bundi Gap, on road Keglsugl-Bundi, subalpine forest, 2800 m, 4 Aug. 1992, A. Aptroot 32189 (Holotype, B). Synonym: Psoroma pannarioides Aptroot & Diederich, nom. illeg. , Biblioth. Lichenol. 64: ISO (1997) - non Henssen (1983). 704 1. Hattori Bot. Lab. No . 100 200 6

Fig. 6. Pannaria papuana, part of fertile material from Borneo (TNS).

This most characteristic and unusual species, which Aptroot et al. (1997: ISO) recog­ nized as very Pannaria-like (as indicated by the original epithet), was placed in Psoroma since they regarded the blue-green parts as cephalodia, a view recently repeated by Elve­ bakk & Bjerke (2005). However, as shown by J0rgensen (200 I b), the choice of photobiont is not important taxonomically at genus level. Fertile material exhibits apothecia (Fig. 6) with the typical characters of Pannaria, a deep blue hymenial reaction and asci without apical amyloid ring structure. The rather thin squamules with single-layered cortex and the thallus without pannarin place it in the Australasian subgenus Lepidoleptogium (A. L. Smith) P. M. J0rg. (the Pannaria immixta-complex, J0rgensen 2004b) as the only known species which contains both green and blue-green photobionts, though P crenulata P. M. J0rg. & D. 1. Galloway gives the impression of being a blue-green counterpart of such a species. Sterile material without the green marginal squamules is easily mistaken for a Parmeliella, especially P polyphyllina. Habitat and distribution: This is a fairly common and widespread species in the moun­ tains of New Guinea (Eastern Highlands, Madang, Simbu and Western Highlands Provinces) in mossy forests at c. 2500 m and upwards into the lower subalpine region shrubbery at 2900 m. It is recorded here as new to Borneo, and is a Malesian element. Specimens examined (additional to those cited in Aptroot et aI. , 1997): MALAYSIA, Borneo, Sabah, Mt. Kinabalu, southern slope, between Kambaranga radio station and second radio station, 2 146-2500 m, 18 May \963, M. Mitzutani 2639 (TNS); Mesilau trail to W. Mesilau river, 1800m, 1965, M. E. Hale 29205 (US). PAPUA NEW GUINEA, Eastern Highlands Prov., Daulo Pass, 18 km NW of Goroka, 2600m, 7 Apr. 1982, H. Streimann 18049 (CANB). Morobe Prov., Ekuti Divide, Bulolo­ Aseki road, 33 km WSW of Bulolo, 2260 m, 17 June 1982, H. Streimann (B, CANB, H, LAE). Southern Highlands Prov., Pavende Logging area, 15 km NNW of Ialibu, 2400 m, 14 Dec. 1982, 1. A. Elix & H. Streimann (B, CANB). Western Highlands Prov. , Mur Mur Pass, 7 km NE of Tambul, 2800 m, 27 June 1982, H. Streimann 21202 (B, CANB). P. M. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 705

Pannaria prolificans Vain. This isidiate species differs from all others by its elongated marginal lobes resting on a distinct black prothallus, and its marginal, coralloid isidia. A detailed description is pro­ vided by J0rgensen (2003b: 253- 254), who reports that the cited South American material is rather different from the typical material from the Philippines. The Malesian specimens have more strongly developed rhizohyphae, partly with unusual, small-squamulose thallus and knobbly isidia. We interprete these at the moment as extreme expressions of this vari­ able species, but they may prove to represent a distinct taxon when more and better-devel­ oped material becomes available. Habitat and distribution: This corticolous species of moist, mossy forests at c. 2500 m is basically an Antillean element. Previously it was only once recorded with certainty from Australasia (Philippines, Mindanao, J0rgensen 2003b: 254). Specimens examined: MALAYSIA , Borneo, Sabah, Kota Belud distr., S-slope of Mt. Kinabalu, area below Paka cave, 2500-2600 m, 18 May 1963, Z. Iwatsuki (TNS); below Layang Layang radio station, 2400m and on summit trail, c. 3200 m, H. Sipman & B. Tan 31241, 31330 (B). PAPUA NEW GUIN EA, Northern Prov., Owen Stanley Range, Myola, 0-1 km along trail from guesthouse towards NE, on right bank ofIora river, 2100- 2400 m, 15 Oct. 1995, H. Sipman 38264 (B). Simbu Prov., Mt. Wilhelm area, Bundi Gap, on road Keglsugl-Bundi, 2800 m, 4 Aug. 1992, H. Sipman 35504 (B).

Pannaria aff. rubiginella P. M. ]0rg. & Sipman The recently described P rubiginella is characterized by its squamulose-crustose thal­ lus resting on a thin prothallus, the weak PD+ reaction (if any) and the subglobose, rugu­ lose spores (see J0rgensen & Sipman 2004). The PNG material differs in its rather vivid PD reaction, but since all specimens exhibit all the other characteristics of the species, we refer them tentatively to this species, rather than regarding them as extreme forms of P an­ dina. Habitat and distribution: This species is known from the upper parts of the montane forests, in remnants of forest towards the subalpine grasslands, at c. 2500 m. This Pacific element is otherwise known from cool, moist habitats on the Pacific coast of the Americas. Specimens examined: PAPUA NEW GUINEA, Eastern Highlands Prov., Daulo Pass, 18 km NNW of Goroka, 2600 m, 7 Apr. 1982, H. Streimann 18060 (CANB). Simbu Prov., Mt. Wilhelm, Bundi Gap, on road Keglsugl-Bundi, 2800 m, 4 Aug. 1994, H. Sipman 35567 (B).

Pannaria tenuis P. M. J0rg. & Sipman sp. novo (Fig. 7) Pannariae molkenboeri similis, sed thallo tenuissimo, lobis angustissimis apicibus rev­ olutis, sine acido lichenoso (PD-). Type: Malaysia, Borneo, Sabah, Ranau Distr., Kinabalu Park, S-slope of Mt. Kinabalu, sur­ roundings of Headquarters, in valley along Silau Silau trail, 1650 m, 14 May 1989, H. Sip man & B. Tan 31378 (B, holotype). Thallus foliose, grey-blue, to 2 cm wide, composed of narrow, to I mm broad, deeply divided lobes with protruding, coarse blackish-blue rhizohyphae; in section 100- 150 pm thick; upper cortex 30-40 pm thick, cellular, of rectangular cells, above a distinct, to 30 pm thick cyanobiont layer of small-celled, "violet" Nostoc in chains, with a loose, 40- 90 pm thick medullary layer below gradually merging into the long rhizohyphae. Apothecia and 706 1. Hattori Bot. Lab. No. 100 2 0 0 6

Fig. 7. Pannaria tenuis, holotype (B). pycnidia unknown. Chemistry: All reactions negative, no substances detected by TLC. This characteristic species is difficult to place within this genus since it is sterile and exhibits unusual characters, namely very narrow lobes which are revolute apically, and no pannarin. However, the well developed rhizohyphae and the anatomy clearly link it to Pan­ naria molkenboeri, the Goroka form of which is morphologically quite close, but differs in anatomy (thicker) and chemistry (presence ofpannarin). Habitat and distribution: Known only from the type specimen, which was collected on mossy stems of a tree at fairly low altitude, indicating a possible wider distribution in the lowlands of Malesia.

Parmeliella MUll. Arg. This is the most complex genus of the Pannariaceae in the region, with several poorly understood species complexes, particularly in the lowlands. The montane taxa are also dif­ ficult to classify, but the rich material available does allow some conclusions which show that there are surprisingly many, little known taxa in the region, and more may be hiding among sterile, poorly developed collections. A key to their identification is difficult to make, but we have nevertheless made a first attempt, mainly using characters of the apothecia which unfortunately are not always pre­ sent. When the specimens are sterile one has to try both possibilities (at 4) in the key. I. Thallus with yellow or orange pigments ...... 2 Thallus grey-brown or blue-grey, without yellow/orange pigments ...... 3 2. Thallus brown, not isidiate; medulla orange; often fertile, spores citriform ...... P endomilta Thallus grey with a yellowish hue, sparingly isidiate; medulla whitish; rarely fertile, spores el- lipsoid ...... Pflavida 3. Thallus very thin (l ess than 100 .urn), foliicolous ...... P./oliicola Thallus thicker ( I 00 .urn or more), not foliicolous ...... 4 4. Apothecia with thalline margin ...... 5 P M. J0RGENSEN & H. SIPMAN : Lichen family Pannariaceae in the montane regions of New Guinea 707

Apothecia only with proper margin ...... 10 5. Thallus very thick (c. 400,um), with prominent, brownish rhizohyphae ...... P. pannarioides Thallus thinner (at most 200 ,urn), with inconspicuous or blackish rhizohyphae ...... 6 6. Thallus flat, resting on prominent prothallus ...... 7 Thallus cushion-forming, with at least some ascending lobes/squamules, prothallus less promi- nent ...... 8 7. Thallus large (at least 5 cm diam.), on spongy prothallus; lowland ...... P. mariana s.lat. * Thallus smaller (not more than 5 mm diam.), on crustose prothallus; montane ...... P. montana 8. Thallus coralloid, usually shiny, in dense cushions ...... P. nitida Thallus squamulose, dull, mainly in loose cushions ...... 9 9. Squamules uneven, appearing gnarled, often with exposed, bluish lower surface ...... Pannaria papuana cyanomorph. Squamules smooth, regular; lower surface, if showing, white ...... P. polyphyllina 10. Thallus with enlarged, radiating marginal lobes ...... P. piundensis Thallus of more or less equal squamules ...... 11 11. Thallus nearly crustose with neatly incised lobes resting on blackish crustose prothallus ...... P. philippina Thallus distinctly squamulose, prothallus irregular or poorly developed ...... 12 12. Thallus with numerous lobules in lace-pattern; apothecia with thick, permanent proper margin (> 100 ,urn); spores warted ...... P. laceroides Thallus mostly without lobules, forming flat rosettes; apothecia with thin proper margin, often excluded at maturity « 100 J1m); spores with smooth exospore ...... P. hawaiiensis * Difficult lowland complex, not treated further.

Parmeliella endomilta (Vain.) Makhija & Adawadkar Syn. nov.: Parmeliella endoferruginea Aptroot (Bib!. Lich. 74, 1997); Pannaria pannosa var. erythrocardia MiiI!.Arg. (Flora 70, 1887). This is a rare, often fertile (apothecia with thalline margins), small-squamulose species, which is easily recognized by the thin, skin-like thalli, the orange-red medulla and the citriform spores (see Aptroot et al. 1997: 128). The type of Pannaria endomilta is ster­ ile with numerous secondary lobules, but otherwise with the typical thallus of the species. Habitat and distribution: A species of moist, tropical mountain forests, commonest below 1500 m. Evidently a paleotropical element, mainly lowland in PNG, now also known from Borneo, Java, Madagascar and Mauritius, in addition to the previous records from PNG (Aptroot et al. 1997). Specimens examined: INDONESIA, Java, corti cola, without collector (L, holotype of var. erythro­ cardia). MALAYSIA, Borneo, Sabah, Mt. Kinabalu, between Kambaranga Radio station and Water­ falls, 2000-2146 m, 17 May 1963, Z. Iwatsuki 550b (TNS). Sarawak, Gunung Mulu National Park, 4th Div., Baram distr., valley ofUlu Jerneh, 500m, 5 Apr. 1978, B. Coppins 5257, 5317 (E). MADA­ GASCAR, Diego-Ruarez, Col de Ramada, 60 km S of Anthalaha, 500 m, A. Aptroot & R. Henssen (Herb. Aptroot). MAURITIUS, Savanne, along road between Mt. Cocotte and Basin Blanc, 550-580 m, H. Krog & E. Timdal MAU 58/08 (0). PAPUA NEW GUINEA, Central Prov., near Dbamura on Owert's Corner Road, 40 km NE of Port Moresby, 580 m, 10 Febr. 1981, H. Streimann & E. K. Naoni 14927 (CANB); K. B. sawmill, Ehu Creek, 12km SW ofSogeri, 750m, 16 Febr. 1981, H. Streimann & E. K. Naoni 16630 (CANB). Gulf Prov., Werr River, Kaintiba, 640 m, 24 Jan. 1983, H. Streimann 33578 (CANB). Eastern Highlands Prov. , Waiopa, Aiyura-Omaura road, 13 km SE of Kainantu, 1450 m, 8 708 1. Hattori Bot. Lab. No. 100 2 0 0 6

Dec. 1982, 1. A. E1ix & H. Streirnann 12402 (CANB). Morobe Prov., Aseki-Manyamya road, 6 km NW of Aseki, 1950 rn, 5 Dec. 1982, 1. A. E1ix & M. Toia 12110 (CANB); Pouyu village, 2 km SE of Aseki, 1500 rn, 24 Jan. 1981 , H. Streirnann & E. Tarnba 12664 (CAN B); Herzog Mts, 15 km WSW of Lae, 760 rn, 13 Jan. 1981, H. Streimann & T. Urnba 11051 (CANB). Southern Highlands Prov., Piribu sawmill, Tari-Komo road, 3 km SW of Tari, 1650 m, 15 Dec. 1982, 1. A. Elix & H. Streimann 13199 (CANB); Tari-Komo road, 6km NW of Komo, 1480m, 16 Dec. 1982, 1. A. Elix & H. Streirnann 13199 (CANB). PHILIPPI NES, Mindanao, Davao, Mt. Apo, 6000 ft. , 21.4 1904, E. B. Copeland 1090 p.p. (TUR-VA[N 12 [68, ho[otype of P. endomilta).

Parmeliella jiavida P. M. J0rg. This recently described species (J0rgensen 2003a) is easily recognized by its faint yel­ lowish colour (due to an unknown xanthone). The rather small PNG sample is very spar­ ingly isidiate, which is not taxonomically important as the known Philippine material is quite variable in this character. Habitat and distribution: Corticolous on trunk in lower montane forest. A most sur­ prising discovery of a species thought to occur rather locally in the Philippines (Luzon, Basilian) (J0rgensen 2003a). Specimen examined: PAPUA NEW GUINE A, Morobe Prov. , 7 km SE of Bulolo, 1200- 1320 m, 21 Dec. 1973, H. Kashiwadani 10830 (TNS).

Parmeliella /oliicoia Aptroot & P. M. J0rg., sp. novo (Fig. 8) Thallus praecipue foliicola, placodioideus, radiatus, asteroideus. Apothecia saepe ag­ gregata, marginibus thallinis variabilibus; ascosporis verrucosis, IS- 22(- 30) X 10- 12 ,urn. Type : Papua New Guinea, Eastern Highlands, Mt. Gahavisuka Prov. Park, 11 km N of Goroka, along trail to outlook, 2400 rn, 3 Aug. 1992, P. Diederich 10665 (LG, holotype).

Fig. 8. Parmeliellafoliicola, holotype (Herb. Diederich). P. M. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 709

Thallus placodioid, consisting of radiating, grey-blue to brownish lobes which are confluent in the centre, to 0.2 mm wide, and 50(-70),um thick, often with upturned tips or lobules, resting on a thin, felty, cream to blackish hypothallus. Apothecia sessile, flat, round to lobate, 0.5-1.2 mm diam., brown with paler, cellular proper margin, sometimes obscured by thalline squamules; hymenium 100,um high, brown in upper parts, I + blue; subhymemium, pale brown, of intricate hyphae, 50,um high; ascospores simple, hyaline, broadly ellipsoid, I 5- 22(- 30) X 10-12 ,um, ornamented with coarse, rounded warts. Chem­ istry: All reactions negative, no substances detected in TLC. This is a most characteristic species due to its thin thallus with confluent lobes which are upturned apically, and its main habitat. Habitat and distribution: This is the only species of the genus which is known to be predominantly foliicolous (sometimes also on culms of grasses, or rarely on smooth bark). It is fairly common in upland forests at c. 2500 m and rather widespread in the region, pos­ sibly a SE Asian element. Specimens examined: INDON ES IA, lrian Jaya, Star Mts, Ok Bon, van Zanten 434, 437 CL, herb. Aptroot). Sumatra, western part, Mt. Singalang, reg. nubium, 1400-1600 m, 25 July 1894, V Schiffn­ er 3414 (TNS, W). MALAYSIA, Borneo, Sabah, Ranau Distr., Kinabalu park, S-slope ofMt. Kinabalu, surroundings of Headquarters, in valley along Silau Silau, 14 May 1989, H. Sipman & B. Tan 29462 (B). PAPUA NEW GUINEA, Eastern Highlands Prov., Mt. Gahavisuka Prov. Park, 2300-2450 m, 3 Aug. 1992, P. Diederich 10616, 10676. (Herb. Diederich), H. Sipman 354632 (B). Madang Prov., S side of Ramu valley, Bundi village, on slopes towards Mt. Pizetara, 1300-1 600 m, 8 Nov. 1995, H. Sipman 392\0, 39368 (B). Northern Prov. , Owen Stanley Range, Myola, near guesthouse along lora river, 2\OOm, 14 Oct. 1995, H. Sipman 38172 (B). Simbu Prov. , Bundi Gap, c. 2800 m, 4 Aug. 1992, P. Diederich 1 \068 (Herb. Diederich), Serusiaux 13856-32 (LG), H. Sipman 35535 (B); Mt. Wilhelm, Pindaunde valley, near the hut on the shore of Lake Piunde, c. 3600 m, 9 Aug. 1992, P. Diederich 10274 (Herb. Diederich); Mt. Wilhelm area, 11 km on road under construction from Gembogl to Goroka, 9 Aug. 1992, P. Diederich 11264 (Herb. Diederich). Southern Highlands Prov., laro river, Onim, 14km NNW of lalibu, 2230m, 11 Sept. 1982, H. Streimann 23862 (CANB); Tari, Mt. Ambua, Kalkman 5198 (L). Western Highlands Prov., Mt. Giluwe, Hoffmann 87- 511 (LG).

Fig. 9. Parmeliella hawaiiensis, specimen from Borneo (TNS). 710 1. Hattori Bot. Lab. No. 100 2 0 0 6

Parmeliella hawaiiensis H. Magn. This little known species, which has hardly been reported since its description (Mag­ nusson 1955), is characterized by its neat, fiat, grey-blue rosettes, to 1.5 cm diam., with centrally located apothecia (Fig. 9), up to 0.5 mm broad with narrow distinct proper exci­ pie, which is excluded at maturity when the apothecia become convex. Habitat and distribution: A corticolous species of tree bases, probably a lowland ele­ ment just reaching the lower parts of the montane forests. P hawaiiensis is possibly a SE Asian element, reaching eastwards as far as Hawaii, and certainly not endemic to that is­ land. Specimens examined: PAPUA NEWGUINEA , Morobe Prov., 15 km SW of Bemard Camp, Idenburg River, 1600m, 1939, L. J. Brass (UPS); Spreader Divide, 12km NW of Aseki, 2000 m, 21 Jan. 1981 , H. Streimann & E. Tamba 11991 (CANB). USA, Hawaii, Femda1e, May 1909, Faurie 900 (UPS, ho1otype).

Parmeliella laceroides P. M. Jorg. & Sipman sp. novo (Fig. 10) Parmeliellae philippinae similis, sed squamulis non-adpressis, laceratis, in parte ad­ scendentis. Type: Papua New Guinea, Central Prov., Owen Stanley Range, Kagi village, along Kokoda trail towards Gap, 1700 m, 20/21 Oct. 1995, H. Sipman 38594 (B, ho1otype). Thallus grey-blue to brownish, spreading irregularly, to 5 cm diam., resting on a thin, crustose, blackish prothallus; individual squamules 1-3 mm diam., neatly incised, often lace-like with partly ascending lobes; in section C. \ 00 pm thick with \- 2 layered cortex, 20- 30 pm thick, the upper cells of which are clearly smaller than the lower. Apothecia 1- 2 mm diam., pale brown with a lighter, distinct, cellular proper margin; ascospores wart­ ed, ellipsoid, 15- 19 X 8- 10 mm. Chemistry: All reactions negative, no substances detected byTLC. When sterile, this species may remind one of a form of P polyphyllina, but it is thin-

Fig. 10. Parmeliella laceroides, holotype (B). P. M. J0 RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 711

Fig. 11. Parmeliella montana, holotype (B). ner and finer due to its neat, squamulose, laceroid thallus. It certainly belongs in the Parmeliella nigrocincta group as obvious from the apothecia. Habitat and distribution: So far only known by a few specimens from mossy and stunted montane forest at 1700-3100 m. Probably it represents a southern Malesian ele­ ment. Specimens examined: MALAYSIA, Sabah, Kota Belud distr., Kinabalu Park, S-slope of Mt. Kina­ balu, along summit trail, 2500-3100m, 9 May 1989, H. Sipman & B. Tan 30970, 31059 (B). PAPUA NEW GUINEA, Simbu Prov., Mt. Wilhelm area, Bundi Gap, on road Keglsugl-Bundi, c. 2800 m, 4 Aug. 1994, H. Sipman 35584 (B); c. 16 km on new road under construction from Gembogl to Goro­ ka, c. 2800 m, 9 Aug. 1992, H. Sipman 35928 (B).

Parmeliella montana P. M. J0rg. & Sipman sp. novo (Fig. 11) Parmeliellae marianae similis, sed thallo minore et prothallo crustaceo, sporis magnis. Type: PNG, Morobe Prov., Mt. Kaindi, Wau, 2100 m, 19 Dec. 1973, H. Kashiwadani 10541 (TNS, holotype). Thallus squamulose, brown; individual squamules 2-3 mm diam., to 100,um thick, usually with distinct, paler margins, resting on a thin, crustose, blackish prothallus. Apothecia frequent, round, 1- 2 mm diam. with distinct thalline margin and brown disc; spores simple, colourless, with distinct, smooth exospore, broadly ellipsoid, (15- )20- 26 X I 0-12 ,urn. Chemistry: All reactions negative, no substances detected by TLC. A species of the P mariana group, with a crustose prothallus and small squamules, reaching high altitudes. Habitat and distribution: Corticolous in montane forests from 2100 m to 3600 m and probably a Malesian element. Specimens examined: MALAYSIA, Borneo, Sabah, Kota Belud distr. , Kinabalu Park, S-slope of Mt. Kinabalu, along summit trail, c. 2900 m and 3600 m, 10 May 1989, H. Sipman & B. Tan 31030, 31049, 31198 (B). PAPUA NEW GUIN EA, Eastern Highlands Prov. , Mt. Gahavisuka Prov. Park, 11 km N 712 1. Hattori Bot. Lab. No. 100 200 6

Fig. 12. Parmeliella nitida, holotype (B). of Goroka, 2300 rn, 17 March 1987, H. Siprnan 22179 (B). Morobe Prov., Mt. Kaindi, 5 km W of Wau, 2360 rn, 13 March 1982, H. Streirnann 17625 (CANB); Mt. Sarawaket, southern Range, 4 km NEE of Lake Gwarn, 2850 rn , 4 July 1981 , T. Koponen 31580 CH). Sirnbu Prov., Mt. Wilhelrn, Pin­ daunde valley, near the hut on the shore of Lake Piunde, 3600 rn, 5 Aug. 1992, P. Diederich 10280, 1221 8 (Herb. Diederich). Southern Highlands Prov., Pavende Loggi ng Area, 15 km NNW of lalibu, 2400 rn , 14 Dec. 1982, 1. A. Elix & H. Streirnann 13084 (CAN B).

Parmeliella nitida P. M. J0rg. & Sipman sp. novo (Fig. 12) Parmeliellae polyphyllinae similis, sed lobis vulgo nitidis, coralloideis; ascosporae breviores et latiores, 15- 18 X 9-12 pm, perisporio bene evoluto, subverrucoso. Type: Papua New Guinea, Sirnbu Prov., Mt. Wilhelrn, Pindaunde valley, near hut on S-shore of Lake Piunde, W slope of valley, C. 3700 rn , 6 Aug. 1992, H. Siprnan 35688 (B, holotype). Thallus squamulose, dark, shiny, brown, to 5 cm diam., forming thick cushions; indi­ vidual squamules 2- 3 mm diam., to 150 pm wide, dentately incised, beset with gnarled, isidioid lobules, resting on a crustose, blackish prothallus. Apothecia round to irregularly lobed, to 2 mm diam., with strong, crenulate thalline margin; ascospores simple, colourless, broadly ellipsoid, 15- 18 X9- 12pm, with slightly warty exospore. Chemistry: All reactions negative, no substances detected by TLC. This species is closely related to P polyphyllina, but has a coralloid thallus, and more broadly ellipsoid spores, and it occurs at higher altitudes. Habitat and distribution: A corticolous species known only from the scrub along Lake Piunde in New Guinea and a similar site in Mt. Kinabalu on Borneo, probably a Malesian element. P. M. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 7 13

Fig. 13. Parmeliellapannarioides, holotype (TNS).

Specimens examined: MALAYSIA , Borneo, Sabah, distr. Ranau, Kinabalu Park, S-slope of Mt. Kinabalu, along Summit trail, c. 3200 m, 12 May 1989, H. Sipman & B. Tan 31242 (B). PAPUA NEW GUINEA, Simbu Prov., Mt. Wilhelm, near the hut on the shore of Lake Piunde, 3600 m, 5 Aug. 1992, P. Diederich 10051, 10235, 10272, 10276, 10281,10301 , 11234 (Herb. Diederich), H. Sipman 21974, 22062 (B).

Parmeliella pannarioides P. M. Jorg. & Sipman sp. novo (Fig. 13) Thallus fuscus, squamulosus, ad 2 cm latus, ad 400 flm crassus, rhizohyphis fuscis prominentibus. Apothecia ad 2 mm diam., excipulo thallino distincto, sporis ellipsoideis, rugolosis, 25- 30X9- 12flm. Type: Papua New Guinea, Western Highland distr., Mt. Wilhelm, en route from Kombugo­ manubo to the Pindaunde Lakes, 3400-3650 m, 3 Jan. 1974, H. Kashiwadani 11064 (TNS, holotype). Thallus squamulose, up to 2 cm diam., brown and thick, to 400 flm with cellular, 50-60 flm thick cortex, producing prominent brownish rhizohyphae from the naked lower surface, resting on a crustose, blackish prothallus. Apothecia round, to 2 mm diam., with flat, brown disc and distinct thalline margin; ascospores simple, colourless, rugulose, 25- 30X9- 12 flm. Chemistry: All reactions negative, no substances detected by TLC. This species is superficially very similar to a Pannaria (it has been called Pannaria obscura in Aptroot et al. 1997), particularly in the form with radiating, thick lobes. In a­ ddition to the hymenial characters and the lack of pannarin (PD-), it is recognizable by the distinctly brownish rhizohyphae, a character not found in any Pannaria. It also has a blackish-blue, crustose prothallus, which is visible at the margins of the rhizohyphae cushions. In its more small-squamulose forms it is more similar to a Psoroma from which it is readily distinguished by the cyanobiont and the amyloid hymenium. It is a unique species. Habitat and distribution: Growing on branches in dense scrub, and as yet only known 714 1. Hattori Bot. Lab. No. 100 2 0 0 6

Fig. 14. Parmeliella philippina, detail of Sip man 35926 (B).

from around Lake Piunde in PNG, at high altitudes. Possibly endemic, with no obvious close relatives. Specimens examined: PAPUA NEW GUI NEA , Simbu Prov. , Mt. Wilhelm, Pindaunde valley, near hut on the shore of Lake Piunde, 3600 m, 5 Aug. 1992, P. Diederich 10282, 10275, 10304, 11096 (Herb. Diederich), 12 March 1987, H. Sipman 21971 ; en route from Kombugomanubo to the Pin­ daunde Lakes, 3400-3650 m, 3 and 6 Jan. 1974, H. Kashiwadani 11475, 11481 (TNS); en route from Pindaunde Lakes to the summit of Mt. Wilhelm, 3600 m, 31 Dec. 1973, H. Kashiwadani 11363, 11387 (TNS).

Parmeliella philippina (Vain.) P. M . JeJrg. This species has long been mistaken for the subantarctic P nigrocincta from which it differs in having a small-squamulose (Fig. 14), adpressed thallus with secondary lobules, as well as by the longer spores, 16- 18 Jim. Habitat and distribution: A quite common and widespread, corticolous species of branches in montane forests, particularly common at c. 2500 m but reaching into the sub­ alpine scrub, as high as 3600 m. It is a southern Asian element. Specimens examined: INDONESIA , Java, Mt. Gedeh, 1916, W. Seiferth (0, W). MALAYSIA , Bor­ neo, Sabah, Ranau distr., Kinabalu Park, S-slope of Mt. Kinabalu, along summit trail, c. 3600 m, II May 1989, H. Sipman & B. Tan 31196 (B). PAPUA NEW GUI NE A, Madang Prov. , Huon Peninsula, Fin­ isterre Range, Yupna valley, Teptep village, trail in NNW and deep valley in N direction, 2500 m, 30 July 1992, P. Diederich 10885 (Herb. Diederich), H. Sipman 35377 (B). Morobe Prov., Aseki­ Menyamya road, 6km NW of Aseki, 1950m, 5 Dec. 1982,1. A. Elix & M. Toia 12110p.p. (CANB); Huon Peninsula, Saruwaged Range, Honzeukngnon village, S of Denim airstrip in Timbe valley, 2100 m, 7- 8 March 1987, H. Sipman 24454 (B); Mt. Kaindi, Wau, 2200 m, H. Kashiwadani 10579 (TNS). Northern Prov. , Owen Stanley Range, Myola, near guesthouse, along Iora River, 2100 m, 14 P. M. J0RGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 715

Fig. 15. Parmeliellapiundensis, holotype (B).

Oct. 1995, H. Sipman 38163 (B). Simbu Prov. , Mt. Wilhelm area, 16 km on new road under construc­ tion from Gembogl to Goroka, 2800 m, 9 Aug. 1992, H. Sipman 35926 (B); Bundi Gap on road Keglsugl-Bundi, 2800 m, 4 Aug. 1992, H. Sipman 35584 (B); Pindaunde valley, near hut on shore of Lake Piunde, 3600 m, 5 Aug. 1992, P. Diederich 10283, 10284 (Herb. Diederich). PHILIPPINES, Luzon, Benguet subprov., 1500m, May 1911, E. D. Merrill (TUR-VAIN 12345, holotype); Baguio, Mt. St~. Thomas, 2200 m, Febr. 1987, A. Aptroot 20349 (Herb. Aptroot).

Parmeliella piundensis P. M. J0rg. & Sipman sp. novo (Fig. IS) Parmeliellae nigrocinctae similis, sed lobis marginalibus radiato-elongatis, apotheciis maturis convexis. Type: PNG, Simbu Prov., Mt. Wilhelm, Pindaunde valley, near hut on S-shore of Lake Piunde, 3400 m, 6 Aug. 1992, H. Sipman 35742 (B, holotype). Thallus squamulose, forming colonies up to S cm diam.; individual squamules to 2 mm, grey-blue to grey-brown, appressed on a crustose, black prothallus, SO- lOO j1m thick, with single-layered, cellular upper cortex, the marginal ones enlarged and radiating, crowded in central parts and there with a tendency to form isidioid outgrowths. Apothecia numerous, in the centre of the thallus, often aggregated, proliferating, O.S- I mm diam., ini­ tially witH a broad proper exciple, becoming convex at maturity; ascospores simple, colour­ less, ellipsoid, faintly verrucose, IO- lSX4- S j1m . Chemistry: All reactions negative, no substances detected by TLC. An easily recognized species by the elongated marginal squamules and the numerous, small, convex apothecia with distinct proper margin. Habitat and distribution: Growing on branches of dense scrub near Lake Piunde in PNG at C. 3S00 m, and in a similar habitat on Mt. Kinabalu in Borneo. Possibly a Malesian taxon in the P nigrocincta-complex, and then with subantarctic affinity. Specimens studied: MALAYSIA, Borneo, Sabah, distr. Ranau, Kinabalu Park, S-slope of Mt. Kin­ abalu, along summit trail, C. 3600 m, II May 1989, H. Sipman & B. Tan 31184 (B). PAP UA NEW 716 1. Hattori Bot. Lab. No. 100 200 6

GUINEA, Simbu Prov., Mt. Wilhelm, en route from Kombugomanubo to Pindaunde Lakes, 3400- 3650 m, 3 Jan. 1974, H. Kashiwadani 11491 (TNS); Pindaunde valley, near hut on S-shore of Lake Piunde, on W-slope of the valley, 14 March 1987, H. Sipman 22090 (B).

Parmeliella polyphyllina P. M. ]0rg. For a description see ]0rgensen (200 I b). However, its variation is greater than that de­ termined from the few specimens then available. The lobules are sometimes finely incised and are not always ascending, and may be flatly imbricate, forming a layered thallus. In its typical form P polyphyllina is characterized by the numerous, fragile, erect lobules and may be mistaken for either Pannaria papuana, which has thinner, more bluish lobules, some of which contain a green photobiont, or Parmeliella nitida (see above for differences) found at higher altitudes. P laceroides may also have ascending lobules, but not as pro­ nounced as in P polyphyllina; when fertile it is always separable by the apothecia, which have a proper margin only and contain warty spores. Habitat and distribution: A corticolous species, mainly found on the trunks of trees in the montane forests, usually below 3000 m, which is otherwise known only from rain forests in Queensland, Australia, from where it was first described. It is possibly an Aus­ tralasian element. Specimens examined: MALAYSIA, Borneo, Sabah, di str. Kota Belud, Kinabalu Park, S-slope of Mt. Kinabalu, along Summit trail, c. 3100 m, 10 May 1989, H. Sipman & B. Tan 31066 (B). PAPUA NEW GUINEA: Central Prov. , K. B. Sawmill, Ehu Creek, 12 km SW of Sogeri, 750 m, 16 Febr. 198 I , H. Streimann & E. K. Naoni 16635 (CANB). Eastern Highlands Prov., Mt. Gahavisuka Prov. Park, I I km N of Goroka, along trail from parking place to orchid house, 2300 m, 3 Aug. 1992, H. Sipman 35462 (B). Madang Prov., Huon Peninsula, Finisterre range, Yupna valley, Teptep village, 2300 m, 30 July 1992, H. Sipman 35217 (B). Morobe Prov. , Aiuwa-Bakia track, 3km W of Aseki, 1500m, 22 Jan. 198 I , H. Streiman & E. Tamba (B); Angabena Ridge, 4 km NNE of Aseki, 1750 m, 1. A. Elix 12770 (CANB); Mt. Kaindi, 5 km W ofWau, 2350m, 13 March 1982, H. Streimann 17538 (CANB); Spreader Divide, 12km NW of Aseki, 2000m, 21 Jan. 198 1, H. Streimann & E. Tamba 11915 (CANB) ; Aseki-Menyamya road, 9km NW of Aseki, 2180m, 5 Dec. 1982, 1. A. Elix & M. Toia 12028 (CANB). Simbu Prov., Mt. Wilhelm area, c. I I km on new road under construction from Gem­ bogl to Goroka, 2800 m, 9 Aug. 1992, H. Sipman 35812 (B). Southern Highlands Prov., Onim forestry station, Taro River, 14 km NNW ofIalibu, 2280 m, 19 Dec. 1982, 1. A. Elix & H. Streimann 13427 (CANB); Tari-Komo road, 6 km N of Komo, 1480 m, 16 Dec. 1982, 1. A. Elix & H. Streimann 13253 (CANB). PHILIPPI NES, Luzon, Prov. Benguet, Mt. Santo Tomas, S of Baguio, along road from telecommunication station to Baguio, 19 Febr. 1987, H. Sipman 2 I 748 (B).

Psoroma Michx. This much misunderstood genus only comprises small-squamulose species, mostly without lichen acids, with asci with a distinct ring structure (]0rgensen 200 I b). In New Guinea there is only one species belonging to the genus so defined, which is newly de­ scribed herewith.

Psoromafilicicola P. M. ]0rg. & Sipman sp. novo (Fig. 16) Psoromatis tenue similis, sed thallo melius evoluto, squamulis corticatis, imbricatis, dentatis, sine acido lichenico; apothecia cupuliformia, ascosporae simplices, hyalinae, P. M. J0RGEN SEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 717

Fig. 16. Psoromafilicicola, holotype (H).

14- 18 x 10- 12 pm. Type : Papua New Guinea, Morobe Prov., Sarawaket southern Range, 4 km SW of Lake Gwam, 3350 m, 5 July 1981 , T. Koponen 31828 (H, holotype). Thallus squamulose, forming rosettes of imbricate, incised, slightly elongated, green to brownish squamules, to I mm long, c. 300 pm thick with cellular, to 50 pm thick upper cortex; cephalodia common, bluish, granular, containing Nostoc in clusters, forming hemisphaerical structures among the squamules, which contain Myrmecia. Apothecia cupuliform, to I mm diam., flattening with age, with brown disc and dentate thalline mar­ gin, densely tomentose below; ascospores simple, colourless, ovoid, 14-18 X 10-12 pm, with rugulose exospore. Pycnidia not observed. Chemistry: All reactions negative, no sub­ stances detected by TLC. This new species is closely related to P tenue Henssen, but differs in a number of characters, most remarkably in the cupuliform apothecia, which only flatten when old, and in the much better developed, corticated thallus which has no lichen substances. In this lat­ ter character it resembles the generitype, P hypnorum (Vahl) S.F. Gray, which, however, has larger apothecia with squamulose margins and larger, warted spores. Additionally P filici­ cola deviates markedly from both these species in its ecology (see below). Habitat and distribution: Presently known only from montane regions of PNG, at c. 3000 m, confined to stems of tree-ferns (Cyathea) in subalpine grasslands where the spongy trunks contain moisture most of the time, but probably not as much as the substrate for the other species of the genus, moist debris along cold brooks in arctic/subantarctic re­ gions. The unusually thick upper cortex may be an adaptation to the drier conditions. Specimens examined: PAPUA NEW GUINEA, Morobe Prov., Sarawaket, southern Range, 3 km E of Lake Gwam, 3500m, 8 July 1981 , T. Koponen 32466 (H); 4km NNE of Lake Gwam, 2850m, 4 July 1981 , T. Koponen 31581, 31584 (H). Northern Prov., Owen Stanley Range, Myo1a, c. 3 km NE of guesthouse in deep valley (frost hollow), 2700 m, 16 Oct. 1995, A. Aptroot 37648 (Herb. Aptroot). Simbu Prov., Bismarck Range, Mt. Wi1helm, 1mbuka ridge above Lake Aunde, 3500 m, 26 June 1968, W. Weber & McVean (COLO, CBG, UPS); Pindaunde valley, near the hut on the shore of Lake Piunde, 5 Aug. 1992, P. Diederich 10242 (Herb. Diederich); near Lake Piunde, 3500 m, March 1987, 718 1. Hattori Bot. Lab. No. 100 200 6

A. Aptroot 18591 (Herb. Aptroot), H. Sipman 22088 (B).

CONCLUSIONS The Pannariaceae of the New Guinea mountains comprise according to our studies 37 species in nine genera. Ten species appear to be endemic to the island, most of them in the genus Fuscopannaria. The fact that different collectors in the same locality have found different sets of species is an indication that some of these species grow in small niches and are easily over­ looked. Therefore their actual distribution is still uncertain and in need of further research. The results indicate, however, a close link between the Pannariaceae occurring on higher altitudes on Mt. Wilhelm in New Guinea and those of similar altitudes on the disjunct Mt. Kinabalu in Borneo, both among the highest mountains in the region (over 4000 m). The concentration of possible endemic species at Lake Piunde is probably an artefact resulting from the presence of a hut which provides the sole lodging possibility on the way up to Mt. Wilhelm. An evaluation of the distribution of these species indicates patterns, however, which mostly coincide with the vegetation zones. In the treeless upper region (c. 4000 m) there are few species, growing on the ground, namely the endemic Fuscopannaria cacuminum, and Pannaria papuana which is also found further down in the subalpine scrub. This scrub is species rich and contains the highest number of endemic species, just as the Andean paramo bush (Jorgensen & Arvidsson 2004, Sipman 1995) with which some surprisingly disjunctive species are shared, e.g., Erioderma gloriosum and Pannaria andina. This phy­ togeographical anomaly (Jorgensen & Sip man 2002a) is hard to explain because the North­ ern Andes mountains emerged in late Cretaceous times (Taylor 1995) so that these species must have made the long crossing over the Pacific against prevailing wind systems. Otherwise the representatives of the family are, with few exceptions, identical with, or closely related to, the of the neighbouring islands, often with an Australasian distri­ bution (e.g., Pannaria molkenboeri). The general impression is that of a paleotropical rela­ tionship. However, in high altitude habitats, there are mainly australlsubantarctic elements, such as Degelia, Fuscoderma and Psoroma, and also rarely a northern temperate one (Pan­ naria conoplea). At lower altitudes (c. 1250m), one gets glimpses of the lowland element (e.g., Parmeliellaflavida and P. philippina) restricted to the SE Asian forests, just like Eri­ oderma tomentosum Hue (Jorgensen & Sipman 2002a).

ACKNOWLEDGEMENTS During the preparation of this paper we have been assisted by Mr 1. Berge, Mrs A. Botnen, Mrs B. Ingvartsen and Dr T. Tonsberg to whom we are most thankful. We are also grateful to the directors and curators of the herbaria for promptly providing loans, particu­ larly to A. Aptroot, Soest and P. Diederich, Luxembourg who have put collections from their private herbaria to our disposal. H. S. is indebted to several persons who accompanied him in the field or arranged field trips, namely Andre Aptroot, Paul Diederich, Peter Lamb­ ley, Emanuel Serusiaux and Benito Tan, and to the Deutsche Forschungsgemeinschaft for P. M. JORGENSEN & H. SIPMAN: Lichen family Pannariaceae in the montane regions of New Guinea 719 financial support. Prof. M. R. D. Seaward kindly took care of linguistic corrections.

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