J Hattori Bot. Lab. No. 91: 289-294 (Nov. 2001)

WEYMOUTHIA COCHLEAR/FOLIA (SCHWAGR.) DIX. (-MUSCI) IN AUSTRALIA, AND SOME NOTES ON W MOLLIS (HEDW) BROTH.

HEINAR STREIMANNi t

ABSTRACT. Weymouthia cochlearifolia is revised and illustrated. W billardieri (Hampe) Broth. is a new synonym. New synonyms, Meteorium molle Wils. var. majus Wils. and Pilotrichella mollis (Hedw.) Jaeg. var. major (Wils.) Jaeg., are reported for W mollis. Altitude-latitude figures are pre­ sented for both species together with a discussion of their distributions and habitat preferences.

When the genera of the were revised for Australia doubts were ex­ pressed (Streimann 1991) about the position of the genus Weymouthia in the Meteoriaceae. Weymouthia mollis was retained in the Meteoriaceae with reservations, while W cochleari­ folia was omitted on the basis that the characters of this species were more akin to those of the Lembophyllaceae than to the Meteoriaceae. Subsequently Tangney (1997), transferred Weymouthia to the Lembophyllaceae in his generic revision of this family. Thus to com­ plete the revision of the genus for Australia the study of Weymouthia cochlearifolia is pre­ sented here.

Key to Australian species of Weymouthia (after Scott & Stone 1976) 1. firm; leaves sub-orbicular, very concave, wrinkled when dry...... (I) W cochlearifolia Plants soft; leaves oblong-spathulate, predominantly U-channelled, not wrinkled ... (2) W mollis

Weymouthia cochlearifolia (Schwagr.) Dix., New Zealand Inst. Bull. 3: 255. 1927. Hypnum cochlearifolium Schwiigr., Spec. Musc. Suppl. 1(2): 221. 1816. Type: 'In Australasia lectum communicavit cl. La Billardiere'. Holotype: G. lsothecium cochlearifolium (Schwiigr.) Mitt., Hooker's 1. Bot. Kew Gard. Misc. 8: 264. 1856. Stereodon cochlearifolia (Schwiigr.) Mitt., 1. Linn. Soc., Bot. 4: 88. 1860. Coelidium cochlearifolium (Schwiigr.) Rchdt, Reise Osterr. Freg. Novara Bot. 1(3): 191. 1870. Porotrichum cochlearifolium (Schwiigr.) Mitt., Trans. & Proc. Roy. Soc. Victoria 19: 84.1882. Lembophyllum cochlearifolium (Schwiigr.) Lindb., Ind. Bryol. 718. 1897. Hypnum cochlearifolium var. minus Hampe, Linnaea 25: 715. 1853. Type; 'Van Diemensland (Stu­ art). Lembophyllum cochlearifolium var. minus (Hampe) Par., lnd. Bryol. 718.1896. Meteorium molle Wils. var. majus Bastow, Pap. & Proc. Roy. Soc. Tasm. 1886: 81. 1887. Type: 'Brown's River' . Neckera billardieri Hampe, Linnaea 30: 637. 1860, syn. novo Type: 'Apollo Bay, F. Mueller 1857'. Holotype: BM!; Isotypes: BM!, MEL (32257)! Pilotrichella billardieri (Hampe) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76: 259.1877, syn. novo Meteorium billardieri

1 Estonian Natural History Museum, Kopli St 76, Tallinn, EE 004, Estonia. t Heinar Streimann died on 29 August 2001. 290 1. Hattori Bot. Lab. No. 91 200 1

(Hampe) Mitt., Trans. & Proc. Roy. Soc. Victoria 19: 82. 1882, syn. novo Weymouthia billardieri (Ham pe) Broth., Nat. PfI. 1(3): 812. 1906, syn. novo W cochlearifolia (Schwagr.) Dix. var. bil­ lardieri (Hampe) Dix., New Zealand Inst. Bull. 3: 255 . 1927, syn. novo

Plants generally robust, rarely slender, loosely tufted, creeping or pendulous, yellow­ brown, yellow-green or pale green, often ageing to golden-reddish colour, glossy. Stems flexuose, rarely filiform. Branches often pinnate, rigid, freely branching with long flexuose branches or short, tips with spreading leaves, leaf size decreasing to branch apices which gives a flagellate appearance. Branch leaves tightly imbricate, tumid, usually wrinkled or terete, sub-erect when dry or moist, sub-orbicular, cochleariform, very concave, to 1.0-1.3 X 0.6- 2.0 mm, apex cucullate with hooded appearance, base wide, margin entire, or apex generally weakly denticulate or crenulate, costa absent or very faint. Cells smooth, apical cells variable, to 35 X 5 ,urn, incrassate, median cells linear-sigmoid with pointed ends, (25)30- 80 X 4- 6 ,urn , cell size decreasing, width increasing nearer leaf base, incras­ sate, porose; alar groups small, conspicuous, generally with reddish or yellow tinge or colourless, cells quadrate or rectangular to 25 X 12 ,urn, with granular content, very incras­ sate; basal marginal cells above alar cells smaller, narrower, not forming border. Stem leaves to 2.1 X 1.8 mm, often wider than long, denticulation at apex very weak or none, cells similar to branch leaves. Rhizoids very weakly papillose, 13,um wide, cells to 175,um long, margins thickened, ends sometimes overlapping. Perigonia not seen. Perichaetia elongated, bud-like, scattered on branches. Inner perichaetial leaves ovate, to 0.90 X0.45 mm, shortly acuminate, margins entire; ecostate. Papillae absent, upper and median cells to 50 X 3 ,urn, incrassate, +sigmoid, basal cells vari­ ably rectangular ( + 25 X 7 ,urn), porose, reddish at base. Archegonia + 16 per bud, reddish­ brown, to 450X40 ,urn, neck to 20,um wide, cells irregularly rectangular. Paraphyses to 900,um long, 13,um (1 cell) wide, cells rectangular, thick-walled, to 63,um long. Inner perichaetial leaves at base of capsule similar, but lanceolate-ovate, to 2.7 X 1.0 mm, acumi­ nate, upper and median cells to 63 X4 ,urn, more incrassate. Paraphyses similar to those sur­ rounding archegonia but l5,um wide, cells to 100,um long. Capsules brown oblong to oval, rarely asymmetrical, to I.7 XO.8mm; setae at bases of branches, very long, to l5.6mm long, slight anti-clockwise twist. Operculum acutely or obtusely conical, about half capsule length. Peristome yellow-brown, teeth to 400,um long, transversely striolate to 240,um from base; margin of striolate section with semi-circular indentation corresponding with each zig-zag. Processes keeled. Cilia 2, nodulose, shorter than processes. Spores 20- 25 ,urn . Chromosome number: no reports. Illustrations: Fig. I; Allison & Child 1971: 108 & pI. 25; Beever et al. 1992: 119; Brotherus 1906: 866, 1925: 206 (Lembophyllum cochlearifolium); Dixon 1927: 373; Schwagrichen 1816: fig . 88. Ecology: Habit, habitat and altitudinal data are lacking for many older collections, but it appears that Weymouthia cochlearifolia prefers cool temperate and Nothofagus forests where the most common substratum is tree trunks (Nothofagus and Lagarostrobus). This species has also been reported on logs and wood. The altitudinal range (Fig. 3) ap­ pears to be from sea-level to 750 m in Tasmania while in eastern Gippsland in Victoria it H. STREIMANN: Weymouthia cochlearifolia in Australia 291 has been reported from the lowlands to 1280 m. Distribution: Southeast Australia: - Victoria and Tasmania (Fig. 2). It also occurs on Lord Howe Island, in New Zealand, and Chile. It possibly occurs in mainland New South Wales; but collections at CANB could not be checked, nor obtained on loan. Types: The type of Weymouthia cochlearifolia could not be located. The type of Neckera billardieri (BM) was examined and proved to be within the range for variation ob­ served for Weymouthia cochlearifolia. Sainsbury (1955) found that the characters Dixon (1927) used to delineate this variety were unsatisfactory: 'longer, more flexuose stems; shorter setae; and almost globose capsules'. However, these characters show considerable intergrading throughout a single . Dixon (1927) showed very clearly the weaknesses in attempting to segregate this species and its variety and it is surprising that he decided to retain them as separate entities. Australian collections studied exhibited an analogous de­ gree of variation. Perpetuation of this confusion could perhaps be explained by the paucity of collections and in the rarity of fertile material. Notes: Specimens of Weymouthia cochlearifolia have often been mis-determined as Lembophyllum divulsum causing considerable confusion. The main differences in these to taxa are outlined below. Weymouthia cochlearifolia Lembophyllum divulsum I) Cells long 40-80 X6.urn with tapering Cells short 20 X6.urn rhornboidal & variable sigmoid ends 2) Costa absent, faint or double, generally Costa commonly wide at base, forked, generally les than 114 ofleaf length 112 of leaflength 3) Leaf base very obtuse, wide Leaves narrowed at insertion forming a waist 4) Cell ends porose throughout leaf Cells porose only at base, rarely at tip 5) Alar cells short, square or oblong Alar cells rounded 6) Leaf tip pouch-like Leaf tip often pouch-like 7) Branch leaf size variable branches Branch leaves even sized throughout often flagelliform 8) On tree trunks On rocks, soil, rarely on trees

In Scott & Stone (1976) descriptions for the leaf tips of these two species are de­ scribed as being different: Weymouthia - 'entire or at most slightly crenulate'; Lembophyllum -'crenulate-denticulate'. However, this was found to be variable and not a reliable character. Weymouthia cochlearifolia has also been confused with Acrocladium chlamydophyl­ lum, but the latter species has pointed shoots, flatter leaves and large inflated empty hyaline cells in the auricles. Both Lembophyllum and Acrocladium usually occur at the base of trees, or on the ground, while Weymouthia is generally found on tree trunks. Weymouthia cochlearifolia can develop filiform tips to the branches where the leaves are very small and widely spaced (F.v. Mueller-MEL 32264 & J.H. Willis-MEL 1034367). Larger leaves on these branches are similar appearance to those of W. mollis. Perigonia and perichaetia were extremely hard to find because the rounded branch leaves completely covered these organs. It appears that sporophytes mature between August and October. 292 1. Hattori Bot. Lab. No. 91 200 1

Wijk et al. (1969) list W cochlearifolia as occurring in Asia (not confirmed by Noguchi 1976), and for the Pacific - Oceania (not substantiated by Miller et al. 1978). Brotherus (1906, 1925) does not list Weymouthia for Hawaii, but Sainsbury (1955) stated that the genus occurs in the Sandwich Islands (Hawaii). No doubt he based this on the work of Brotherus, where W desmoclada (c. Miill.) Broth., was reported for Hawaii, but is now considered conspecific with the Hawaiian endemic Pilotrichella mauiensis (Sull.) Jaeg. Etymology: from the Latin, cochlea = spoon shaped andfoliatus=leaves, referring to the rounded cucullate leaves. Specimens studied: 93 . NEW SOUTH WALES: Lord Howe Island: Mt Gower Summit, 820m, low vegetation dominated by Metrosideros nervulosa, Zygogynum howeanum, Dysoxylum pachyphyllum, Dracophyllum, tree ferns and palms, on shaded treelet (Dysoxylum) stem, H. Streimann 56146 (NY), 11 Feb 1995; VICTORIA: Rooty Break Track, Goonmirk Rocks Rd, Scenic Reserve, 1150m, dense mountain forest, epiphyte, A. W Thies FN 1471M (MEL), 28 Mar. 1986; Mt Ellery, E. Merrah (BM, NY), 1887; Croajingolong National Park, Camp Ck, S of Cicada Trail, A. C Beauglehole 31993 (MEL), 22 Nov. 1969; Mt Drummer, R. Melville & NA. Wakefield 28320 (MEL, NSW), 12 Jan. 1953; Tarwin, F.v. Muel/er 122 (BM, NY), no date; Sealers Cove, F. v. Muel/er 112 (BM, NY), no date; Ettas Glen, Black Spur, near Healesville, W W Watts V436 (NSW), 21 Dec. 1886; Cement Ck, Warburton, N A. Hansen (MEL 38462), Jan. 1947; Junction Paradise Road and Paradise Track, Wombat State Forest, 20 km SSE of Daylesford, 650 m, disturbed wet sclerophyll forest dominated by Bedfordia, on Bedfordia stem, H. Streimann 39021 (NY, B, HO, PRE, Mahu) 16 Nov 1987; Skenes Ck, Apollo Bay, Mott (MEL 32258), 29 Oct. 1891 ; Boundary Ck, Yeodene, SE of Co lac, A.C Beauglehole 1963 (MEL), Nov. 1949; Turtons Pass, 13 km E of Beech Forest, 430 m, temperate rain forest, on tree trunk, H. Streimann 2428 (CANB, H, L, MO, NICH), 24 Aug. 1975. TASMAN IA : Ulver­ stone, Ladder (MEL 32279), Mar. 1886; Denison Gorge, Scottsdale railway line, W A. Weymouth 1898 (NY), 8 Feb. 1894; Hellyer R., H. P Ramsay 1860 (UNSW), 8 Dec. 1981 ; Goulds Country, A. W Robinson s. n. (as Bastow 385 - MEL, NSW), Aug. 1886; Lyell Highway, shaded log, wet sclero­ phyll forest, D. H. Norris 33430 (COLO, NY), 31 Jan. 1974; Philip Is ., Macquarie Harbour, dense forest, Anon. 1315 (NY), 1 Nov. 1946; Gordon R., forest, Milligan 785 (NY), no date; Mt Field Na­ tional Park, Nothofagus forest, D. H. Vitt 29129 (ALTA), 6 Dec. 1981 ; Sandy Bay, W A. Weymouth 2363 (NY), 22 Oct. 1910.

Weymouthia mollis (Hedw.) Broth., Nat. Pfianzenfam. 1: 812.1906. After the review of this species (Streimann 1991) it became apparent that the follow­ ing taxa were also synonymous. As not all type collections were located, no lectotype was designated. Meteorium molle Wils. var. majus Wils., Fl. Tasm. 2: 203 . 1859. Type: BM. 'In dense woods, pendent from the branches of trees. (J.D.H.) (Gunn, 1602!, 40!) (Lyal/ 102!) Brown's River Gully. (Old­ field, 69!) Cheshunt, Archer!.' Pilotrichella mol/is (Hedw.) Jaeg. var. major (Wils.) Jaeg., Ber. Tatigk. St. Gallischen Naturwiss. Ges. 1875-76: 260. 1877. Bastow (1887: 81) refers to 'Stereodon molle Mitt., var. B. majus', which has tumid branches. Dixon (1927: 255) placed this taxon as Meteorium molle var. majus Bastow into synonymy with Weymouthia cochlearifolia. However, Wijk et al. (1964: 356) placed Mete­ orium molle var. majus Wils. as a synonym of Weymouthia cochlearifolia citing Dixon's 1927 reference. That was not correct as the var. majus Bastow is a robust with tumid leaves and the var. majus Wils. has slender stems. Therefore Meteorium molle var. majus H. STREIMANN: Weymouthia cochlearifolia in Australia 293

Wils. is a synonym of Weymouthia mollis while, while Meteorium molle var. majus Bastow is a synonym of Weymouthia cochlearifolia.

Habitat preferences of Weymouthia This genus is very common in southern Victoria and Tasmania (Fig. 3), particularly in the Nothofagus and Atherosperma dominated (cool temperate) forests. Weymouthia mol/is often forms dense, delicate trailing masses which have the appearance of Papillaria colonies. This latter genus is more common further north in similar, but warmer habitats. Weymouthia mollis appears to be more common than W cochlearifolia and forming larger colonies. Both species occur at sea-level in Tasmania and both occur at higher altitudes in Victoria than in Tasmania. W mollis has rarely been found in New South Wales, but so far W cochlearifolia has not. In southern New South Wales there are several areas of cool tem­ perate forest where W cochlearifolia may be found, but they have not been investigated. Weymollthia mollis is quite common in areas receiving diffuse light in gaps or at the edge of the forest. As the vegetation becomes denser then the species is more common in the tree crowns, especially on the smaller branches.

ACKNOWLEDGEMENTS My deep gratitude is extended to Dr H.P. Ramsay for all the encouragement and assis­ tance she has given me in Australian bryology. The assistance given by I. Gleadhill, then the librarian at the Australian National Botanic Gardens, was greatly appreciated, especial­ ly in locating obscure Australian and European literature. Dr H.P. Ramsay, Prof. lA. Elix and E.M. Canning offred valuable comments on the text. Often I had to seek reprints from overseas and here Drs WR. Buck (NY), T. Koponen (H), and B. van Zanten (GRO) were always ready to assist promptly. I wish to thank the curators of the following herbaria BM, H, JE, L, H, HO, MEL, NSW, NY, S for their assistance in promptly providing loans of col­ lections. Help given by C. Hilliker, of the Australian National University; in the prepara­ tion of the maps and figures is also gratefully acknowledged.

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