Taxonomic Studies on Australian Meteoriaceae (Musci). 2: the Genera Aerobryopsis, Barbella, Floribundaria, Meteoriopsis, Meteorium and Weymouthia
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Journ . Hallori BOI . Lab. No. 69: 277- 312 (Jan. /991) TAXONOMIC STUDIES ON AUSTRALIAN METEORIACEAE (MUSCI). 2: THE GENERA AEROBRYOPSIS, BARBELLA, FLORIBUNDARIA, METEORIOPSIS, METEORIUM AND WEYMOUTHIA. HEINAR STREIMANN 1 ABSTRA CT Excluding PapiJIaria, dealt with in a separate publication, a total of 6 genera with 8 species are recognised in Australia with no endemic species. Weymouthia cochlearifolia is omitted from the family and BarbefJa nilens is not accepted in that genus. Most of the taxa extend from southeast Asia to the Pacific Islands. All taxa are ill ustrated. INTRODUCTION The genera Aerobryopsis, Barbella, Floribundaria, Meteoriopsis, Meteorium, Pilotrichella and Weymouthia, as listed in Index Muscorum (Wijk et al. 1959- 1969), have not been critically examined previously. The genus Papillaria has already been revised (Streimann 1990). Noguchi (1976) in his revision of the east Asiatic species dealt with the above genera except for Pilotrichella and Weymouthia. Manue! (1977) published a world revision of the genus Meteoriopsis, citing only two Australian collections. With the exception of Weymouthia (Scott & Stone 1976), none of the Australian species have been reviewed. Streimann & Curnow (1989) deleted Chrysocladium phaeum (Mitt.) Fleisch. and Pilotrichella conferta Ren. & Card. from the Australian moss flora as these reports were erroneous, thus leaving 8 species in 6 genera (Table I) for Australia. The status of Barbella nitens will be discussed separately in Part 3. MORPHOLOGICAL STUDIES All collections illustrated are at CBG except for Ramsay (UNSW), Schofield (UBC), Stone (MELU) and Watts (NSW) . General morphology The genera studied have distinct primary and secondary stems and have long leaf tips, especially on the pendulous branches. Plants are generally dull to dark green, I Cryptogamic Herbarium, Australian National Botanic G ardens, GPO Box 1777, Canberra City, A.C.T ., 2601 , Australia. 278 Journ. Haltori Bot. Lab. No. 69 199 TAillE I. Meteoriaceae taxa (except Papillaria) reported for Australia prior to this revision with present status and distribution (*endemic). Previously recognized taxa Changed taxa Australian distribution Aerohryopsis wallichii (Brid.) Fleisch. QLD Barbella cuhellsis (Milt.) Broth. QLD, NSW, LHI B. nilellS (Hook. f. & Wils.) Nog. Not Barhel/a * B. perpinnata (Broth.) Broth . Brachyli1eciul11 ?sa/ebrosul11 (Web. & Mohr) B. S. G . F/orihundaria florihunda QLD (Dozy & Molk.) Fleisch. F. pscudof/oribunda Fleisch. QLD * F. rohuslu/a Broth. & Watts Meleoriopsis rec/inala F wa/keri (Ren. & Card .) Broth. QLD M eleoriopsis reclinaIa QLD (C Mucll.) Fleisch * Melcorium haileyi (Broth.) Broth. M. bllchananii M . buchallallii (Brid.) Broth. QLD *M. compressum Mitt. Barbel/a cllbensis * Pi/olrichella dil110rpha Barbel/a nilens (C Muel!.) Jaeg. * P. recurl'u/a C M lIcll . Ca/yploli1ecium recurl'u/um (Broth.) Broth. WeYl110ulh ia cochicari/olia Not Meteoriaceae (Schwaegr.) Dix. W. c. var. billardieri (Hpe) Di x. Not Meteoriaceae W. mo/lis (Hedw.) Broth. LHI, VIC, TAS but lighter in Floribundaria and Weymouthia. Stem and branch cross-sections Stems usually have a central strand (Fig. 1- 8), but in branches this is not so common, being more prominent nearer the stem and weaker towards the end of the branch. In Floribundaria species the cells became angular nearer the ends of branches. In mature stems and branches the outer cortex consists of golden yellow thick walled sclereids while the cells become more regular in shape and size towards the centre where cell walls are thinner. In Meteoriopsis reclinata the outer and inner cortical cells are bigger than in other species while the cortical cells of Floribundariajforibunda are all large and conspicuous. Rhizoids Rhizoids are clustered in leafaxils and arise from the stem or branches, but rarely from the leaf base (Barbel/a cubensis, King 777). They are smooth, reddish brown and very similar in all species. Rhizoid cell length varies with the shortest in Aerobryopsis (1l2 Ilm). Cells greater than 150llm are more prevalent, being observed in Barbella, Meteoriopsis and Meteorium. H. STREIMANN : Taxonomic studies on Australian Meteoriaceae. 2 279 FIG. 1- 8. Stem cross-sections: I Aerohryop.I'i.\' wallichii (HS 28975); 2 Barhella cubensis (HS 29677): 3 Floribundariafloribullda (Schojield 8(175); 4 F. pseudojlorihunda (Walls Q624pp); 5 F. walkeri (Watts Q579); 6 Meteoriopsis ree/illata (HS 29499); 7 MeteoriunI hucliallallii (HS 29680); 8 Weymouthia mol/is (TlIrnhlll/ H520). Sexuality Meteoriaceae are generally dioicous, except Barbelfa which is occasionally autoicous. Perigonia and perichaetia are rare, with the former appearing to be more frequent, but they were not found for Barbella and are extremely rare in Meteorium. The scarceness of sexual parts may account for the lack of sporophytes and asexual reproduction may be the main method of propagation. That may be the reason why some species are rare or do not form large colonies (e.g. Floribundaria spp.). Perigonia are rounded to shortly ovate and are found on the branches, but in larger leaved species they are covered by the leaves and consequently harder to find . The size and shape of perigonialleaves (Fig. 9- 16) show only slight variations, being between 1.0- 1.4 mm. The ratio of perigonialleaf length to antheridiallength is generally between 2.2 and 3.1. The shape of perigonial leaves is similar in all species, except those of Floribundaria pseudofioribunda which tend to be narrower. Generally periogonial leaves are ecostate and without papillae except in Aerobryopsis and 280 Journ. Hattori Bot. Lab. No. 69 199 I FIG. 9- 16. Perigonial leaves: 9 Aerobryopsis lI'allichii (Schofield 80052); 10, 11 Floribundaria pseudojloribunda (Walls Q655, Q624pp); 12 Meteoriopsis reciinata (Stone 15193); 13 ,14 Meteorium buchananii (HS 29680); 15 , 16 Weymouthia mol/is (Turnbull H 520). 200.um FIG. 17- 22. Antheridia: 17 Aerobryopsis wallichii (Schofield 80052); 18 Floribundaria pseudojloribunda (Watts Q655); 19 Meteoriopsis reciinata (Stone 15193); 20 Meleoriul11 buchananii (HS 29680); 21 , 22 Weymouthia mol/is (Turnbull H520). H. STREIMANN: Taxonomic studies on Australian Meteoriaceae. 2 28 1 Weymouthia where a few faint papillae may be present. Leaves closest to the antheridia are not papillose, but those further away tend to develop papillosity. Margins are entire and a border one cell wide of longer narrower cells was noted in all species. Some perigonial leaves in Weymouthia had a band of yellowish brown cells across the base. The number of antheridia per perigonium varies and is not consistent within the genera. Meleorium (21) had the most and Floribundaria pseudofloribunda (6) the least. Antheridial length (Fig. 17-22) was commonly between 450-600llm, except in F. pseudofloribunda (380 Ilm) and in Weymouthia (410 Ilm). All antherdia have the normal club shape with some being slightly curved. All species have a short stout stalk except Weymouthia where a thin stalk to 30llm long (Fig. 22) was noted. Antheridial size related to the size of the plant. Paraphyses are the same number as antheridia or numerous in all species except in Floribundaria pseudofloribunda which had few. Width varied from 171lm (Aerobryopsis) to 251lm (Floribundaria pseudofloribunda), while cell length varied between 80-136Ilm, again the extremes are for the same species. Perichaetia are rare, ovate to elongate on branches and often project from between FIG. 23- 35. Perichaetial leaves: 23 Aerobryopsis wallichii (Kanlak 953); 24 Barbella cubensis (Walls LH! 42/); 25, 26 Floribundaria .floribunda (Schofield 80/75); 27- 3 1 Meleoriopsis rec/inala (Wails Q541 A, Q54! A, Q669, Q669, Q541 A); 32, 33 Meleorium buchananii (HS 29498); 34, 35 Weymouthia mollis (Curnow 1368). 282 Journ. Haltori Bot. Lab. No. 69 1 9 9 1 the leaves. except in Barhel/a where they align with the stem and are very similar to the leaf buds. Perichaetial leaves (Fig. 23- 35) are generall y longer than perigonialleaves (except in Weymoulhia where they are equal) and have a greater range of shapes and sizes. The longest are in Meleorium (2.4 mm), and the shortest in Weymoulhia (1.2 mm). The leaf widths show very little variation between the species. Ratio of perigonial leaf length/archegonial length is more variable than perigonial leaflength/antheridiallength and could be separated into two d istinct categories: A ratio> 4.3: Meleoriopsis (4.3), M eleorium (4.4), Floribundaria floribunda (5 .0). B ratio < 3.5: Weymouthia (2.9), Barbel/a (3.0), Aerobryopsis (3 .4). The inner perichaetial leaves are of two kinds in Meleoriopsis (Fig. 28, 30), and possibly also in Barbel/a, being very long and narrow, distantly sharply serrated, especially near the apex, but otherwise similar. In Aerobryopsis cell s near the base are distinctly porose, especially so the small er outer leaves. The margins are entire in all species, but are rarely denticulate in Floribundaria pseudofloribunda. As with the perigonial leaves papillae are lacking, but those further from the archegonia tend to develop papill osity. Again all leaves are ecostate with a margin of narrower rectangular cells. The number of archegonia per bud varied from 9 in Me/eorium to 24 in Aerobryopsis. Archegonial length (Fig. 36-43) varies from 300.um in Floribundaria floribunda to 550.um in Me/eorium. Shape shows very little variation, with Floribundaria floribunda and Weymouthia having the narrowest (0.3.um) archegonia and Meleorium having the widest (0.65 .um). Archegonia in all species have a long stout stalk. 200.urn .- 39 41 37 38 40 42 43 FIG. 36-43. Archegonia: 36 Aerohry psis l1'al/ichii (Kalltak 953): 37. 38 Barhel/a cuhensis (O/iI'er NSW MI83259): 39 Floribundaria .tforibunda (Sc/t(Jfield 801 75) : 40, 41 M eleoriopsis reclillala (WallS Q669): 42 Meleorium huchananii (HS 29498); 43 Weymoulhia m ol/is (Curn OlI' 1368). H . STREIMANN: Taxonomic studies on Australian Meteoriaceae. 2 283 TAB LE 2. Reported chromosome numbers in Meteoriaceae (from F ritsch (982). - ----- Chrom. No. Species Author (n) Aerohryopsis suhdivergells (Broth.) Broth. I1 Inoue & Momii 1971 Barhella(lagellifera (Card.) Nog. 10 Inoue 1965 [as B.