J Hattori Bot. Lab. No. 75: 179- 182 (Feb. 1994)

A NEW MASTIGOPHORA (HEPATOPHYTA) FROM

DONNA HALL MILLER AND HARVEY A. MILLER 1

ABSTRACT. Mastigophora tuberculata sp. nov. from the Carstensz Mountains differs from M. dicla­ dos in having tuberculate leaf cells and tufted or branched paraphyllia.

The genus Mastigophora was described by Nees in 1838 to accommodate Jungerman­ nia woodsii Hooker. Subsequently, species now included in several genera, mostly in Her­ bertaceae and Lepidoziaceae, were placed in Mastigophora. Modern studies of the genus (e.g., Inoue, 1971; Piippo, 1984) have resulted in removal of the discordant elements and have demonstrated the potential plasticity within the genus, particularly for the widely dis­ tributed Malesian-Pacific species M. diclados. Four species of Mastigophora were recognized by Inoue (1971) with only a single species reported for New Guinea, the widely distributed and variable M. diclados. Howev­ er, he excluded four "species" described from New Guinea which were assigned to Mastigophora as new by Stephani (1900- 24). The excluded taxa, all collected by Leder­ mann and with specimens thought to be lost, are: M. appendiculata Steph. 1922. Spee. Hep. 6: 368. M. guineensis Steph. 1923. Spee. Hep. 6: 369. M. pyramidana Steph. 1923. Spee. Hep. 6: 369. M. valida Steph. 1923. Spee. Hep. 6: 369. Piippo (1984) reviewed the genus for New Guinea including the descriptions provided by Stephani and agreed with Inoue (1971) that too little detail is provided to arrive at a sat­ isfactory conclusion about their identity. Neither mentions a review, in the absence of spec­ imens, of the Stephani manuscript "lcones" where Stephani often provided clues to his de­ scriptions which Verdoorn (in litt.) noted were often based on the drawings. From the draw­ ings and the descriptions, M. guineensis and M. valida would seem to fall within the range of variability of M. diclados and we so consider them. On the other hand, M. appendiculata and M. pyramidana appear to be discordant elements in this genus as we understand it, but we do not venture a guess to their proper placement. The new species was collected in the vicinity of Punjak Jaya (Carstensz Peak) in the tropicalpine zone in a grassland with scattered small trees and shrubs growing in a peaty soil. Bryophytes formed large masses on branches of trees and shrubs as well as creating deep polsters around their bases. The Mastigophora was collected in admixture with other bryophytes. At first glance, the color and habit are so much like Mastigophora diclados that little attention was paid to it until sorting of the mass collections was begun in the laborato­ ry. We noticed that a specimen from valley in the headwaters of the Sungai Tsing River (Hope et al., 1976) looked somewhat different from M. diclados in that the leaf

1 Department of Biology, University of Central Florida, Orlando, Florida 32816, US.A. 180 J. Hattori Bot · La b ·No. 75 D. H. MILLER & H. A. MILLER : A new Mastigophora (Hepatophyta) from New Guniea 181 surface appeared to be roughened when viewed at 20 X or greater magnification. The dor­ sal surface of each leaf cell has a prominent dome over the whole lumen unlike other Mastigophora species. Another feature which is unlike anything we have seen in M. dic/a­ dos is the presence of tufted or branched paraphyllia on the ventral surface of the primary axis.

Mastigophora tuberculata sp. nov. Planta major, rufo brunnea plus minusve fusca vel nigricans caespitosa. Caulis ad plus minusve l 0 cm longus parum divisus crassus durus, grandifolius, ramis primariis remotius­ culus parvifoliis, longe paucipinnatis, pinnis minutifoliis plus minus longis capillaceis. Folia caulina approximata vel imbricata, parum incuba non transverse inserta, concava caulique appressa apicibus erectiusculis, ad medium triloba vel interdum quadriloba, lobis in piano porrectis sinu angusto obtuso discretis, inaequalibus, acutis, antico majore, basi antica calcarata. Folia ramulina minora, biloba, lobis subaequalibus, basi similiter appen­ diculatus. Cellulae superae 20--32 µm, basales 25-40 X 48- 80 µm, parietibus crassis nodu­ lose trabeculatis, papillis dorsalibus conspicue tuberculatis vel mammosis. Amphigastria caulina approximata vel imbricata appressa oblonga, basi calcarata, infra medium bifido, laciniis anguste lanceolatis acutis vel obtusis. Amphigastria ramulina bifida, laciniis angus­ tioribus. Paraphyllia fasciculis ventralibus, Iaciniis gracilibus interdum ramosis. Reliqua desunt. Specimen typicum in valle Ngga Pulu ad flumen Sungai Tsing, alt. 3320 m.s.m., Irian Jaya, in­ sula New Guinea, H. A. Miller 22107, I Feb. 1991 lecto (specimen typicum in hb MU conservatum). Plants in deep polsters near the bases of trees, or shrubs or grass hummocks from the upper montane and subalpine forest zone to the lower alpine zone. Leafy plants, deep red­ dish, blackish reddish to blackish reddish brown above becoming brownish in the older parts deep in the cushion. Stems mostly 4- 10 cm or more long, glossy reddish black dorsal­ ly and paraphyllose below, mostly bipinnately branched with an occasional single or bipin­ nate lateral branch arising at nearly right angles to the primary axis and slightly secund to­ wards the ventral side; branches attenuate towards the tip and frequently microphyllous and flagelliferous at the end and with a few distal rhizoids. Paraphyllia nearly lacking dorsally but numerous in small clumps of2-several laciniae on the ventral side of the stem; the indi­ vidual laciniae simple or toothed or branched. Leaves on the main axis usually three-lobed or with an occasional reduced fourth lobe; imbricate, semi-amplexicaul, weakly incubous to nearly transverse, concave, roundish-quadrate, divided two-thirds to three quarters or more, sinus rounded; the antical segment largest, erecto-patent, cordate to reniform below

Fig. 1. Mastigophora tuberculata D. H. and H. A. Miller A. Habit of the plant. B. Leaves from the main axis. C. Underleaf from the main axis. D. Branch leaves. E. Branch un­ derleaves. F. Paraphyllia tufts. G. Stem cross-section with an emerging paraphyllial tuft. H. Cellular detail from a single paraphyllium outlined immediately below. I. Cellular detail of complete leaf segment tips. J. Leaf segment margin. K. Cellular detail of leaf base near the middle of the largest segment as viewed from a stem cross-section. L. Leaf cross section. Scale bars: A. I 0 mm; B- E. 0.5 mm; F. 0.1 mm; G- L. 0.05 mm. All figures drawn from the type. 182 J. Hattori Bot. Lab. No. 75 I 9 9 4 becoming lanceolate in the upper half, margin of the basal disc involute becoming both in­ volute and incurved in the lanceolate portion of the segment; the next two segments nar­ rower, ovate-lanceolate, acute to acuminate with a tendency towards becoming cucullate distally; the leaf insertion long with the antical line merging to the stem but with the posti­ cal line of insertion, numerous paraphyllia-like appendages are found on the lower margin forming a laciniate tuft or a series of irregular spinosedenate teeth at the ventral angle. Leaf margins of segments often irregular and involute and mostly mamillate crenulate by virtue of the bulging, rounded gibbouspapillate, blade cells. Branch leaves generally smaller, mostly bifid with appendages or an extra vestigial lobe (or two), with areolation similar to that of primary stem leaves. Underleaves about half as long as the leaves, erecto-patent, the lobe margins plane to incurved, bifid half or more, sinus rounded, lobes ovate-lanceolate, acute to short acuminate, the margin crenulate by bulging cells, the underleaf base shortly recurved and appendiculate with one to several irregularly branched or curved laciniae. Leaf cells along the margin of segments rounded, nearly isodiamteric, 18- 28 µ.m in diame­ ter; segment tip region cells rounded and 20-32 µm with some elongated cells below the tip up to 30 X 40 µm; median cells of the leaf segments oval to rounded, 25- 35 µm in di­ ameter; median basal leaf cells 25-40 X 48-80 µ.m becoming shorter distally; cell lumens more or less stellate because of pits through the very thick yellowish cell wall; cells bulging to form a conspicuous tuberculate to mamillate papillate dorsal surface on the leaf. Androecia and gynoecia not seen. Specimens examined: New Guinea, Irian Jaya, Western Sudirman Range, Carstensz Mountains, valley east from Ngga Pulu peak, alpine meadow, altitude 3320 m.s.m. Harvey A. Miller 22107, I February 1991 (typus hb MU). Irian Jaya, Western Sudirman Range, south ridge from Gn Platen Spitz, 4 km south of , subalpine rainforest, altitude ea. 3050 m.s.m. Harvey A. Miller 22709,22724,22725,22740,22746, 12August 1992.

LITERATURE CITED Hope, G. S., J. A. Peterson, l. Allison, & U. Radok (eds.) 1976. The Equatorial Glaciers of New Guinea, Results of the 1971- 73 Australian Universities' Expeditions to Irian Jaya: Survey, Glaciology, Meteorology, Biology and Palaeoenvironments. A. A. Balkema, Rotterdam. Pp. i-xii; 1- 244; 3 maps. Inoue, H. 1971. Memoir of the genus Mastigophora Nees. Bull. Nat. Sci. Mus. Tokyo 14(4): 603-608. Piippo, S. 1984. Bryophyte flora of the Huon Peninsula, New Guinea. III. Ann. Bot. Fennici 21 : 21-48. Stephani, F. 1900-1924. Species Hepaticarum. 6 vols. Geneva. Stephani, F. (undated). /cones lneditae. Manuscript drawings traced and distributed by Johanna Stephani; originals at hb G.