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Characterization of Myhogramme Lnida, Myriogrammeae Trib

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Characterization of Myhogramme Lnida, Myriogrammeae Trib J. I’hyrol. 33, 106-121 (1997) CHARACTERIZATION OF MYHOGRAMME LNIDA, MYRIOGRAMMEAE TRIB. NOV (DELESSEIUACEAE, FWODOPHYTA) Max H. Hommersand‘ and Suzanne Fredericf Department of Biology, University of North Carolina, Chapel Hill, North Carolina 27599-3280 ABSTRACT ideae, and 40 genera, many of them new. Most ge- The Myriogramme group of Kylin was found to contain neric names proposed by Kylin had been used pre- two distinct clusters of genera that m’trecognition at the viously by J. Agardh (1876, 1898) for subgenera or tribal level. In this paper, we establish the tribe Myriogram- “tribes” (sections) in Nitophyllum or Delesseria. My- meae based on a study of the type species of Myriogram- riogramme Kylin (1924:55),type of the Myriogramme me, M. livida, from the Southern Hemisphere. The My- group Kylin (1924:53), for example, was based on riogrammae is charact~zedby 1) marginal and dijfuse Tribus 111 Myriogramma of subgenus I1 Aglaophyllum intercalaly meristems; 2) nuclei arranged in a ring bor- in Nitophyllum u. Agardh 1898:42). Kylin designated dm‘ng the side walls of vegetative cells; 3) microscopic veins Nitophyllum lividumJ. D. Hooker et Harvey 1845 as absent; 4) procarps scattered, fmed opposite one another the type species of Myriogramme because, in his opin- on both sides of the blade postm.or to one or more vegetative ion, it included Nit~hyll~mgrayanum J. Agardh pm’central cells {covcr cells) and consisting of a carpogo- (1876:449), the first species listed by J. Agardh nial branch, a one-/to two-celled lateral stmde group and (1898) under Mynogramma. An element of confu- a one-celled basal sterile group; 5) auxiliav cell diploidized sion was introduced from the beginning in that, al- by a connecting cell cut o~postm.olaterallyfrom the fiil- though Kylin selected M. livida as the type species, ized caqogonium; 6) gonimoblast initial cut off distally he based his concept of Myriogramme primarily on a from the auxilialy cell, generating one distal and one to study of M. minuta Kylin (1924:56), a new species two lateral gonimoblast $laments that branch in the plane from Naples, Italy. Subsequent investigations of re- of the expanding qstocarp cavity and later fuse to fm lated taxa placed in Nitophjllum Greville 1830 (nom an extensive, branched fusion cell; 7) spermatanpal and cons.), Myriogramme Kylin 1924, Haraldiophyllum Zi- tatrasporanpal sm’fmed inside the margzn on both sides nova 1981, or Hideophyllum Zinova 1981 have not of the blade by re.sumption of meristematic activity; and 8) resolved the many ambiguities arising from Kylin’s tetrasporangza produced primarily from the central cells. original characterization of Myriogramme based on The Myriogrammae currently includes Myriogramme nontype material (Kylin 1925, 1929, 1934, 1956, Kylin, Gonimocolax Kylin, Haraldiophyllum A. Zi- Hollenberg and Abbott 1965, Abbott 1969, Norris nova, Hideophyllum A. Zinova, and a possible unde- and Wynne 1969, Mikami 19’72, Zinova 1981, Wynne scribed genus from Pacajic North and South Amka. Gen- 1983, 1986, 1996, Millar 1990, 1994, Millar and era are separated based primarily on features of gonimo- Huisman 1996). blast and caqosporangial development. We began our study of Myriogramme by surveying procarp and cystocarp development in type and Kty index words: Ceramiales;Delesseriaceae; morphology; field-collected material of potentially interesting spe- Myriogramme; Myriogrammeae tib. now.; Rhodophy- cies placed in the Nitophyllum group, the Crypto- ta; subantarctic; systematics; taxonomy pleura group, and the Myriogramme group in the North Atlantic Ocean, Pacific North and South In 1923, Kylin investigated the vegetative and re- America, Australia, and New Zealand. Our prelimi- productive development of the type species of each nary observations indicated that the Myriogramme of five prominent European genera of Delesseri- group requires systematic revision. In this paper, we aceae using material stained with hematoxylin. He describe the developmental morphology of type and divided the Delesseriaceae into two subfamilies field-collected material of M. livida, the type species based on the position of procarps and cystocarps of ~yriogramm~,and propose a new tribe Myriogram- (pre-/and postfertilization stages of the female re- meae trib. nov., to contain Myriogramme, Gonimoce productive apparatus): along the midrib of a fertile lax, Haraldiophyllum, Hideophyllum, and a few species blade in Delesserioideae versus scattered randomly currently placed in Nitophyllum. over the blade surface in Nitophylloideae. The fol- lowing year, Kylin (1924) published a comprehen- MATERIALS AND METHODS sive taxonomic revision of the Delesseriaceae in Type collections of Myriagrainme lavida and M. crozicri used in which he divided the family into 11 “Gruppen” this study were kindly loaned by The Natural History Museum (groups), 5 in Delesserioideae and 6 in Nitophyllo- (Loridoil). Morphological observations were largcly niade on ma- terial fixed in 8-10% fornialiti-s~awater and preserved in 5% for- malin-seawater. Whole-mount and sectioned rnatcrial either was Received 25 June 1996. Accepted 16 October 1996. stained with aniline blue and mounted in browsyrup or glycer- Authot- for reprint requests. ine or was treated with Wittmann’s aceto-iron-hcniatoxylin-chlo- ’< Present address: Department of Biology, P.O. Box 42451, Uni- ral hydrate (Wittmann 1965) and mounted in 50% Hoyer’s versity of Southwestern I.ouisiana, Lafayette, Lousiaria 70504. mounting medium, as described in Hommersand et al. (1992). 106 MYRIOGRAMMEAE TRIB. NOV. 107 Whole-mount preparations were sometimes cleared for critical cell, afterward radiating upwardly and bearing pri- observations. Slides were placed in an alcohol chamber for 12- mary carposporangia either terminally or in chains; 24 h after destaining in 45% glacial acetic acid, transferred by stages (% absolute ethanol, 34 xylerie; y3 absolute ethanol, *h xy- secondary carposporangia, where present, develop- lene) into xylene, and mounted in Piccolytem (Ward’s Natural ing sympodially from inner gonimoblast cells and Science Establishment, Inc., Rochester, New York). Drawings were either terminal and solitary, or forming clusters or made with a camera lucida, and photographs were taken with a branched chains. Fusion cell formed progressively, Zeiss Photomicroscope 111. Herbarium abbreviations follow Holm- gren et al. (1990). incorporating the supporting cell, central cell, resid- ual auxiliary cell, and primary gonimoblast cell and OBSERVATIONS extending to include inner gonimoblast cells and cells in floor of cystocarp. Sterile groups either ex- Mynogramme Kylin (1924:55) cluded from, or included within, fusion cell. Cysto- Description: Plant body foliose, erect and basally carp cavity formed schizogenously, covered by a attached, or repent and attached secondarily by rhi- multilayered pericarp with a single ostiole overlying zoids; entire, sinuous, lobed, lacerate, or deeply di- sterile group 1. Tetrasporangia bome in small cir- vided; membranous above, thickening toward the cular, elliptical, or rectangular sori scattered ran- base, sometimes with a basal nerve, or the nerves domly bordered by a sterile margin in upper parts subdichotomously branched in lower parts; estipi- of blade. Sori formed by renewal of meristematic tate or with a compressed to cylindrical stipe. Blade activity, mostly five cell layers thick at maturity. Pri- monostromatic or tristromatic at margin, polystro- mary tetrasporangia arising two (to four) per cell matic toward base and in basal thickenings, nerves from central cells. Secondary tetrasporangia also cut and stipes. Microscopic veins absent. Cells polygonal off from central cells and possibly from inner cor- in surface view and rectangular in sectional view, ar- tical cells. Division of tetrasporangia tetrahedral. ranged in horizontal tiers and vertical rows in po- lystromatic parts. Growth diffuse, by marginal and Mynogramme Ziuida u. D. Hooker et Harvey) intercalary meristems, the divisions often perpendic- Kylin (1924:58) ular to one another, producing clusters of rectan- (Figs. 1-55) gular cells that later become polygonal in shape. Description: Thalli erect from a compact basal Chloroplasts parietal, simple or dissected in margin- rhizoidal system to 40 cm high, bluish red to rose al cells, later breaking into small units of defined red or reddish brown, foliose with acute to rounded size and shape linked initially by fine strands. Nuclei tips and smooth or ruffled margins, undivided, la- one to many in meristematic cells, at first arranged ciniate, lobed or deeply dissected; blade mostly in a median plate or ring alongside lateral walls, lat- monostromatic, polystromatic toward base, estipitate er forming two rings bordering side walls on either or with a compressed to cylindrical stipe and basal side of plane passing through middle of blade, ul- nerve, or the nerves subdichotomously branched be- timately lying in a broad band bordering side walls low. Central cell layer of stipes and polystromatic between chloroplasts. Primary and secondary pit parts typically broader than the cells on either side, connections initially median, usually one between or with several cell layers in the center distended each adjoining cell, sometimes with finer secondary forming a central core of larger diameter cells. Sper- pit connections formed later near cell surfaces. matangia borne in elliptical to irregularly shaped, Spermatangia produced in small circular to elliptical separate or confluent sori
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