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The Genus Leptostomum R. Brown (Musci) in Southern South America

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The Genus Leptostomum R. Brown (Musci) in Southern South America Journ. HallOri BOI. Lab. No. 69: 257- 264 (Jan. J99/) THE GENUS LEPTOSTOMUM R. BROWN (MUSCI) IN SOUTHERN SOUTH AMERICA CELINA M. MATTERI! The genus Leptostomum R. Brown comprises only eight species in the world, according to Hyv6nen (1987), of which two are recognized in Southern South America: L. menziesii R. Brown and L. splachnoideum Hook. et Arnott. The genus is currently treated as the only member of the family Leptostomataceae within the Bryineae sensu Brotherus (1924). TraditionalJy the species have been distinguished by gametophytic and sporophytic characters, though the latter are usuaIJy difficult to examine due to the lack of recently mature sporophytes (Koponen & Norris 1985, Shaw 1985, Hyv6nen 1987). Observations based on herbarium material are impracticable because drying capsules lose spores and peristomes. During my revision of the genus for the Fuegian moss flora abundant fruiting material became available. Even when the local species are easily separated by vegetative characters, an attempt was made to present as ac­ curate observations as possible on the structure of the sporophytes, which exhibit con­ siderable differences at the specific level. More importantly these studies are expected to help in the disputed placement of the genus (Andrews 1951, Sorsa 1976, Koponen 1978). In particular the spore morphology of the local species examined by SEM reveals some relationships to the Bartramiineae rather than the Bryineae. Both species have large thick-walled spores, with relatively large, secondarily sculptured verrucae or pila-like processes with a slight external evidence of polarity, all characteristic features of the Bartramiineae. Thus, these observations reinforce the suggestion of various authors (Sorsa 1976, Koponen 1978) in that the family Leptostomataceae should be better placed within the suborder Bartramiineae. PhytogeographicalJy the genus shows a Gondwanic distribution, occurring mostly in Southern South America and Australasia with some ingression in SE Asia. These species are endemic with the sole extension of L. menziesii in the Juan Fernandez Islands (Robinson 1975). In this paper I take the opportunity to present fuIJ descriptions of both local species since they did not exist other than the picturesque translations of Montagne's descriptions in Gay (Montagne in Gay 1850). Leptostomum R. Brown, 1811, nom. cons. The genus Leptostomum in South America is easily distinguished from its relatives by its big size, growing always in compact short turfs, the stem leaves tufted on upper parts of stem, piliferous and with short elongate ceIJs. Fertile female plants typicaIJy 1 Museo Arg. Cs. Nat. B. Rivadavia, Av. A. Gallardo 470, 1405 Buenos Aires, Argentina. 258 Journ. Haltori Bot. Lab. No. 69 1 99 I show a bunch of long perichaetial aristae at the apex. The sporophytes with elongate setae, big erect capsules, a reduced peristome and spores with large processes are addition! diagnostic features. KEY TO SPECIES \. Pl ants robust (up to 3cm). Stem leaves up to 4mm long, margins revolute, costa excurrent into a strong wavy arista. Spores up to 381lm in diameter . .. .. .. .... .... L. menziesii I. Plants shorter. Stem leaves up to 2 mm long, margins plane, costa subexcurrent, with a thin. articul ate arista formed by ex tension of the lamina. Spores up to 26 pm in diameter ... .. .. .. ..... .. ..... .. .. .. .... .. .. .. .. L. splachnoideum I. Leptostomum menziesii R. Brown Fig. 1- 7, 18- 21 Trans. Li nn. Soc. London 10: 321. 181 1. Type: "America Australis, Staten Land, A . Men=ies, anno 178T (BM) . Gnnllostomul1Il1Iell=iesii (R. Brown) Hook., Musci Exo!. I: 6. 1818. Leptodol1lilll1l bullocki Thcr.. Rev. Chil. His!. Na!. 33: 517. pI. 28, fig. 1- 9. 1929. Type: ··Chile. Cordillera Nahuelbuta. D . S. Bullock. IV. 1929" (pc -Ther.), (err. typ. pro Leptostol1lul1l). Leptodolltilll1ll1le/desii( R . Brown) Ther. Rev. Chil. His!. Nat. 33: 515. 1929 (err. typ. pro LeplOslomul1I). Plants robust, light green to yellowish, densely tufted with thick red-brown tomentum. Stems mostly single, \.5- 3.0 cm tall, densely tomentose throughout, rhizoids papillose. In cross-section stems angular, epidermal ce ll s small and thick-walled, in 2 3 rows, inner cortex of larger hyaline cells. central strand distinct. Leaves somewhat spirally arranged and tufted in upper 2/3 of stem, erect to erect-patent when wet, concave, oblong-ovate to obovate, with narrow bases, I -4 mm long., 0.5- 1 mm wide, widest just below the middle, acute to graduall y narrowed apices; margins occasionally planc, generally revolute from widest point to near apex, entire or wi th a few apical teeth; lamina extending a longside the excurrent costa; costa strong, long exeurrent in a wavy arista o f 0.5- 0.8 mm long, occasiona ll y weakened below the apex but always linked with the arista by several elongate cells. In cross-section, costa with 3 4 guide cells, adaxial and abaxial stereid bands and thick-walled epidermis. Median leaf cells isodiametric to elongate (14)22(38) x (6) 12( I 6) pm, irregularly inerassate with ± st raight angles, 2- 3 rows of marginal elongate cells with thicker transverse walls forming an indistinct border, cells larger near the base and costa. Dioicous. Male plants un branched or with short erect branches, 1. 5- 2 cm ta ll , perigonia terminal on main stems or branches, gemmiform. Setae elongate, (9) I 7(22) mm long. Capsule golden ye ll ow, elongate and typically arched upward, (6)7( 10) mm long, plane or curved on one side and concave on the opposite, with a horizontal mouth, rugose when dry; exothecial cells inerassate, elongate, 80- 120 tlm long, shorter toward the mouth, stomata somewhat sunken but otherwise phaneropore on a scarcely distinct neck . Annulus incomplete, with scattered cells around the mouth. Peris/ome formed by a thin yellow membrane, extremely delicate and fragile, granulose throughout, 200-250 pill long, irregularly cracked at free border. Spores ye ll owish, subspherical with flallened proximal faces (26)32(38) I'm in diameter, densely covered by relatively large uneven verrucae, smaller on proximal fa ce; operculum convex with a short beak. This is a unique species, not to be mistaken for any other moss in the Fuegian region where it is usually found with its big sporophytes, Valdivian small , sterile forms of the species could hardly be mi staken for L. splachnoideum. However, L. menziesii is usually recognized by its bigger plants, with strongly recurved leaf margins, toothed C. M . MATTERI : Genus Leptostomum in southern South America 259 1 4 . '. ..... ' . ... ~. ..: . ' ". .., ,". .. ~ . ., :. \. ;" .. l ", ~ . FIGS. 1- 14. Leptostomum spp. 1- 7. L. m e n~iesii. I . Habit offemale plant. 2. Wet capsule. 3. Part of peristomial membrane. 4. Spore. 5. Stem leaf. 6. Median leaf cells. 7. Exothecial cells. 8- 14. L. spJachnoideum. 8. Habit of female plant. 9. Wet capsule. 10. Pa rt of peristomial membrane. 11. Spore. 12. Stem leaf. 13. Median leaf cells. 14. Exothecial cells. 1- 7 from CM 3576; 8- 14 from Gay 2238 (pc). 260 Journ. Hattori Bot. Lab. No. 69 199 I at apex, and a thicker arista which is an extension of the costa. Forms with a weaker subapical costa also occur but the combination of characters mentioned above should distinguish them. In this connection it should be noted that, in my opinion, the specimen Kalela 191k (H) mentioned and figured in Hyvonen (1987) is L. menziesii judged solely by gametophyte characters. Consequently, Fig. 3d, e & g from Hyvonen (1987) belong to this species and not to L. splachnoideum as stated there. Morphological details of the peristomial membrane were obtained by dissection of 14 ripe capsules, though its density and granulated nature prevented any counting of the number of layers. The whole cell pattern is strongly irregular. The conspicuous spore ornamentation which was shown by Sorsa (1976) as hollow, reveals some similarities to some of the Bartramiaceae, such as some species of Conostomum (Hirohama 1977, Matteri 1984) and Breutelia (Hirohama 1977). Large, secondarily sculptured spore processes are also common features on this family. Ecology and distribution. Leptostomum menziesii is a common and locally fre­ quent element from the wettest Nothofagus forests in well-lit microhabitats. It usually grows on living tree trunks - various species of Nothofagus, Pilgerodendron uviferum or Drimys winteri - or on shrub twigs - Pernettya mucronata, Berberis spp., Chilio­ trichum diffusum - at ca. 0- 1 m from ground level. It is a dominant epiphyte on branches of Nothofagus antarctica and Berberis ilicifolia in flooded open areas in the rain forests of Southeastern Fuegia, and seems to be more abundant there than fur­ ther North. Less often it is found on rotten logs or on rocks and it usually shares the substratum with Lepyrodon /agurus, Leptotheca gaudichaudii and/or Tortula anderssonii. It is known from sea level in the South to ca. 1500 m a.s.1. in Northern Patagonia. This is a widely ranging endemic occurring from ca. 56° S in Fuegio­ Magellanic region to ca. 37° S in North Patagonia and in Juan Fernandez Islands, both East and West of the Andine Cordillera. Specimens examined. Argentina. Neuquen. Lago Tromen, 39° 32' S, 71 ° 27' W, Ka/e/a /9/k (H); Lago Currhue, 39° 52' S, 71 ° 28' W, Eskuche M12 (CM 4784*); Lago Ortiz Basualdo, 40° 59' S, 71° 52' W, P. Moreau (CM 4738). Rio Negro. Lago Nahuel Huapi, 40° 58' S, 71 ° 30' W, Dusen 713 (H). Chubut. Lago Vintter, 43° 55' S, 71 ° 30' W, Eskuche /941-16 (BA) . Santa Cruz. Depto. Lago Argentino, Bahia Onelli, 50° 08' S, 73" 26' W, Matleri 5124 (CM); Laguna Torre, 49° 20' S, 73° 03' W, Schiavone (CM 4875); Seno Mayo, 50° 22' S, 73 ° 16' W, Malleri (CM 49(7); Seno Spegazzini, 50° IS'S, 73° IS' W, P. Moreau (BA) . Tierra del Fuego. Depto. Ushuaia. Isla de los Estados. Pt. Punta Roca, 54° 44' S, 64° IS' W, Malleri (CM (273); Pt.
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