Ulota larrainii Garilleti, Mazimpaka & F. Lara, 2015

Garilleti, Ricardo, Mazimpaka, Vicente & Lara, Francisco, 2015, Ulota larrainii (Orthotrichoideae, Orthotrichaceae, Bryophyta) a new species from Chile, with comments on the worldwide diversification of the genus, Phytotaxa 217 (2), pp. 133-144 : 133-141

Ulota larrainii Garilleti, Mazimpaka & F. Lara , sp. nov., Figs. 1–19

Plantae caulibus principalibus procumbentibus; folia sinuosa, non contorta, linearia ad linear-lanceolata basi dilatata, marginalibus cellulis in 1(2) seriebus differentiatis et dilatata basi restrictis. Vaginula pilis 2(3)–seriatis curtis, coloratis papillosisque. Seta 1.6–1.8 mm longa. Capsula exserta, urna in sicco vacuoque curte cylindrica ad ovatam. Peristomium duplex. Exostomii dentes 16, aurantiaci, opaci, revoluti. Endostomium hyalinum, evolutissimum, 16 latis planisque segmentis, in sicco incurvatis. Calyptra conica , plicata, luteola obscuro rostro munita, pilosa. Sporae multicellulares, obscure virides, sphaericae, 100–140 μm.

Type:— CHILE. Provincia Capitán Prat: Caleta Tortel, comienzo de “ruta turística”, en tundras de Donatia , Astelia y Lepidothamnus fonkii ; lat. 47º47’32”S, long. 73º31’48”W; alt. ca. 90 m; epífito en Nothofagus dombeyi, J. Larraín 26884 A, con R. Vargas, 20 Ene[ro] 2007 (holotype: VAL-Briof, isotype: CONC).

Plants small, prostrate, forming creeping patches, not crisped. Rhizoids reddish, smooth, on basal parts of stems. Leaves erect when dry, sinuose, erect-patent when moist, linear to linear-lanceolate, with widened base, 1.30–1.60(–1.85) mm long. Leaf base elliptical, sometimes ovate-rounded, slightly concave, gradually narrowing to a long triangular to almost linear variably keeled lamina. Apex long acuminate. Leaf margins plane, frequently one of them slightly recurved in the transition between base and lamina, entire to scarcely crenulate. Costa ending below apex, 25–45 μm wide at base, 20–25 μm at middle leaf. Basal leaf cells rectangular to irregularly trapezoidal, strongly thick-walled, with linear to somewhat vermicular lumen, (13–)18–30(–40) × 7–13 μm, wider and coloured orange-red near leaf insertion; differentiated marginal cells in 1(2) rows limited to the base, quadrate to shortly rectangular, with longitudinal and transversal walls somewhat and equally thickened. Median and upper laminar cells isodiametric to elliptic, thick-walled, smooth, 11–20 × 9–12(–14) μm.

Goniautoicous. Perigonia lateral on main stems and branches. Perichaetia terminal on main stems and branches. Perichaetial leaves distinctively differentiated, longer than the vegetative ones, 2.0– 2.5 mm long, lanceolate with the base not sheathing, gradually narrowing to a somewhat enlarged lamina that ends in a blunt apex. Vaginula hairy, with hairs short, stout, yellowish to coppery, bi–triseriate, uniseriate at apex, papillose. Seta 1.6–1.8 mm, relatively broad. Capsule exserted, shortly cylindrical to ovate when dry, globose to pyriform when moist, 1.0– 1.2 mm long, narrowed to seta through a short neck, not constricted below or at the orifice, mouth rounded in shape. Urn sulcate along all its length by 8 well marked furrows. Exothecial bands eight, formed by 2–4 rows of shortly rectangular cells with moderately thickened longitudinal walls, separated from the mouth by a ring of 2–4 rows of oblate cells. Stomata phaneropore, located in the urn base. Peristome double. Prostome inconspicuous, very fragmentary or absent. Exostome formed by 8 pairs of teeth orange-brownish, thick, opaque, revolute when dry, generally touching the capsule wall at the teeth tips, but sometimes not reaching the exothecium, 240–300 μm long. External side of teeth (Outer Peristomial Layer, OPL) ornamented by a tall, bulging papillose reticule, less prominent towards the tip; internal side of teeth (Primary Peristomial Layer, PPL) with very low, sometimes almost inconspicuous, longitudinal lines along most of the tooth, papillose in the upper part. Endostome of 16 broad, plane, triangular, hyaline to light-orange segments, joined at their bases by 1–2 rows of cells; main segments biseriate, intermediate segments uniseriate, sometimes partially biseriate, 2/3–3/4 as long as teeth; endostome PPL smooth, internal side of segment (Inner Peristomial Layer, IPL) ornamented in the lower 2/3 by an irregular pattern of lines, sometimes with scattered papillae or dominated by low papillae, in the upper 1/3 papillose. Calyptra campanulate, hairy. Spores multicellular, deep green, mostly spherical, sometimes shortly ovoid, thick-walled, 100–140 μm, verrucose to almost smooth since the verrucae usually partially detach and remain on the spore sac.

Distribution and Ecology:— Ulota larrainii is only known from the type locality, Caleta Tortel, located by the sea in Aysén Region, an area with annual rain average over 3,000 mm, mean relative humidity of 90% and an average temperature of 8ºC. The new species was collected on Nothofagus dombeyi ( Mirbel 1827: 467) Oersted (1871: 354) in a tundra area with pulvinate shrubs of Astelia pumila ( Forster 1789: 30) Brown (1810: 291) , Donatia fascicularis Forster & Forster (1776: 10) , Oreobolus obtusangulus Gaudichaud (1825: 99) , Gaimardia australis Gaudichaud (1825: 100) , etc. The surrounding vegetation consists of a hyperhumid evergreen forest including, among others, Nothofagus dombeyi , N. nitida (Philippi 1857: 44) Krasser (1896: 163) , Weinmannia trichosperma Cavanilles (1801: 45) , Drimys winteri Forster & Forster (1776: 84) , Podocarpus nubigenus Lindley (1851: 264) , Amomyrtus luma ( Molina 1782: 352) Legrand & Kausel ( Kausel 1948: 146) , and Caldcluvia paniculata ( Cavanilles 1801: 44) Don (1830: 92) , with a dense understory formed mainly by Chusquea spp. , Desfontainia spinosa Ruiz & Pavón (1799: 47) , Berberis spp. , and Fuchsia magellanica Lamarck (1788: 565) .

Etymology:— Named after Juan Larraín, the first collector of this moss, and as a tribute to his contribution to Chilean Bryology.

Discussion

Taxonomic differentiation:— Ulota larrainii can be distinguished from other species of the genus by a combination of several sporophyte and gametophyte characters ( Figs. 1–19 ): 1) multicellular, large spherical spores; 2) thick seta;

3) capsule shortly exserted, very shortly cylindrical and not constricted below the mouth, with a short neck, and the urn sulcate with wide ribs; 4) exostome formed by 8 pairs of orange to brownish teeth that are revolute and usually touch the exothecium only at the tips; 5) endostome of 16 hyaline, very broad segments joined at the bases by 1–2 rows of cells; 6) vaginula with short, stout papillose hairs; 7) prostrate habit; 8) sinuose, not crisped leaves; and 9) basal differentiated marginal cells in 1(2) rows, shortly rectangular to quadrate.

Some of the distinctive characters that differentiate Ulota larrainii from other species in the genus can be found in both the sympatric U. billbuckii and U. magellanica (Montagne 1843: 242) A. Jaeger (1874: 164). These are the prostrate habit, the leaves that are merely sinuose and do not give the plant a crisped aspect, the hyaline differentiated marginal cells of the leaf base in 1(2) rows, the endostome of 16 broad, shining, hyaline to pale orange segments, and the large spores. The differences between these three species are, however, numerous and include a set of characters that have clear discriminatory value in the genus. A synthesis of these differences is shown in Tab. 1. A full account of U. billbuckii was published in Garilleti et al. (2012), but no modern illustration of U. magellanica is available, so we present one here ( Figs. 20 , 22–31 ) to facilitate its differentiation from U. larrainii . In addition, the following dichotomous key should assist with efficient discrimination between the three mosses:

1. Spores unicellular, 40–50 μm in diameter, capsules cylindrical longer than 1.2 mm; vaginula hairs long, smooth or with very sparse papillae, frequently protruding between the perichaetial leaves ................................................................................. U. magellanica

1’. Spores multicellular, 100–140 μm in diameter, capsules short cylindrical up to 1.2 mm long; vaginula hairs short, papillose, hidden in the perichaetial leaves....................................................................................................................................................................2

2. Exostome of 8 pairs of teeth completely split into 16, yellowish to whitish; endostome with a remarkably tall continuous basal ring 3–5 cells high; spores oblong or irregularly elongate ...................................................................................................... U. billbuckii

2’. Exostome of 8 pairs of teeth, orange to brownish; endostome without a tall continuous basal ring; spores mostly rounded in shape.................................................................................................................................................................................. U. larrainii

The only other Ulota species with multicellular spores from Chile is U. streptodon . This species is easily separated from U. larrainii by its long ellipsoid to almost fusiform capsule, the mouth somewhat puckered and star-shaped, and a very different peristome, consisting of white or pale brown exostome teeth formed of 16 teeth joined at their bases and remaining erect or very irregularly curved, and an endostome of 8 rudimentary segments.

Multicellular spores in Ulota :— Until recently ( Caparrós et al. 2011), multicellular spores were thought to be an uncommon feature in Ulota since they were only known to occur in three species of the genus. However, this number has increased significantly in the last few years ( Caparrós et al. 2011, Garilleti et al. 2012), rising to eight species including U. larrainii described here. These species are irregularly distributed across the world ( Garilleti et al. 2012), since they have been reported in several areas around the Pacific Ocean. Three of them are native to Eastern Asia, namely Ulota yakushimensis Iwatsuki (1959: 153) , U. yunnanensis F. Lara, Caparrós & Garilleti in Caparrós et al. (2011: 211), and U. gigantospora F. Lara, Caparrós & Garilleti in Caparrós et al. (2011: 214); U. novo-guinensis Bartram (1959: 91) is only known from Papua New Guinea; another species, U. membranata Malta (1932: 18), occurs in Tasmania and New Zealand. Finally, three species, U. billbuckii , U. streptodon , and the here-described U. larrainii are confined to southern South America.

It is more difficult to estimate the percentage of species in the genus that have multicellular spores, since no updated lists of accepted species of the genus are available. Index Muscorum (Van der Wijk et al. 1969) recognized 74 taxa, 14 of them of infraspecific rank. Since then, estimations of the number of accepted Ulota species have ranged from “over 50” to “somewhat more than 60 taxa” ( Eddy 1996, Crosby et al. 2000, Albertos et al. 2008, Caparrós et al. 2011). Eleven species have been added to Ulota since the publication of Index Muscorum , with the description of nine new species ( Hoe & Crum 1971, Guo et al. 2004, Caparrós et al. 2011, Wang & Jia 2012, Garilleti et al. 2012) and the transfer of two species from related genera ( Goffinet & Vitt 1998, Albertos et al. 2008). Additionally, selected regional checklists and floras (see references in Fig. 32) provide data on different taxa that are likely to gain eventual acceptance, giving a list of 69 taxa that are currently accepted or have not been synonymized (67 species and 2 varieties: see Appendix). Consequently, species with multicellular spores currently represent 11.6% of the Ulotae and we can no longer consider that multicellular spores are exceptional. However, species with this type of spore are not widespread. All of them are endemic to restricted areas of the world, and so the phenomenon within the Ulotae should be considered rare from a biogeographic perspective.

Ulota distribution and diversification:— Ulota has a bipolar distribution preferentially linked to oceanic environments in temperate areas, which leads the genus to be more common in middle to high latitudes. It prefers, with few exceptions, epiphytic biotopes. It is rare in tropical or subtropical areas, where it finds refuge in mountainous areas with mitigated average temperatures. These few insights are the only biogeographical perspectives of the genus we currently have, as no general assessments of the worldwide distribution of its species are available. In an attempt to address this, we have prepared to map the different areas of the world where Ulota species are present, relating them to the number of recognized and endemic taxa. As the map shows ( Fig. 32), there are 7 main diversification centres for the genus, each of them comprising 7–22 taxa: Western North America (a), Eastern North America (c), Europe and its surroundings (d), North Eastern Asia (g), Subtropical China and its surroundings (h), Southern South America (n), and Southern Australasia (r). Two additional areas involve 4 species: Altai and the neighbouring areas of central Asia (f), and Papua New Guinea (m). The number of endemics ranges from 0 to 18 but endemism rates are high in most of these areas, and outstanding in four of them: Papua New Guinea (75.0%), Subtropical China (77.8%), Southern Australasia (78.6%), and Southern South America (81.8%).

Another ten areas in the world contain a single species, in most cases very narrow endemics of tropical mountain zones. Exceptions are: 1—northern territories of North America (b) and Eurasia (e) where one widespread species, Ulota curvifolia (Wahlenberg 1812: 365) Liljeblad (1816: 546) , has a scattered distribution; 2—Madagascan region (p) where one endemic species thrives over a relatively wide tropical area; 3— Cape Peninsula in South Africa (o) because of its Mediterranean climate; and 4—Kerguelen Island (q) due to its Subantarctic climate. The species reported in this latter locality is Ulota phyllanthoides (Müller 1884: 80) Kindberg (1889: 96) ; it has not yet been revised and probably corresponds to U. phyllantha Bridel (1819: 113) , which is known from other Subantarctic areas, and is the only species in the genus present in both hemispheres.

Southern South America, and particularly the hyperoceanic area linked to the Andean zone of Patagonia and Fuegian regions, is not only the world nucleus with the highest endemicity for Ulota , but it is also the richest area in number of species (22), far exceeding the next richest, Southern Australasia (14 taxa). While this could reflect the lack of modern taxonomic revisions of the genus in that area, we think that the revision by Malta (1927) still retains its relevance. The description here of U. larrainii , along with the other recently described new species from the same area ( Garilleti et al. 2012), tells a different history that points towards a very diverse and poorly explored area.

Additional studied specimens:— Ulota magellanica . ARGENTINA: Neuquén –Villa La Angostura, orillas del lago Nahuel Huapi, 40º48’22’’S, 71º35’55’’W, 790 m, bosque de Nothofagus dombeyi antropizado, sobre Nothofagus dombeyi, F. Lara & San Miguel. 18-XII-2005 ( MAUAM).

CHILE, Provincia Capitán Prat: Caleta Tortel, en las pasarelas del pueblo, lat. 47º’47’57” S, long. 73º32’49”W, ca. 5 m, epífito en gran Nothofagus dombeyi junto a la pasarela, J. Larraín 26927 A & R. Vargas, 20/01/2007 (VALBriof). Provincia Capitán Prat: en bosque de Nothofagus dombeyi ribereño junto al río Carrera. lat. 47º397’33”S, long. 72º58’47”W. ca. 100 m, J. Larraín 26594 & R. Vargas 16/01/2007 (VAL-Briof).

CHILE, Provincia General Carrera: Puerto Bertrand, ca. 5 km al sur del pueblo por la carretera, lat. 47º02’15”S, long. 72º49’09”W ca. 210 m, bosque de Nothofagus dombeyi en pendiente junto al camino, epífito en Nothofagus dombeyi, J. Larraín 26756 B & R. Vargas, 18/01/2007 (VAL-Briof).

CHILE, Región de Magallanes y de la Antártica Chilena. Provincia Antártica Chilena, Comuna de Cabo de Hornos: Parque Nacional Alberto de Agostini; north-central west of Isla Hoste, ca 5 km W of the eastern tip of Isla Gordon along SW arm of Beagle Channel in unnamed sound, 55º00’22”S / 069º12’11”W. 5 m a.s.l., arborescent formation of Nothofagus betuloides with Drimys winteri and scarce N. antarctica , branches of Nothofagus antarctica near the shoreline, Garilleti 2012 –018b, 19/01/2012 (VAL-Briof).

Ulota billbuckii . CHILE. Región X, Provincia Osorno: P. N. Puyehue, zona interfronteriza Los Pajaritos, 40º40’47”S, 71º59’53”W, 1021 m, bosque de Nothofagus pumilio y N. dombeyi , en troncos de N. pumilio, Lara & San Miguel , 31 diciembre 2005 ( MAUAM).

CHILE, Región XI, Provincia Capitán Prat: Caleta Tortel, comienzo de la “ruta turística”, en tundras de Donatia , Astelia y Lepidothamnus fonkii , 47º47’32”S, 73º31’48”W, alt. ca. 90 m, epífito en Nothofagus dombeyi, Larraín 26884 B & Vargas, 20 enero 2007 (VAL-Briof).

CHILE, Región de Magallanes y de la Antártica Chilena. Provincia Antártica Chilena: Comuna de Cabo de Hornos, Parque Nacional Alberto de Agostini, north-central west of Isla Hoste, ca. 5 km W of the eastern tip of Isla Gordon along SW arm of Beagle Channel in unnamed sound, 55º00’22”S, 069º12’11”W, 2 m a.s.l., branches of Nothofagus antarctica, Garilleti 2012 –018a, 19 January 2012 ( NY, Holotype); Parque Nacional Alberto de Agostini, SE coast of Isla Gordon in unnamed sound ca. 10 km W of eastern tip of island on the Brazo Sudoeste of the Beagle Channel, 54º58’38’’S, 069º20’38’’W, 10 m a.s.l., extense, dense forest of Nothofagus betuloides along NNE exposed slopes, trunk and branches of young Nothofagus betuloides, Garilleti 2012 –019, 19 January 2012 (VAL-Briof).

Acknowledgements

We are deeply grateful to Juan Larraín, who sent us many specimens of Orthotrichaceae from Chile, among which we discovered the new species, as well as for his valuable commentaries on many ecological aspects of Aysén Region. We warmly address our thanks to Tom Blockeel for revising and improving the text of the manuscript. The curators and staff of CONC and NY are gratefully acknowledged for sending us several specimens fundamental for our study. This work was funded by the Spanish Ministries of Economy and Competitiveness ( CGL 2013–43246– P) and Science and Innovation ( CGL 2011–28857/ BOS).

Figures

FIGURES 1–9: Ulota larrainii. 1. Habit. 2. Detail of capsules. Those at the lower left with spores retained by the endostome segments. 3. A sporophyte with the differentiated perichaetial leaves erect and appressed to the seta; note the different aspect of the vegetative leaves on the stem at the right side of the sporophyte. 4. Detail of the peristome; note the broad and shiny endostome closing the capsule mouth and the revolute exostome teeth touching the capsule wall only at the tips. 5. Base of a vegetative leaf showing the 1 cell wide hyaline marginal area. 6–7. Vegetative leaves. 8–9. Perichaetial leaves. All from holotype. Photographs by R. Garilleti.

FIGURES 10–18: Ulota larrainii. 10. Inner view of the peristome where the endostome formation can be observed; the dark small structures on the endostome segments are verrucae detached from the spore walls. 11. External view of an exostome tooth with an endostome segment to the back on the right; a very tall papillose reticulum forms the ornamentation of the main part of the external side of the tooth. 12, Detail of the ornamentation from the basal part of the inner surface of the exostome. 13–14. Two variants of the ornamentation of the endostome inner surface. 15. Calyptra. 16–17. Spores. 18. Detail of the vaginula showing the typical short and stout hairs.All from holotype. Photographs by R. Garilleti.

FIGURES 19–21. Comparison of capsules. 19. Ulota larrainii. 20. U. magellanica 21. U. billbuckii. (U. billbuckii and U. larrainii, from their respective holotypes, U. magellanica from Larraín 26927A). Photographs by R. Garilleti.

FIGURES 22–31: Ulota magellanica. 22. Habit. 23. Mature capsule with the endostome erect, a common position of this structure in this species. The wide, flattened neck gives a longer aspect to this specific capsule. 24. An operculate capsule. 25. Calyptra. 26. A vaginula from a dry specimen, with the conspicuous vaginula hairs naturally exposed. 27. Detail of the mainly uniseriate, smooth vaginula hairs. 28–29. Leaves. 30. Perichaetial leaf. 31. Spores. (Figures 22, 24, 28 & 29, from Garilleti 2012–018b, Figures 23, 25 & 26, from Larraín 26594 & R. Vargas, Figures 27 & 31, from Larraín 26756b & R. Vargas, Figure 30 from Lara & San Miguel 18-12-2005). Photographs by R. Garilleti.