Research Article |
Corresponding author: John R. Spence ( bryum500.js@gmail.com ) Academic editor: Matt von Konrat
© 2021 John R. Spence.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Spence JR (2021) Studies in Austral Bryaceae (Bryopsida). III. A Preliminary Account with Keys to Rosulabryum J.R. Spence in Chile. PhytoKeys 185: 87-98. https://doi.org/10.3897/phytokeys.185.70407
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A preliminary study of the genus Rosulabryum J.R. Spence in Chile is presented, with brief species descriptions, notes on ecology and distributions, and a taxonomic key. The following 12 species are confirmed with vouchered specimens; Rosulabryum andicola (Hook.) Ochyra, Rosulabryum billarderii (Schwägr.) J.R. Spence, Rosulabryum campylothecium (Taylor) J.R. Spence, Rosulabryum capillare (Hedw.) J.R. Spence, Rosulabryum coloratum (Müll. Hal.) J.R. Spence, Rosulabryum densifolium (Brid.) Ochyra, Rosulabryum longidens (Thér.) J.R. Spence, Rosulabryum macrophyllum (Cardot & Broth.) Ochyra, Rosulabryum perlimbatum (Cardot) Ochyra, Rosulabryum puconense (Herzog & Thér.) J.R. Spence, Rosulabryum rubens (Mitt.) J.R. Spence, and Rosulabryum torquescens (Bruch ex De Not.) J.R. Spence. Rosulabryum canariense (Brid.) Ochyra is tentatively excluded as the Chilean material can be referred to R. coloratum. Similarly, Rosulabryum viridescens (Welw. & Duby) Ochyra is tentatively excluded since the Chilean plants do not match the African type, but instead appear to be atypical plants of R. campylothecium.
Bryaceae, Chile, mosses, Rosulabryum
With a known moss flora of more than 900 species, Chile has a rich but incompletely documented bryoflora (
The Bryaceae is a large world-wide family that occurs in a wide variety of habitats and climates and can represent as much as 10% of the species richness in local and regional moss floras. Chile has an extremely rich and incompletely known Bryaceae flora, with an estimated 85 species, based on revisions and recent collecting (
One of the most easily recognized and common genera is Rosulabryum J.R. Spence, which is well represented in the southern hemisphere. The genus currently consists of ca. 75 described species (J. Spence unpublished data), with 12 well documented species in Chile. However, much of the country remains under collected, especially in the north and in the Andes, thus it is likely that additional species will be found with more intensive field work. Of particular note is the presence of a diverse montane Rosulabryum flora in the northern and central Andes which may extend southward into the Chilean Andes.
Rosulabryum occurs in a clade with Brachymenium sect. Brachymenium, sister to a clade consisting of Plagiobryum Lindb., Plagiobryoides J.R. Spence and Ptychostomum Hornsch. (
Collections were obtained from various herbaria, primarily CONC, MO and NY. Additional field work in the Tierra del Fuego region of Chile (Magellanes Province) has been completed and collections are currently being studied in the author’s personal herbaria. Much of the recent floristic work has been done in the southern regions of Bío Bío, Aisén and Magallanes (e.g.,
Of the 12 species documented in this study, their distributions include four main elements, 1) widespread temperate, 2) southern temperate, 3) subantarctic/Nothofagus, and 4) Neotropical montane. The widespread element includes R. capillare, R. rubens and R. torquescens. Southern temperate species include R. billarderii, R. campylothecium, and R. macrophyllum, with the first two extending to Australasia. The subantarctic element is represented by R. perlimbatum, which extends to southern New Zealand. The Neotropical montane element includes R. andicola and R. densifolium. An endemic element may also exist, as currently R. longidens and R. puconense are endemic to Chile, with potentially one record of the former reported from Argentina which needs to be verified. Rosulabryum coloratum has an anomalous distribution, as it occurs in montane regions of Bolivia as well as lower elevations of south-central Chile. In general, these distributions conform at least in part to well documented and similar distributions among the vascular plant flora of Chile (
Rosulabryum is characterized by a combination of predominantly gametophyte characters, including mostly rosulate habit to sometimes evenly foliate stems, ovate to obovate or spathulate leaves, a well-developed limbidium, serrulate, serrate or denticulate distal leaf margins, costa with well-developed stereid band, nodding capsule, well-developed peristome with appendiculate cilia, small spores, rhizoidal tubers, and filiform gemmae in the leaf axils (
1 | Filiform gemmae present, usually in axils of sterile shoot leaves | 2 |
1’ | Gemmae absent | 3 |
2 | Plants large, at least some leaves > 3 mm, distal margins sharply serrate; costa of innovations short-excurrent; predominantly terricolous | Rosulabryum andicola (in part) |
2’ | Plants small, leaves < 2 (2.5) mm, distal margins serrulate, costa of innovation leaves long excurrent; terricolous or lignicolous-epiphytic, often on tree trunks and large branches | Rosulabryum longidens |
3 | Plants small, leaves mostly < 2 (2.5) mm, spirally twisted around stem when dry, costa excurrent in long ± straight awn | Rosulabryum capillare |
3’ | Plants small to robust, if spirally twisted around stem then leaves > 3 mm or costa short-excurrent | 4 |
4 | Plants small, leaves < 2 mm, distal leaf margins smooth to serrulate, stems rosulate to somewhat evenly foliate, leaves often with reddish tints; rhizoidal tubers red to orange or red-brown, cell walls protuberant | 5 |
4’ | Plants medium to large, leaves mostly > 2.5mm, distal leaf margins finely serrulate to sharply serrate or denticulate, stems variable but often rosulate, leaves mostly green to yellow-green, rarely somewhat reddish-tinged, rhizoidal tuber color various, cells smooth, lacking protuberant walls | 6 |
5 | Sexual condition dioicous; costa excurrent in medium-length awn, limbidium distinct, leaf margins serrulate, rhizoidal tubers red, cell walls strongly protuberant, diameter to ca. 260 (280) µm; capsules 2–3 mm long | Rosulabryum rubens |
5’ | Sexual condition synoicous or polyoicous with single sex shoots, costa percurrent to short excurrent in short awn, limbidium weak, sometimes absent distally, margins smooth to finely serrulate; rhizoidal tubers red-brown, red to orange, cell walls weakly protuberant, diameter to 600 µm; capsules 4–6 mm long | Rosulabryum puconense |
6 | Limbidium weak distally, of 1 row or sometimes nearly absent; leaves concave; distal laminal cells firm-walled to often incrassate | 7 |
6’ | Limbidium distinct distally, usually of 2 or more rows; leaves flat or weakly concave; distal laminal cells thin to firm-walled but usually not incrassate | 10 |
7 | Plants in tight rosulate tufts when dry, leaves imbricate, golden-yellow or yellow-green; costa excurrent in long ± straight awn; rhizoidal tubers lacking | Rosulabryum campylothecium |
7’ | Plants in loose rosulate to comal tufts or stems evenly foliate, leaves somewhat imbricate to shrunken or twisted when dry, colors various but not golden-yellow; costa excurrent in short often recurved awn; tubers present | 8 |
8’ | Plants ± evenly foliate; leaves spirally twisted around stem when dry; leaf apex acute, costa excurrent in short recurved awn; tubers brown to red-brown | Rosulabryum coloratum |
8 | Plants in one or more interrupted rosulate tufts; leaves weakly imbricate to irregularly contorted when dry; leaf apex broadly acute to rounded or obtuse; costa excurrent in short recurved or straight awn; tubers red, scarlet, brown or yellow-brown | 9 |
9 | Sexual condition autoicous or rarely synoicous; leaf broadly acute at tip, costa tapering towards leaf tip, leaves not strongly keeled; rhizoidal tubers bright red to scarlet | Rosulabryum canariense |
9’ | Sexual condition dioicous or rhizo-autoicous; leaf rounded-obtuse at tip, costa very thick to apex, leaves strongly keeled; rhizoidal tubers brown to yellow-brown | Rosulabryum viridescens |
10 | Sexual condition synoicous or rarely autoicous; plants medium-sized, leaves 2–3 mm, obovate, costa excurrent in medium-length awn; rhizoidal tubers scarlet to red | Rosulabryum torquescens |
10’ | Sexual condition dioicous or rhizo-autoicous; plants medium to robust, leaves mostly > 3 mm, ovate, obovate to spathulate, costa percurrent to excurrent in short awn; rhizoidal tubers dark red, brown to red-brown | 11 |
11 | Limbidium very wide, (3) 6–8 rows, often imparting whitish-hyaline border to leaf | 12 |
11’ | Limbidium narrower, (2) 3–4 rows, colored to clear, not imparting whitish-hyaline border to leaf | 13 |
12 | Leaves spirally twisted around stem when dry, distal margins sharply serrate; limbidium 3–4 rows; filiform leaf axil gemmae usually present | Rosulabryum andicola (in part) |
12 | Leaves irregularly contorted to somewhat imbricate when dry, distal margins serrulate; limbidium 4–8 rows; filiform leaf gemmae absent | Rosulabryum perlimbatum |
13 | Plants elongate evenly foliate, to 8–10 cm, leaves narrowly ovate, distal margins sharply serrate to denticulate, teeth often double | Rosulabryum densifolium |
13’ | Plants rosulate, not evenly foliate, mostly < 2 cm long, leaves obovate to broadly ovate, distal margins serrulate to singly serrate | 14 |
14 | Leaves ovate to oblong, ±imbricate when dry; distal margins serrulate | Rosulabryum macrophyllum |
14’ | Leaves obovate to spathulate, irregularly contorted when dry; distal margins sharply serrate | Rosulabryum billarderii |
A widespread African-Neotropical species found from Chile north to the Southwestern US. This species replaces R. billarderii in warmer climates in Chile and is rare south of the Maule Region. It is found principally on damp to dry soil or soil over rock, rarely on fallen logs and tree stumps. The species is characterized by dioicous sexual condition, strongly rosulate stems, large leaves that are spirally twisted around the stem when dry, a costa excurrent in a short awn, a very well-developed wide limbidium, sharply serrate distal leaf margins, and brown filiform gemmae in the leaf axils of sterile shoots. The rhizoidal tubers are red to red-brown and can be > 1 mm in diameter. (Illustrations:
Representative specimen examined: Region VIII, Bío Bío Province, Ecological Reserve Coligual, 37°23'S 71°40'W, ca. 630 m, on forest floor, clay banks along road with adjacent native forest, R.R. Ireland & G. Bellolio 35234, 21 Nov. 2002 (MO).
The common widespread species of large Rosulabryum from central Chile south to Aisén. The species is also found in Argentina, the Falkland Islands, Australia, New Zealand and Macquarie Island (
Aisén Region, Provincia Capitán Prat, cruzando pasarela Río Ñadis, en bosque de Nothofagus dombeyi, 47°29'49"S, 72°56'51"W, alt. ca. 70 m, sobre tronco caído en claro del bosque, J. Larraín & R. Vargas 26799, 19 Ene 2007 (CONC): Region VIII, Bío Bío Province, road from Tomeco to Florida, 2 km N from Hwy 0–50, 36°57'S 72°40'W, alt. ca. 190 m, on soil bank, R.R. Ireland & G. Bellolio 32401 (MO).
A common temperate species in the central and south regions of Chile from Coquimbo Region to at least the Los Lagos Region but absent from colder subantarctic climates. It also occurs in Australia and New Zealand (
Region VIII, Prov. Ñuble, Cobquecura, rock cliffs near ocean, 36°05'S 72°48'W, alt. ca. 0 m, on cliff, R.R. Ireland & G. Bellolio 32374, 9 Oct 2001 (MO): Region VIII, Prov. Concepción, Puda Beach, 36°29'S 72°54'W, alt. ca. 0 m, R.R. Ireland & G. Bellolio 32267, 7 Oct. 2001 (MO).
A worldwide temperate species found in a wide variety of habitats, but most often on damp shaded soil or fallen logs and tree stumps. Its distribution in Chile remains poorly understood but it is common in the more temperate southern regions such as Bío Bío (
Region VIII, Prov. Concepción, Bellavista Creek, San José Farm, 36°41'S 72°56'W, alt. ca. 90 m, on rotten log, stream with adjacent native forest with predominantly Nothofagus, Aetoxicon punctatum & Peumus boldus, R.R. Ireland & G. Bellolio 33903, 23 Nov. 2001 (MO): Region VIII, Bío Bío Province, Saltillo del Itata, small falls on Itata River, 37°04'S 72°09'W, ca. 210 m, on brick wall beside poplar trees by river, R.R. Ireland & G. Bellolio 34950, 28 Oct. 2002 (MO).
A relatively uncommon species found principally in semi-arid regions in the central and northern regions of Chile and adjacent areas of Bolivia, found on sandy soil or soil over rock, often near streams. The species is characterized by its dioicous sexual condition, evenly foliate stems, large narrowly ovate leaves that are spirally twisted around the stem when dry, a weak limbidium distally, and brown to red-brown rhizoidal tubers.
Aisén Region, Provincia Capitán Prat, río Baker, pasando balsa Baker y luego camino río abajo, en bosque en cerro detrás de la casa de don Delmiro, junto a riachuelo, 47°15'46"S, 72°42'54"W, alt. ca. 100 m, en el suelo junto al arroyo, J. Larraín & R. Vargas 26697, 18 Ene 2007 (CONC); Region VIII, Prov. Ñuble, road from Cobquecura to Quiríhue, 24 km SE of Cobquecura, Mengel Creek, 36°13'S 72°36'W, alt. ca. 360 m, on sandy soil beside creek, R.R. Ireland & G. Bellolio 32323, 8 Oct 2001 (MO).
A robust species distributed throughout the Neotropics, primarily along the mountain chains from Mexico south to Chile, and throughout the Andes (
Bío Bío Region, Prov. Ñuble, Renegado River at Aserradero bridge, 36°54'S 71°28'W, alt. ca. 1000 m, on muddy soil bank beside river, R.R. Ireland & G. Bellolio 30831, 9 Dec 2002 (MO).
A common species in the central regions of Chile, from the Bío Bío north to at least Coquimbo, usually on tree trunks and larger branches, stumps and fallen trees in forests, occasionally on soil. The species is characterized by dioicous sexual condition, small leaves, rosulate fertile stems, with numerous rosulate to evenly foliate innovations with leaves spirally twisted around the stem when dry, strong excurrent costa in medium to long awn which is sometimes red, brown to red-brown filiform gemmae in the leaf axils of sterile shoots, and small red to red-brown rhizoidal tubers.
Region VIII, Bío Bío Province, Lago Falls as south side of Laja River, 37°12'S 72°22'W, alt. ca. 170 m, on tree stump, R.R. Ireland & G. Bellolio 34815, 23 Oct. 2002 (MO); Region VIII, Prov. Arauco, Huillinco Falls, 37°45'S 73°22'W, alt. ca. 130 m, on base of large Nothofagus sp., falls and rock cliff in native forests, R.R. Ireland & G. Bellolio 33662, 12 Nov. 2001 (MO).
An uncommon species of moist to wet soil in the subantarctic moorlands and Nothofagus forests in southern Chile, reaching north to the Aisén Region (
Región de Aisén, Provincia de Aisén, Comuna de Cisnes Parque Nacional Queulat, sector Angostura Risopatrón, sendero Los Colonos, bosque de Amomyrtus luma-Nothofagus nitida-Laureliopsis philippiana, sobre tronco podrido en la orilla del lago, 44°14'04"S, 72°30'24"W, elev. 130 m, J. Larraín 43684, with R. Vargas, E. Muñoz & J.F. Croxatto, 14 dic 2019 (CONC).
A common and widespread species of the subantarctic forests and moorlands of the Magallanes and Aisén Regions on damp to wet soil, fallen logs, and tree stumps (
Prov. Antártica Chilena, Comuna Cabo de Hornos, Parque Nacional Alberto de Agostini, Isla Hoste, Península Dumas, Bahía Ibáñez, Caleta Yekadahby, 55°03'47“S, 68°25'19“W, on soil in shade in moist Nothofagus betuloides-N. pumilio-Maytenus magellanica forest on E-facing slope with extensive rock outcrops, J.R. Spence 6059, 15 January 2013(CAS).
This species was synonymized under R. capillare by
Prov. Antártica Chilena, Comuna Cabo de Hornos, Isla Grande de la Tierra del Fuego, Bahía Yendegaia, NNE shore opposite Caleta Ferrari, 54°50'28“S, 68°47'52“W, on dry exposed soil over rock outcrops near ocean, J.R. Spence 6042, 13 January 2013 (CAS): Region VIII, Bío Bío Province, Saltillo del Itata, small falls on Itata River, 37°04'S 72°09'W, ca. 210 m, on soil bank among poplar trees by river, R.R. Ireland & G. Bellolio 34940, 28 Oct. 2002 (MO).
A rare species generally found in disturbed habitats, especially on disturbed soil and concrete, and possibly introduced from the northern hemisphere. There is only one record from Chile in disturbed habitats in the capitol Santiago (
A widespread warm temperate to subtropical species in the northern hemisphere, Africa, Australasia and South America (
Los Lagos Region, Chiloé, communa de Ancud, Estación Biológica Senda Darwin, Al fondo de al “ciudad de los Muertos”, Sobre tronco caido ed bosque Quemado, 41°52'S 73°39'W, 300 m, J. Larraín 23424, 2 Feb. 2003 (CONC): Region VIII, Prov. Concepcion, Park “Jorge Alessandri” (Compania Manufacturera de Papeles y Cartones), on soil in clearing, 36°56'S 73°09'W, ca. 200–490 m, R.R. Ireland & G. Bellolio 32819, 19 Oct. 2001 (MO).
Rosulabryum canariense (Brid.) Ochyra.
Rosulabryum viridescens (Welw. & Duby) Ochyra.
Robert Ireland, William Buck and Juan Larraín supplied collections for study, for which I am grateful. Thanks to John Atwood and the staff at MO for providing loans and material for study.