BFNA Title: Aloina
Author: C. Delgadillo M.
Date: Jan. 2001
Edit Level: R
Version: 4

Bryophyte Flora of North America, Provisional Publication
Buffalo Museum of Science
1020 Humboldt Parkway
Buffalo, NY 14211 USA


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ALOINA (J. K. A. Mller) Kindberg, Bih. Sv. Ak. Handl. 6(19): 22. 1882, conserved name * [Alo + ina, characteristic of; fleshy leaved like Alo]

Claudio Delgadillo M.

Barbula sect. Aloina J. K. A. Mller, Syn. 1: 596. 1849


Plants in thin turfs or scattered. Stem 0.5--6 mm, central strand present. Leaves forming a rosette, lingulate, suborbicular or deltoid, deeply concave, 0.5--2.8 mm, infolded, partly 2-stratose, base auriculate, sheathing, margins entire to irregularly crenulate distally, apex cucullate, mucronate or piliferous; costa broad, percurrent to long-excurrent, in section hydroids absent, stereid band abaxial, occasionally wanting, filaments present on costa and adjacent 2-stratose area of leaf blade, filament cells cylindrical to subspheric, thin-walled, terminal cell subspheric to nearly conic, with an apical thickening; cells of leaf base thin-walled, rectangular, sometimes differentiated marginally, medial and distal leaf cells thick-walled, mostly transversely elongated, smooth. Sexual condition monoicous or dioicous, perigonia distinct, with non-cucullate unfolded leaves, perichaetia sometimes partly differentiated, the inner leaves with unfolded blades. Seta 4.5--17 mm, capsule cylindric to ovoid-cylindric, erect or slightly inclined, annulus of 1--2 rows of vesiculose cells, deciduous or persistent, operculum conic to long-rostrate, erect or inclined, peristome generally long and twisted, papillose, with a basal membrane. Calyptra cucullate. Spores spheric, finely papillose, 9--25 m.


Species 9 (5 in the flora): mostly on calcareous soils in the Mediterranean Basin, southern Africa, Australia; sparsely distributed from Alaska to Argentina.


This is generally a very distinct genus characterized by differentiated leaf bases, infolded partly 2-stratose laminae, broad costae, and filaments ending in a terminal cell with an apical thickening; the filaments cover the costa and part of the lamina. Plants of Aloina sometimes resemble Indusiella (Grimmiaceae) in habit and general leaf structure although no filaments are found on the leaves of the latter.


SELECTED REFERENCES: Delgadillo M., C. 1973. A new species, nomenclatural changes, and generic limits in Aloina, Aloinella, and Crossidium. Bryologist 76: 271--277. Delgadillo M., C. 1975. Taxonomic revision of Aloina, Aloinella and Crossidium (Musci). Bryologist 78: 245--303.


1. Leaf apex piliferous; costa undifferentiated in the distal two thirds, in section with no stereid band, but with a few stereid or substereid cells in mid-costal region 1. Aloina bifrons

1. Leaf apex muticous; costa percurrent to subpercurrent, in section with 1--7 stereid layers.

2. Leaf base without a differentiated margin of thin-walled hyaline cells.

3. Costa with strong solid papillae on the abaxial side at and near the apex; capsule ovoid-cylindric, peristome short (to 900 m), nearly straight 2. Aloina hamulus

3. Costa smooth abaxially; capsule cylindric, peristome long (600--1200 m) and twisted 3. Aloina aloides

2. Leaf base with a marginal area of narrow thin-walled hyaline cells.

4. Leaves suborbicular to short-lingulate; synoicous (or dioicous); capsule cylindric, operculum conic to short-rostrate 4. Aloina brevirostris

4. Leaves ligulate; dioicous; capsule ovoid-cylindrical, operculum long-rostrate 5. Aloina rigida


1. Aloina bifrons (De Notaris) Delgadillo, Bryologist 76: 273. 1973


Tortula bifrons De Notaris, Mem. R. Acc. Sc. Torino 40: 305. 1838; Aloina pilifera (De Notaris) H. A. Crum & Steere; Aloina rigida var. pilifera (De Notaris) Limpricht


Plants to 3 mm. Leaves lingulate to ovate-lingulate, 1--2.5 mm, margins entire to serrulate, undifferentiated at base, apex piliferous, open, broadly obtuse or rounded; costa undifferentiated, with few or no stereid cells, filaments of 4--8 cells, cells subspheric to cylindric; cells of leaf base 18--37 m, medial and distal cells 9--22 m, papillae none. Sexual condition dioicous. Seta 6--14 mm, urn cylindric to ovoid cylindric, 1.7--3.2 mm; operculum conical, rostrate, erect or slightly inclined, 0.9--1.4 mm; peristome 800--1230 m, strongly twisted; spores 9--24 m.


Capsules mature Jan.--July(--Aug.). Sunny sandy soil or soil over limestone in dry areas; 300--800 m; Calif., Ariz., Idaho; Mexico (Baja California, San Luis Potos, Nuevo Len, Puebla and Zacatecas); s South America; Australia; Pacific Islands (New Zealand); sw Asia (Jordan), s Europe; Africa (Egypt, South Africa).


Aloina bifrons is readily identified by the hyaline hair of the leaf apex and the poorly developed costa that is only evident toward the leaf base, but inconspicuous in the distal half of the leaf. No other North American species of the genus exhibits a piliferous leaf apex.


2. Aloina hamulus (J. K. A. Mller) Brotherus, Nat. Pfl. 1: 428. 1902


Barbula hamulus J. K. A. Mller, Bull. Herb. Boiss. 5: 192. 1897


Plants to 6 mm. Leaves lingulate, 0.6--2 mm, margins entire to irregularly crenulate, undifferentiated at base, apex cucullate; costa subpercurrent to percurrent, filaments of 3--7 cells, cells cylindric to subspheric; cells of leaf base 13--59 m, medial and distal cells 9--26 m, large solid papillae on abaxial surface distally. Sexual condition dioicous. Seta 7--16 mm, urn ovoid-cylindric, 1.1--3 mm; operculum conical to long rostrate, erect or inclined, 0.5--1.2 mm; peristome to 900 m, twisted to nearly straight. Spores 9--13 m.


Sporophytes unknown in flora area. Sandstone or calcareous rocks, sandy soil in dry sunny sites; 760--1800 m; Colo., Ill., Kans., La., Nebr., Okla., Tex.; Mexico; Central America (El Salvador, Guatemala).


The undifferentiated marginal cells of the leaf base and the large abaxial solid papillae near the leaf tip distinguish Aloina hamulus from A. rigida, which, in addition, has a strongly twisted and longer peristome. The ovoid-cylindric capsule and abaxially papillose leaves distinguish this species from A. aloides var. ambigua. Young leaves should be checked when papillae are not evident in mature leaves.


3. Aloina aloides (Schultz) Kindberg, Bih. K. Svensk. Vet. Ak. Handl. 7: 136. 1883


Trichostomum aloides Schultz, Nov. Act. Leop. Car. 11: 197. 1823; Barbula aloides (Schultz) Bruch; Tortula aloides (Schultz) De Notaris


Varieties 2 (1 in the flora): North America; Mexico; Europe; n Africa; Asia, Atlantic Islands (Canary Islands); Australia.


3a. Aloina aloides var. ambigua (Bruch & W. P. Schimper) E. J. Craig in A. J. Grout, Moss Fl. N. Amer. 1(4): 214. 1939


Barbula ambigua Bruch & W. P. Schimper, Bryol. Eur. 2: 76. 139. 1842


Plants to 2.5 mm. Leaves ligulate to lingulate, 1--2.8 mm, margins entire, undifferentiated at base, apex cucullate to nearly open; costa subpercurrent to percurrent, filaments of 3--6 cells, cells subspheric; cells of leaf base 15--64 m, medial and distal cells 11--24 m, occasionally with large solid papillae at leaf tip. Sexual condition dioicous or rhizoautoicous. Seta 5.5--12.5 mm, urn cylindric, 1.5--3 mm; operculum campanulate, 1--1.4 mm; peristome 600--1200 m, strongly twisted. Spores 13--24 m.


Capsules mature Jan.--May. Low, desert areas, banks and dry washes, soil and limestone; 400--800 m; Ariz., Calif.; Mexico (Baja California); Europe; n Africa; Asia (India, Israel, Jordan, Lebanon, Turkey); Australia.


The cylindrical capsules with a long and twisted peristome are distinctive. Forms with short-lingulate leaves may be confused with A. hamulus when sterile. Outside North America, A. aloides var. ambigua may occur as high as 2100 m.


4. Aloina brevirostris (Hooker & Greville) Kindberg, Bih. K. Svensk. Vet. Ak. Handl. 7: 137. 1883


Tortula brevirostris Hooker & Greville, Edinburgh J. Sci. 31: 289. 1824


Plants 0.5--2 mm. Leaves lingulate to suborbicular, 0.5--1.5 mm, margins entire, undulate-denticulate distally, differentiated at base, apex cucullate to nearly open; costa subpercurrent, filaments of 2--5 cells, cells subspheric to cylindric; cells of leaf base 13--66 m, medial and distal cells 13--40 m, papillae none. Sexual condition synoicous or dioicous. Seta 6--17 mm, urn cylindric, 1--2 mm; operculum conic, short-rostrate, erect or inclined, 0.5--1 mm; peristome 250--900 m, twisted. Spores 15--25 m.


Capsules mature (Mar.--)May--Aug. Bare or disturbed soil or silt, roadside banks, calcareous boulders or gravel; 100--1500 m; Greenland; Alta., B.C., Man., Nun., N.W.T., Ont., Yukon; Alaska, Mont.; Europe; Asia (Siberia).


The leaves of Aloina brevirostris are usually short, and cucullate with a reduced lamina; forms with larger laminae may be confused with A. rigida, but the synoicous condition and the cylindric capsules with a conical operculum are diagnostic. The latter species is dioicous, and has ovoid-cylindrical capsules with a long-rostrate operculum.


5. Aloina rigida (Hedwig) Limpricht, Laubm. Deutschl.1: 637. 1888


Barbula rigida Hedwig, Sp. Musc. 115. 1801


Varieties 3 (1 in the flora): North America; South America; Europe; Asia; Africa.


5a. Aloina rigida (Hedwig) Limpricht var. rigida


Plants to 2.5 mm. Leaves short-lingulate to ligulate, 0.5--2.5 mm, margins entire to irregularly denticulate, differentiated at base, apex cucullate; costa subpercurrent to percurrent, filaments of 3--9 cells, cells cylindric to spheric; cells of leaf base 11--88 m, medial and distal cells 9--40 m, papillae none. Sexual condition dioicous. Seta 4.5--17 mm, urn ovoid-cylindric, 1.7--3.4 mm; operculum conical to subulate, long-rostrate, 1.2--1.9 mm; peristome 1200--1750 m, strongly twisted. Spores 11--22 m.


Capsules mature June--Aug.(--Oct.). Rocks, banks, clay, sandy or gravelly soil in deserts, plains or conifer forests; 1000--3000 m; Alta., B.C., N.B., N.S., Nun., Ont., Yukon; Colo., Ill., Iowa, Kans., Mont., Nebr., N.Mex., Tex.; Mexico (Nuevo Len, Quertaro, Zacatecas); South America; Europe; Asia; Africa.

The ovoid-cylindrical capsule with a long peristome and long-rostrate operculum as well as the differentiated basal leaf margins are the distinguishing features of A. rigida var. rigida. Despite recent reports from California, this species has not been confirmed from that state.