BFNA Title: Pterygoneurum
Author: R. H. Zander
Date: January 3, 2003
Edit Level: R Brum+
Version: 2

Bryophyte Flora of North America, Provisional Publication
Buffalo Museum of Science
1020 Humboldt Parkway
Buffalo, NY 14211 USA
www.buffalomuseumofscience.org/BFNA/bfnamenu.htm

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PTERYGONEURUM - POTTIACEAE

 

 xxx. PTERY­GO­NEURUM Juratzka, Laubm. Oest. Ungarn 95, 1882, conserved name and orthography  *  [Greek pterux, -ugos, wing + o + Latin neuron, nerve; alluding to the ridge-like costal lamellae]

Richard H. Zander

 

 

Plants bulbiform, gregarious or forming a thin turf, green above, light brown below. Stems buried, usually short, 0.5--5 mm; hyalodermis absent, sclerodermis absent, central strand present, distinct; axillary hairs ca. 7 cells in length, all hyaline or proximal 1--2 cells brown. Cauline leaves appressed when dry, weakly spreading when moist; ovate to ligulate, adaxial surface broadly concave, 2--3.5 mm (inclusive of the 1.5--2 mm awn); base not differentiated in shape; distal margins weakly to broadly incurved to near apex, plane below, entire or denticulate above; apex obtuse to rounded, often weakly cucullate; costa long-excurrent as a smooth or denticulate, hyaline awn, occasionally absent in some leaves, adaxial outgrowths of (0--)3--4 lamellae longitudinally inserted on adaxial surface of costa, ca. 12 cells in height, occasionally lobed or branching, smooth or hollow-papillose, adaxial cells (differentiated as lamellae) in 4--6 rows; transverse section circular, adaxial epidermis apparently absent, adaxial stereid band absent, guide cells 2 in 1 layer, hydroid strand present, usually large, abaxial stereid band present, often weak and round to elliptic in section, abaxial epidermis present; proximal cells differentiated across leaf base, occasionally higher medially, rectangular, little wider than the distal cells or somewhat inflated, ca. 2--5:1, walls of basal cells thin; distal medial cells quadrate to shortly rectangular, often transversely elongated, 10--13(--20) µm wide, (2--)1:1(--2), 1-stratose; papillae absent or small, simple, solid or hollow and present abaxially and medially, 2--3 per lumen, cell walls thin to weakly thickened and weakly trigonous, flat or somewhat bulging on both sides or just abaxially. Specialized asexual reproduction absent. Sexual condition autoicous or paroicous. Perichaetia terminal, interior leaves not sheathing, elliptic, little differentiated, to 1.5 mm. Seta 0.4-3.5 mm. Capsule stegocarpous or rarely cleistocarpous, theca short-cylindric, often macrostomous, variously smooth, sulcate or ridged, 0.5--1.5 mm, annulus when present of ca. 3 rows somewhat vesiculose cells; operculum rostrate to conic-rostrate; peristome teeth absent or when present rudimentary, in a latticework borne on a membrane, weakly twisted, teeth to 300 µm. Calyptra cucullate or mitrate with 3--4 large lobes. Spores 25-38 µm. KOH distal laminal color reaction yellow, occasionally red or orange in spots medially near apex or at leaf base.

 

Species 12 (4 in the flora): most continents, dry climate, soil or occasionally rock.

 

SELECTED REFERENCES Wareham, R. T. 1939. Pterigoneurum [sic]. In: A. J. Grout, Moss Flora of North America North of Mexico.1(4): 208--211. Zander, R. H. 1993. Genera of the Pottiaceae: Mosses of Harsh Environments. Bull. Buffalo Soc. Nat. Sci.32.

 

Pterygoneurum is apparently most closely related to Crossidium, and, like that genus, has chlorophyllose filaments (occasionally) in addition to the adaxial costal lamellae, and the stem is densely clothed with pale, brown rhizoids. Pterygoneurum is common in arid regions of the West, forming a dense, low turf in exposed soil. The hyaline awns and short setae are distinctive in the field.

 

 

1. Capsules immersed or partly exerted, calyptra mitrate.

2.Lamellae ca. 4--12 cells high; capsules immersed to somewhat exerted, stegocarpous (var. kieneri has an annulus but bursts irregularly)  . . . .  3. Pterygoneurum subsessile

2.Lamellae often low, 2--6(--10) cells high; capsules immersed, cleistocarpous. . . . . .   4. Pterygoneurum kozlovii

1. Capsules exerted on an elongate seta, calyptra cucullate or rarely mitrate.

3. Distal lamina smooth or rarely abaxially papillose; lamellae even in shape and size; capsules shortly exerted, ovoid; peristome lacking; cells of operculum in straight rows . . . .  .  2. Pterygoneurum ovatum

3. Distal lamina often abaxially weakly papillose; lamellae irregular in shape and size; capsules cylindric, long exerted; peristome occasionally present, fragile; cells of operculum slightly twisted . . . .  . .  1. Pterygoneurum lamellatum

 

1. Pterygoneurum lamellatum (Lindberg) Juratzka, Laubmfl. Oest. Ungarn 97. 1882

 

Tortula lamellata Lindberg, Oefv. K. Vet. Ak. Foerh.21: 233. 1864; Pterygoneurum arcticum Steere

 

 

Leaves with distal lamina sometimes weakly papillose abaxially; awn smooth; lamellae (0--)8--14 cells in height, irregularly lobed, often bearing filaments. Capsule stegocarpous, exerted, short- to long-cylindric, annulus present, operculum cells in twisted rows; peristome present, fragile and often adherent to the operculum. Calyptra cucullate.

 

Capsules mature late winter, spring. soil, rock faces; 500--1400 m; B.C., Nun.; Alaska, Ariz., Calif., Nev., Utah; Europe; Asia.

 

The operculum of P. lamellatum is commonly long-conic when dry, while that of P. ovatum is generally linear. The peristome may be trellis-like (illustrated by S. Flowers (1973) or a rudimentary arrangement of small plates over a membrane (R. H. Zander 1993: 202), or even absent. Also keying to P. lamellatum is the British Columbian specimen reported by T. T. McIntosh (1989) as Tortula wilsonii (Hooker) R. H. Zander. This proved to be a new species of Pterygoneurum distinguished by broadly recurved leaf margins and multiple papillae on both sides of the leaf above midleaf, these being unique traits for the genus. The adaxial costal lamellae are distinct but occasionally absent in some plants, which then phenocopy Tortula or Hilpertia. This specimen in turn proved to be the same as a Pterygoneurum discussed but not formally described by B. M. Murray (1992) for Alaska; a name will be provided by her in a future publication.

 

2. Pterygoneurum ovatum (Hedwig) Dixon

 

 Gymnostomum ovatum Hedwig, Spec. Musc.31. 1801; Pterygoneurum cavifolium Juratzka, illegitimate name

 

Leaves with distal lamina smooth or rarely papilose; awn smooth or rarely with a few teeth; lamellae 8--16 cells in height, not lobed, seldom bearing filaments. Capsule stegocarpous, emergent to exerted, ovoid, annulus present, operculum cells in straight rows; eperistomate. Calyptra cucullate or rarely mitrate.

 

Spores mature spring. soil (volcanic, dry saline), frost boil, low desert scrub areas; 900--1520 m; Alta., B.C., Man.**s**, N.W.T.**n**, Nun., Ont., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Kans., Mont., Nebr., Nev., N.Mex., N.Dak., Oreg., S.Dak., Tex., Utah, Wash., Wyo.; s South America, Europe, Asia, n Africa, Australia.

 

R. T. Wareham (1939) found the characters of var. incanum, long awns and short setae, inconstant in both American and European material. This is the most common species, and serves to stabilize arid soils (S. Flowers 1973). The setae may reach 3 mm. Faint thickenings reminiscent of a rudimentary peristome can sometimes be found associated with spore sac remnants dug out of the operculum, and the laminal distal cells may by simply papillose abaxially as in P. lamellatum.

 

3. Pterygoneurum subsessile (Bridel) Juratzka, Laubmfl. Oest. Ungarn 96. 1882

 

Pterygoneurum californicum H. A. Crum; P. subsessile var. henrici (Rau) R. T. Wareham

 

Leaves with distal lamina smooth; awn smooth or sharply serrulate; lamellae 10--12 cells in height, not lobed, sometimes bearing filaments. Capsules stegocarpous (or stegocarpous but bursting irregularly), immersed to emergent, short-ovoid, annulus present, operculum cells in straight rows; eperistomate. Calyptra cucullate or rarely mitrate.

 

Varieties 2 (2 in the flora): North Temperate Zone, s South America.

 

This is an abundant moss in the arid West, often occurring with P. ovatum. In some specimens, the perigoniate plants appear separate, but this species and doubtless others are apparently occasionally rhizautoicous. Following the reasoning of R. T. Wareham's (1939) discussion, P. subsessile var. henrici is placed with the typical variety. The characters associated (H. A. Crum 1967) with P. californicum are poor: the spores are finely papillose, the leaf cells do have weak collenchymatous thickenings, and the calyptra is long-mitrate.

 

1. Capsule dehiscent by the fallen operculum . . . . . . .  3a. Pterygoneurum subsessile var. subsessile

1. Capsule bursting irregularly, annulus differentiated but not fissile . . . .  3b. Pterygoneurum subsessile var. kieneri

 

3a. Pterygoneurum subsessile var. subsessile

 

Capsule stegocarpous, spores released after operculum falls.

 

Spores mature late winter, spring. soil (sandy, volcanic), alkali flats; 600--1700 m; B.C., Alta., Sask., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Ill., Iowa., Kans., Minn., Nebr., Nev., N.Mex., N.Dak., Okla., Oreg., S.Dak., Tex., Utah, Wyo.; s South America, Europe, Asia, n Africa.

 

3b. Pterygoneurum subsessile var. kieneri H. Habeeb, Rhodora 51: 128. 1949

 

Capsule facultatively cleistocarpous: annulus differentiated but operculum remains attached, spores released through ruptured capsule walls.

 

Capsules mature July. Grassland soil; 950 m; Nebr.

 

This taxon is P. subsessile with possibly hybrid sporophytes. Two forms with identical gametophytes were present in the type collection, one with typical sporophyte, the other with that of the variety. The operculum is not dehiscent and the capsule bursts irregularly. The sporophyte apparently falls as a unit, breaking off at the base of the seta. The single collection is from Nebraska, Chase Co., 12.5 km W of Champion, W. Kiener 10627 (MO).

 

4. Pterygoneurum kozlovii Lazarenko, Bot. Žurn. Ak. Nauk Uk. RSS 3: 61. 1946

 

Leaves with distal lamina smooth; awn smooth or sharply serrulate; lamellae 2--6(--10) cells in height, not lobed, sometimes bearing filaments. Capsules cleistocarpous, immersed to emergent, short-ovoid; eperistomate. Calyptra cucullate or rarely mitrate.

 

Capsules mature spring. Grassy areas, around alkaline depressions, steppe; moderate elevations; B.C.**s**, Sask.**s**; c Europe.

 

This locally abundant (T. T. McIntosh 1989) western species commonly grows in mixture with other species of the genus; although the gametophyte is much the same as that of P. subsessile, the brown, spheric cleistocarpous capsules with no annulus are unique. Perigonia are borne terminally on short lateral or basal branches.

 

OTHER REFERENCES  

 

Crum, H. A. 1967. A new species of Pterigoneurum [sic] from California. Madroño 19: 92--94.  

 

Flowers, S. 1973. Mosses: Utah & the West. Provo, Utah.  

 

McIntosh, T. T. 1986. The Bryophytes of the Semi-Arid Steppe of South-Central British Columbia. Dissert., University of British Columbia, Vancouver, B.C., Canada. 345 pp.  

 

McIntosh, T. T. 1989. Bryophyte records from the semiarid steppe of northwestern North America, including four species new to North America. Bryologist 92: 356--362.  

 

Murray, B. M. 1992. Bryophyte flora of Alaskan steppes. Bryobrothera 1: 9--33. 

 

Wareham, R. T. 1939. Pterigoneurum [sic]. In: A. J. Grout, Moss Flora of North America North of Mexico. 1(4): 208--211. Newfane, Vt.