Family: Fissidentaceae

Synonyms

According to Pursell and Allen (1996, p. 75) “Bruggeman-Nannenga (1982) conditionally treated [Fissidens appalachensis] as a robust expression of F. pusillus (Wilson) Milde [= F. bryoides Hedw. var. pusillus (Wilson) Pursell in Allen].”

NatureServe Conservation Status

G2G3

Distribution

Endemic to eastern U.S.A. North Carolina, Pennsylvania, South Carolina, Tennessee.

Habitat

Aquatic in mountain streams, ranging from 1200 – 5600 ft elevation, tightly attached to crevices of submerged rocks in swiftly flowing water. Also aquatic or semi-aquatic attached to rocks (usually the lower edges near the stream bottom) in slow moving, shallow water of headwater streams and mountain springs/seeps. More rarely growing just above the water’s surface attached to a thin layer of sandy sediment atop low-lying rocks or stones in quiet streams. Plants may be covered by a foot or more of running water, in which case collecting specimens for documentation may require blindly picking moss bits from underwater or pulling smaller rocks above the water’s surface for inspection. In total submergence of swiftly flowing water F. appalachensis is more commonly found on the downstream nose of rocks especially in crevices of the undercut rock surface where presumably there is calmer water surrounded by eddies and turbulent flow.

In the upper Santeetlah Creek drainage near the headwaters of Johns Branch, robust F. appalachensis occurred on the upper surface of wet rocks (stones) that had accumulated a thin sandy layer of sediment in which the plants were attached. The sediment layer was surely the result of periodic submersion during high rainfall. Here material was collected by cutting the sandy sediment layer away from the solid rock underneath (see specimens Davison & Hicks 2970 and Davison 4003, the later specimen with abundant sporophytes; neither specimen was submerged at the time of collecting).

Brief Description and Tips for Identification

Plants green, becoming blackish below. Stems 5-15 mm long. Leaves in 5-22 pairs. Leaves limbate (with a differentiated border of elongate cells), the limbidium (border) extending to leaf apex or ending a few cells below. The limbidium of 2-5 layers of cells as seen in cross section. Description from Crum and Anderson (1981) and Pursell (2007).

Monoicous. Rhizautoicous or synoicous (Pursell 2007). The only published account of reproductive ecology is Smith (1984): “Well-developed sterile shoots form compact tufts isolated in cracks of rocks or in linear seams along rock fissures in the submerged zone. Fertile shoots occur frequently in the zones above or at the mean water-flow level. Sporophytes mature and spores are liberated from late July through September.”

The genus Fissidens is easily recognizable by its leaves. Unlike most other mosses the leaves are inserted in two rows, one on each side of the stem. Each leaf has a “pocket” or “slot” formed by portion of the leave blade that is split into what is called the vaginant lamina (one lamina lies atop the other with a small space between thus forming a sheath). Beever et al. (2002) explained other specialized terms used to describe the leaf morphology.

Anderson and Zander (1973) listed sixteen species of Fissidens occurring in the southern Blue Ridge Province of which only F. fontanus is truly aquatic. Fissidens appalachensis is typically fully submerged but may be found just above, or at the water’s surface. Regionally, F. appalachensis is distinguished from all other congeners (except F. bryoides) by a limbdium (a leaf border formed of elongate cells) that reaches the leaf apex.

F. appalachensis is potentially confused with forms of F. bryoides, especially F. bryoides var. pusillus that may co-occur with F. appalachensis on wet, streamside rocks. F. bryoides (var. pusillus) is so similar that Richard Zander in the publication naming F. appalachensis included a specimen as F. appalachensis that the late Ron Pursell (Fissidens monographer) reidentified as F. bryoides (see Pursell and Allen 1996). Distinctions setting F. appalachensis apart from F. bryoides var. pusillus include “usually larger plants, longer and comparatively narrower leaves, stronger limbidia, percurrent costae, and infrequently bistratose laminal cells” (Pursell and Allen 1996). In addition, Pursell and Allen (1996) emphasized a habitat distinction: F. bryoides grows on “surfaces of wet stones and rocks near the edges of streams” where submergence would occur only “for a time” during “periods of the year” (presumably during high rainfall). F. appalachensis grows “submerged in crevices among rocks in rapidly running streams” or partially exposed above water on the sides of emergent rocks.

Sérgio, C., & Pursell, R. A. (2001) included F. appalachensis in a complex of aquatic Fissidens that includes the very similar F. jansenii Sérgio & Pursell (Portugal), F. rivularis Schimp. (Europe), and F. geppii Fleisch. (Asia).

Salient Features

  • Aquatic habitat
  • Leaf margin with a band of differentiated cells forming a limbidium (border)
  • Limbidium strong
  • Leaves narrow (a relative concept, see images below comparing to F. bryoides)

References

Anderson, L. E., & Zander, R. H. 1973. The mosses of the southern Blue Ridge Province and their phytogeographic relationships. Journal of the Elisha Mitchell Scientific Society, 15-60.

Beever, J., Malcolm, B., & Malcolm, N. 2002. The Moss Genus Fissidens in New Zealand: Te Puninga Fissidens i Aotearoa: an Illustrated Key. Micro-Optics.

Bruggeman-Nannenga, M. A. 1978. Notes on Fissidens. I and II. I. On the arbitrary nature of the division in section of the genus Fissidens. II. Remarks on some species of the Fissidens bryoides complex. Proceedings Koninklijke Nederlandse Akademie van Wetenschappen. Series C, Biological and Medical Sciences 81: 38 7-402.

Crum, H. A., and L. E. Anderson. 1981. Mosses of Eastern North America (Vol. 1). Columbia University Press.

Pursell, R. A. 2007. 21. Fissidentaceae Schimper. Flora of North America, 27, 331-357.

Pursell, R. A., & Allen, B. 1996. A comparison of Fissidens appalachensis and F. bryoides var. pusillus. The Bryologist, 75-80.

Sérgio, C., & Pursell, R. A. 2001. Fissidens jansenii (Fissidentaceae; Bryopsida), a new aquatic species from Portugal. The Bryologist, 104, 378-381.

Smith, D. K. 1984. New characters and revised distribution of Fissidens appalachensis Zand. The Bryologist, 87, 259-260.

Zander, R. H. 1969. A new species of Fissidens from the Southern Appalachian Mountains. The Bryologist, 406-409.


Habitat

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Fissidens appalachensis

Type locality at Dixon Creek.

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Fissidens appalachensis

Type locality at Dixon Creek.

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Fissidens appalachensis

A small tributary to North Fork Catawba River, location reported in the original publication naming the species (Zander 1969).

Habitat

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Fissidens appalachensis

While not visible above, Porella pinnata also occurs at the point indicated. The inset shows much of the lamina eaten or eroded away while the thicker margin (limbidium) remains partially intact.

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Fissidens appalachensis

Upper portion of Falls Branch, Tennessee

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Fissidens appalachensis

Wolf Creek at Charlie's Creek Rd. Plants occur deeply submerged in the swifly flowing water shown. Plant also occur submerged further downstream where the creek banks are overtopped by evergreen rhododendron, yet there was a small light gap in the middle of the stream where plants were found.

Habitat

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Fissidens appalachensis

Photo attempts to capture the rocky springhead seep that feeds a short stretch of streambed shown in the next photo. Small rivulets in pencil size rock crevices support the species on this wet rocky slope. Roan Mtn., 5600 ft.

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Fissidens appalachensis

Here found on the lower edges of stones where plants stay permanently wet. Roan Mtn., 5600 ft. This stream soon dissappers underground downstream.

Habit

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Fissidens appalachensis

Charlie's Creek

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Fissidens appalachensis

Dixon Creek, type locality

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Fissidens appalachensis

Restricted to the lower crevices of the rock as indicated. The larger mat of bryophytes that are on top of the rock consist mostly of Scapania undulata with lesser amounts of Platylomella lescurii and Racomitrium aciculare. Wolf Creek.

Habit

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Fissidens appalachensis

Charlie's Creek

Morphology

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Fissidens appalachensis

Elongate leaves in two rows, each leaf with a "pocket" in the lower half as typical of Fissidens in general

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Fissidens appalachensis

Costa extends to the leaf apex as does the limbidium

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Fissidens appalachensis

Leaf cross-section showing thickened limbidium

Morphology

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Fissidens appalachensis

Shoots numbered above to correspond to the next three photo compositions

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Fissidens appalachensis

Material from type locality

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Fissidens appalachensis

Material from type locality

Morphology

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Fissidens appalachensis

Material from type locality

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Fissidens appalachensis

Material from type locality

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Fissidens appalachensis

Material from type locality showing variation in leaf tips from the same shoot

Morphology

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Fissidens appalachensis

Material from type locality

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Fissidens appalachensis

Limbidium not extending strongly to the leaf apex, otherwise the limbidium is quite strong

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Fissidens appalachensis

Notice further down the leaf the limbidium appoaches the costa in width

Morphology

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Fissidens appalachensis

Weak expression above and in next two images

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Fissidens appalachensis

Weak expression

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Fissidens appalachensis

Weak expression

Morphology

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Fissidens appalachensis

Leaves covered in diatoms

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Fissidens appalachensis

Portions of diatom covered leaves in cross-section

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Fissidens appalachensis

Limbidium, while not very wide, is confluent at leaf apex

Morphology

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Fissidens appalachensis

This material is from a site where historical specimens were verified by Pursell and Allen (1996). As the next two photo compositions show, the limbidium is not as stong as that from the type locality.

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Fissidens appalachensis

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Fissidens appalachensis

Distinguishing from F. bryoides

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F. appalachensis vs.

F. bryoides

Historical specimens from this locality (Armstrong Creek) were verified as F. appalachensis by Pursell and Allen (1996); however, recent collection of fully submerged plants proved problematic and are tentatively referred to F. bryoides. Two photo compositions follow illustrating the material in question.

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F. appalachensis vs.

F. bryoides

cf. F. bryoides

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F. appalachensis vs.

F. bryoides

cf. F. bryoides. The leaf border (limbidium) is rather weak compared to typical F. appalachensis

Distinguishing from F. bryoides, material from the type locality

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F. appalachensis vs.

F. bryoides

Shoots above were taken from the same palm-sized stone found laying in shallow quiet water (not totally submerged) in a small tributary perhaps 50 ft from the main channel of Dixon Creek. Shoot 1 is F. bryoides and shoot 2 is F. appalachensis as shown in the next two photo compositions. Shoots 3-10 are identified, with some uncertainty, in the images that follow.

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F. bryoides

Leaves are relatively wide and the limbidium fails to approach the costa in width

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F. appalachensis

Leaves are narrow and the limbidium is wide

Distinguishing from F. bryoides, material from the type locality

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F. bryoides

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F. bryoides

Compared to shoot 1, this shoot appears less certain as truly representing F. bryoides.

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F. bryoides

Distinguishing from F. bryoides, material from the type locality

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F. appalachensis

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F. appalachensis

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F. appalachensis