Rolling mosses of Grimmia longirostris Hook. were studied in Páramo Piedras Blancas, a high Andean area in Venezuela, using a geoecological research framework focused on interactions between geomorphic, ecological, and soil processes. Spatial patterns of mosses were analyzed downslope from granitic outcrops on three slopes at 4445–4480 m elevation. Fragments from epilithic mosses on the apex outcrops were the source of other mosses. Distance of moss transport below rocks was measured at all sites; mosses on the gentlest (≤4°), 34 m-long, slope were censused along a belt transect on 17 contiguous quadrats; density of moss growth forms, stone size, percent of bare-soil cover, and surface soil properties were determined. Surface clast morphology and profile characteristics of a periglacial stone pavement at the slope base were also examined. Mosses on this site included four growth forms: epilithic mosses on outcrops; mobile, rolling moss balls; epilithic mosses on loose clasts; and unattached elongated mosses. Moss balls (density: ≤0.87 mosses/m2) lay on upper-transect plots over bare soils (≤92% cover) disturbed by needle ice. Soils here contained ~43–51% fine grains (≤0.063 mm) and 2.9–5.9% organic matter; this made them exceptionally prone to ice segregation and needle-ice formation. Lower-transect plots showed 78–100% stone cover—cobbles and blocks—but little exposed soil; mosses attached to loose clasts were exceedingly abundant (≤31/m2) on this section. Unattached elongated mosses, often resting along boundaries between adjacent stones, were also common (≤5.5/m2). The basal stone pavement showed a shallow (≤7 cm) rock veneer over a 24 cm-thick profile with exceptionally fine soil—up to 83.2% fine grains, ≤8.6% organic matter.A conceptual developmental model emphasizing the close linkages among the spatial distribution of mosses, patterns of surface clasts, and characteristics of soils and stone pavements is proposed: (i) Stones and mosses are transported downhill by needle ice and other frost agents; moss fragments gradually assume a globular shape, forming rolling moss balls. (ii) When these reach the stones and pavements on the basal slope, they cannot be transported any further, and remain immobile. (iii) Some mosses come to rest along boundaries between stones and develop into unattached elongated mosses or, (iv) they may become gradually attached to cobbles on the stone pavement; clasts with epilithic mosses can continue moving toward the basin floor. (v) Some mosses become fragmented and provide a source of elongated mosses, or remain partially attached to clasts. Stone pavements develop by a combination of rock accumulation following needle-ice transport, rock upheaving and sorting by ground frost, slope runoff, and downward soil illuviation.
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