Table 9.4. Structural and floristic characteristics of seven tropical montane cloud forests.
Location
Luquillo Mountains,
Puerto Rico
Volcan Barva,
Costa Rica
Blue Mountains,
Jamaica
Papua New Guinea
Ecuador, Eastern
Andes
Monteverde,
Costa Rica
Forest
Type"Colorado"
Lower
montane
wet
Lower
montane
rain
Mull ridgeLower
montane
rainLower
montane
rain
Lower
montane
wetAnnual
Elevation Rainfall
(m) (mm)725 37251500 34261550 30002500 39601710 nr1480 2500Plot
Size
(m*)400010,0001000 C600 d465 f40,000Tree
Density
(ha-^1 )185 a553 b52 b19 e28 e15078
396 h
1591
2062JBasal Area
(mVha)4029.2nr98nr11,88
9.6h
52.4^1- 8J
Tree
Species
Richness
(sp./ha)40653511959 e111SourceWeaver and
Murphy 1990Heaney and
Proctor 1990Tanner 1977Edwards
1977,
Edwards and
Grubb, 1977
Grubb et al.
1983This chapteraStems > 4 cm DBH.
bStems > 10 cm DBH.
cTen 10 x 10 m plots.
dThree 20 x 10 m plots.
eStems > 20 cm DBH.
fOne 61 x 7.6 m plot.
sStems 2-10 cm DBH.
also suggests disturbance is more frequent at Barva.
Thus, similar elevation and environmental conditions
do not necessarily dictate similar structure and floris-
tics in montane forests.Forest dynamics. The frequency and types of loss of
tree crowns and the death of whole trees affect forest
regeneration, nutrient cycling, and species richness.
Relatively few studies exist for tropical lower mon-
tane forests. The rates and frequency of tree damage
might be expected to be greater in higher elevation
forests because of steeper slopes, less stable soil, and
exposure to more wind. Small-scale forest distur-
bances such as loss of crowns (herein "tree damage")
and treefalls are determined by local climatic forces,
physical characteristics of the substrate, and biologi-
cal attributes of the trees (Putz and Brokaw 1989).
When whole trees and their associated epiphytes fall
to the forest floor, they (1) present pulses of organic
material and nutrients that can subsequently become
available to terrestrially rooted plants (Denslow 1987);
(2) increase the biomass of the forest floor, which
creates additional habitats for terrestrial organisms;
3) reduce resources used by arboreal animals and epi-phytes and create snags for nesting by key bird seed
dispersers (Wheelwright et al. 1984); (4) crush seed-
lings, saplings, and understory plants (Aide 1987);
and (5) affect microclimate of the ensuing gap, which
may subsequently deter or facilitate the germination
of seeds of some species (Putz and Milton 1982).
Although many damaged trees die, some continue
to live by producing new shoots from above- or below-
ground parts. Regeneration from broken plant seg-
ments ("resprouting") has been noted in some forest
trees (Clark and Clark 1989, R. Lawton, pers. comm.)
and shrubs (Kinsman 1990). Resprouting of damaged
individuals might replace lost substrate in the same
location and affect the form and duration of gap re-
generation faster than regeneration from seedlings.
Three censuses of marked trees in the leeward for-
est study area were made in September 1990, 1991,
and 1992 (Matelson et al. 1995). Tree damage and
mortality were divided into five categories: (1) stand-
ing broken stems, classified by the relative height of
the break (high, middle, low); (2) uproots (fallen trees
with exposed root wads); (3) knockdowns (trees fall-
ing as a result of a neighboring tree hitting them); (4)
standing dead trees (stems not broken or uprooted);318 Ecosystem Ecology and Forest Dynamics