wright, "Sex Ratios," pp. 87-88, and Wheelwright, "A
Hypothesis," pp. 281-283). In contrast, demographic
studies have been common at La Selva Biological
Station (Clark 1994) and Barro Colorado Island (Leigh
et al. 1982, D'Arcy and Correa 1985).
Observing that falling branches commonly break
up understory plants in the cloud forest and that these
broken sticks sometimes take root, Kinsman (1990)
tagged broken branches of 28 species of understory
cloud forest shrubs and treelets, left them on the for-
est floor, and censused them annually over a period
of five years to record their survival. Fourteen species
remained alive for five years and some became re-
productive. DNA screening techniques were used to
investigate the role of branch fragmentation on the
population structure of shrubs in the cloud forest (see
Bush, "Clonal Reproduction," p. 88). High genetic re-
latedness within the colonies of the understory shrub
Poikilacanthus macranthus indicated that in large
part they were clones formed by fragmentation. Ad-
aptations for regeneration by fragmentation in species
of Rubiaceae also suggest a possible scenario for these
plants to evolve into epiphytes (see Nepokropff and
Sytsma, "Evolution in Psychotria," pp. 75-78). In land-
slides in the elfin forest at Monteverde, the standing
crop of seeds in the soil was much greater than the
number of seeds falling into traps (Myster 1993). The
taxa that colonize landslides at Monteverde (Table
3.4) are closely related to those found on landslides
in Puerto Rican cloud forests.3.4. BiogeographyThe lower montane zone at Monteverde (the Tilaran
Mountain mass above 1200 m) supports 1708 species
(57% of the total flora of the study area), which is
approximately equal to the native flora of La Selva
Biological Station (Hartshorn and Hammel 1994), but
much higher than that of Santa Rosa National Park,
Barro Colorado Island (Panama), and the lowland
Amazonian forest sites in Manii National Park (Peru)
and Manaus (Brazil) (Croat 1978, Foster 1990, Gen-
try 1990, 1993; Table 3.8). Monteverde's high diver-
sity of epiphytes, especially the 419 species of or-
chids, contributes most to this difference, but the three
life zones above 1200 m also support a large num-
ber of trees (500 plus species). There are more speciesTable 3.8. A comparison of the species richness of floras for several regions and research sites in the neotropics.
Site
Mexico
America north
of MexicoPeruCosta RicaTexasMonteverde area > 700 m
Manii National Park, Perub
Rio Manii, floodplain and
upland terrace
Monteverde area > 1200 m
La Selva Biological StationBarro Colorado Island,
Panama
Adolpho Ducke Forest
Reserve, Manaus
Santa Rosa National ParkAll Species
ca. 20,000
ca. 20,000b17,143aca. 9000ca. 50002985 d
2874
18561723 d
1678 d1207 Cca. 1030603Trees
NA
ca. 700NA>2000NA755
NA
NA442
ca. 312ca. 270NANAFerns
NA
NANA1058NA358
NA
NA175
162803730Orchids
NA
NA1587ca. 1500NA>500
NA
NA291
114782411Area (km^2 )
2,000,000
21,461,100283,52051,100692,408ca. 350
15,320
10058
1616100103Source
Rzedowski 1978
Missouri Botanical
Garden (www.mobot.
org)
Brako and Zaruchi
1993
Mora-Retana and
Garcia 1992,
Dressier 1993, Atwood,
"Orchids," pp. 74-
75, INBIO (www.
inbio.ac.cr)
Rzedowski 1978,
B. Hammel (pers.
comm.)
Haber 1991 (Appendix 1)
Foster 1990
Foster 1990Haber 1991 (Appendix 1)
Atwood 1988, Hartshorn
and Hammel 1994
Croat 1978, Hartshorn
and Hammel 1994
Prance 1990Janzen and Liesner 1980
aDoes not include ferns.
IncludesG ferns and bryophytes.
Native and naturalized species; cultivated species are not included.
Includes pteridophytes.67 Plants and Vegetation