258 A. Elizabeth Arnold
such as those of xylariaceous species and various
Eurotiomycetidae (see Holbet al.2004).
Inoculum volume plays an important role
in determinin gthe infection success of plant-
associated fungi (Agrios 1997), but until recently,
the rate of fungal propagule deposition in tropi-
cal forests was not known. Gilbert (2002), Arnold
(2002), and Arnold and Herre (2003) found that
typical leaves in the forest understory at Barro
Colorado Island (BCI), Panama, receive≥10–15
viable fungal propagules per cm^2 per hour dur-
in gthe mid- and late wet seasons. When adjusted
for mean leaf area and extrapolated to 24 hours,
these data suggest that the average leaf receives
more than 15,000 viable fungal propagules per
day (average based on mean leaf area for 28 tree
species in the understory at BCI; Arnold 2002).
These values consider only fungi capable of grow-
in gon one nutrient medium (malt extract a gar)
and likely underestimate total deposition.
Fungi in the forest air column represent
pathogenic, saprophytic, and endophytic species,
as well as numerous species of unknown eco-
logical roles. The proportion that can form
endophytic symbioses has not been quantified,
but the abundance of viable propagules in air
is positively associated with the frequency of
endophyte infections (Arnold and Herre 2003).
Propagules of fungi at BCI are highly abun-
dant in the air column immediately following
rainfall events, and then decrease as a curvi-
linear function with increasin gtime since si g-
nificant rainfall (Figure 15.2). Both ultraviolet
(UV) radiation and desiccation play an impor-
tant role in the mortality of fungal propagules:
Arnold and Herre (2003) found that the depo-
sition and persistence of livin gpropa gules on
leaves was greater under the forest canopy than
in the laboratory clearin gat BCI. After 1 week of
exposure, sterile plants placed in the lab clearing
accumulated significantly fewer endophyte infec-
tions than plants placed in the forest understory
(Arnold and Herre 2003).
ENDOPHYTE COLONIZATION AND
ABUNDANCE IN TROPICAL
LEAVES
The high abundance of inoculum in the air col-
umn, and the apparently universal receptivity
of tropical plants to colonization by endophytic
fungi (Arnold 2002, Van Bael et al.2005),
leads to high infection rates in mature foliage
of tropical trees. Arnold (2002) recovered endo-
phytic fungi from 100% of mature leaves sampled
5
10
15
20
25
020406080
R^2 = 0.967
Hours since rainfall
CFU per cm
2
per hour
Figure 15.2 Deposition of livin gfun gal
colony-formin gunits (CFU; hyphal fra gments
and both sexual and asexual spores) per cm^2 of
surface area per hour, as a function of hours
since significant rainfall in the understory of
secondary forest at BCI. Each point represents
the mean (±standard error) of six sampling
stations (data from Arnold 2002) and
underestimates total deposition: only fungi
capable of growing on a single medium
(2% malt extract agar) at a given temperature
(23◦C) are represented.