Fog interception by Ball moss (Tillandsia recurvata)

Fog interception by Ball moss (Tillandsia recurvata)
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
www.hydrol-earth-syst-sci.net/15/2509/2011/
doi:10.5194/hess-15-2509-2011
© Author(s) 2011. CC Attribution 3.0 License.
Hydrology and
Earth System
Sciences
Fog interception by Ball moss (Tillandsia recurvata)
A. Guevara-Escobar, M. Cervantes-Jiménez, H. Suzán-Azpiri, E. González-Sosa, L. Hernández-Sandoval,
G. Malda-Barrera, and M. Martı́nez-Dı́az
Universidad Autónoma de Querétaro, Santiago de Querétaro, Querétaro 76010, México
Received: 1 October 2009 – Published in Hydrol. Earth Syst. Sci. Discuss.: 2 March 2010
Revised: 15 July 2011 – Accepted: 5 August 2011 – Published: 12 August 2011
Abstract. Interception losses are a major influence in the
water yield of vegetated areas. For most storms, rain interception results in less water reaching the ground. However,
fog interception can increase the overall water storage capacity of the vegetation and once the storage is exceeded,
fog drip is a common hydrological input. Fog interception
is disregarded in water budgets of semiarid regions, but for
some plant communities, it could be a mechanism offsetting evaporation losses. Tillandsia recurvata is a cosmopolitan epiphyte adapted to arid habitats where fog may be an
important water source. Therefore, the interception storage
capacity by T. recurvata was measured in controlled conditions and applying simulated rain or fog. Juvenile, vegetative
specimens were used to determine the potential upperbound
storage capacities. The storage capacity was proportional to
dry weight mass. Interception storage capacity (Cmin ) was
0.19 and 0.56 mm for rainfall and fog respectively. The coefficients obtained in the laboratory were used together with
biomass measurements for T. recurvata in a xeric scrub to
calculate the depth of water intercepted by rain. T. recurvata contributed 20 % to the rain interception capacity of
their shrub hosts: Acacia farnesiana and Prosopis laevigata
and; also potentially intercepted 4.8 % of the annual rainfall. Nocturnal stomatic opening in T. recurvata is not only
relevant for CO2 but for water vapor, as suggested by the
higher weight change of specimens wetted with fog for 1 h at
dark in comparison to those wetted during daylight (543 ± 77
vs. 325 ± 56 mg, p = 0.048). The storage capacity of T. recurvata leaf surfaces could increase the amount of water
available for evaporation, but as this species colonise montane forests, the effect could be negative on water recharge,
because potential storage capacity is very high, in the laboratory experiments it took up to 12 h at a rate of 0.26 l h−1 to
reach saturation conditions when fog was applied.
Correspondence to: A. Guevara-Escobar
([email protected])
1
Introduction
Atmospheric bromeliads have developed the ability to survive in environments where the rain period is limited. Also,
throughout the dry season they show asynchrony in the leaf
phenology compared to the rest of the community, owing to
distinctive anatomical and physiological traits (Barradas and
Glez-Medellin, 1999). Their crassulacean acid metabolism
(CAM), is characterized by the stomatic absorption of CO2
during darkness, as well as restricted water lost from transpiration (Nobel, 1983). Regarding bromeliads, leaf water
is coupled to the atmospheric water vapor, but the degree
of coupling depends on life form, microclimate and vertical strata within the canopy (Reyes-Garcı́a et al., 2008b).
The bromeliads absorb water through specialized structures
such as foliar trichomes and stomata, when the level of atmospheric water is high or during periods of nocturnal fog
(Benzing, 2000; Reyes-Garcı́a et al., 2008b). Atmospheric
species of bromeliad colonize the tree canopy, rocks and even
cable lines and therefore, the only water available to them
is that detained on their surfaces and the atmospheric vapor
available in the limit layer. The role of trichomes is very
important for water detention; when the humidity content in
the plant is low, the trichome wings are elevated and when
moist, they are folded and stick to the leaf surface (Schmitt
et al., 1989; Stefano et al., 2008). When the trichomes’ wings
are folded a reduction in the contact angle with rain or fog
drops is possible and the runoff from the surface could be decreased. Although the atmospheric bromeliads do not have
roots to absorb water or a tank to capture rainfall, the number
of narrow leave are enough to capture the fog and help satisfy their water requirements (Martorell and Ezcurra, 2007).
Reyes-Garcı́a et al. (2008a) suggested that dew and fog interception are important mechanisms because the main photosynthetic activity of atmospheric bromeliads is during the
rainless time of the year.
Published by Copernicus Publications on behalf of the European Geosciences Union.
2510
On the other hand, hydrological understanding has advanced substantially during recent decades, particularly regarding the natural evaporation process (Shuttleworth, 2007).
A main component of evaporation is rainfall interception loss
generated from the vegetation canopy (Dunkerley, 2000).
Measurement of rainfall interception has been investigated
in temperate and tropical forests, but studies in semiarid
shrubland or grasslands are scarce (Crockford and Richardson, 2000). Nevertheless, Návar and Bryan (1990, 1994)
and Návar et al. (1999) showed that interception losses in a
thorny scrub could exceed 27 % of the annual precipitation.
For dry climates, the magnitude of interception is important
with respect to the annual rainfall, the shortage of water resources and the temperature increases due to global warming
(Méndez et al., 2008).
As with rain, fog interception research mainly describes
tropical and temperate forests. In the Tropical Montane
Cloud Forests fog interception contributes up to 154 % of
annual rainfall and, is an important process during the dry
season (Bruijnzeel, 2001, 2005). The interception capacity
of epiphytes is high but only a fraction of the potential storage could be actually available, because the mosses are usually close to saturation or most of the moss biomass occurs at
sheltered positions in the lower part of the canopy (Hölscher
et al., 2004).
Fog interception measured on epiphytes was correlated
with that measured from fog gauges but was more than an order of magnitude smaller than the actual measurements from
fog gauges (Villegas et al., 2008). By monitoring changes in
their biomass, the epiphytes are used as biosensors to better
quantify the magnitude and mechanisms of fog interception
(Mulligan et al., 2011). For dry climates this measurements
are more important because the small ammounts of atmospheric water, fog or dew are paramount for the biotic diversity (Brown et al., 2008). The availability of water in dry
climates is not abundant, but probably is sufficient to establish an independence from the soils’ water relations; as in the
case of epiphytic bromeliads (Reyes-Garcı́a et al., 2008b).
Maximum water content of epiphytes has been assessed
by submerging samples in water or applying simulated rain
(Pypker et al., 2006). Also, there is a discrepancy in specific
storage measured by dipping vegetation into water compared
to sprinkling experiments, especially when water is blotted
from foliage to simulate windy conditions; storage measured
this way is nearly an order of magnitude lower than under
rainfall simulation (Keim, 2006).
Interception loss is the amount of rainfall detained and
subsequently evaporated from the vegetation, the boundary
layer conductance and canopy storage capacity are the most
important parameters in all models of rainfall interception
loss by forest canopies. Rutter et al. (1971) defined the storage capacity as the depth of water which can be stored or
detained on the plant surfaces in still air. Canopy storage is
added to by intercepted rainfall and depleted by evaporation
and drainage (Rutter et al., 1971). Two primary methods to
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
A. Guevara-Escobar et al.: Ball moss fog interception
estimate interception losses are: (i) calibrated process-based
models of interception and evaporative loss and (ii) direct
measurement (Dunkerley, 2000). The measurements of interception presents some problems because storage is modified by intensity of rainfall, wind speed and direction (Crockford and Richardson, 2000). In the case of fog, Villegas et al.
(2008) showed that interception rates by Ball moss (Tillandsia recurvata) were sensitive to the interaction between low
levels of wind speed and liquid water content of fog. The
interaction between wind speed and fog interception rate results from vertical settlement at low speeds and advectional
impactation at high speeds.
The storage capacity parameters are better identified when
evaporation is low (Vrugt et al., 2003). Since radiation is low
during rainfall, the evaporation rate during storms is predominantly determined by the aerodynamic conductance (Rutter
et al., 1971). Including only storms under zero evaporation
conditions will yield a value which is a better estimate of the
canopy storage than considering storms with non-zero evaporation (Gash, 1979). Storage parameters are calculated using measurementes of the weight gained by a specimen that
is exposed to simulated rain, preferably in still air and for
saturated surfaces (Dunkerley, 2000).
Interception losses are related to precipitation characteristics, evaporation rate and the amount of water stored on vegetation surfaces (C). Two parameters of interception storage
are important: maximum storage (Cmax ), which is the water stored when drainage rate is constant, Cmax includes water temporarily stored and that would be removed by gravity
and; residual or minimum storage (Cmin ), that depth of water
removed only by evaporation (Pitman, 1989; Putuhena and
Cordery, 1996). The value of Cmin is equivalent to soil field
capacity and also corresponds to the minimum quantity of
water required to wet all the canopy surfaces (Rutter et al.,
1971).
In the present work, we used direct methods to measure
interception storage parameters of T. recurvata in laboratory
experiments. Its objectives were to: (i) determine the storage
capacity during daylight using three wetting methods: simulated rain, simulated fog and soaking; (ii) simulate a short
duration fog event during daylight and follow the weight loss
for 12 h; (iii) assess the effect of daylight or dark conditions on the storage capacity of Ball moss and on weight loss
after seven days post-event. The objectives would answer
what is the potential storage with different wetting methods;
how much water would be available at evening after an early
morning fog event and; weather or not stomata opening is
important in the water relations of Ball moss.
This study would improve our knowledge on water balance and ecology of semiarid shrublands because the aerial
epiphyte communities, which grow in association with tree
structures enhance interception of fog and rainfall (Villegas
et al., 2008). Also, Bromeliad species as such as T. recurvata
not only compete for space and light, but intercepted precipitation could have an influence on the amount of available
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A. Guevara-Escobar et al.: Ball moss fog interception
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Fig. 1. Monthly means and standard errors for 24 h records of: (a) air temperature (T ), (b) relative humidity (RH), (c) net radiation and,
(d) rainfall at the study sites.
water and under severe invasion the growth of host species
may be reduced.
2
2.1
Material and methods
Sites
The sites from which the data were taken, located in the Central Highlands of México are described as thorny scrub with
Prosopis laevigata and Acacia farnesiana as dominant shrub
species, both being phorophytes of T. recurvata. The altitude
varies from 1959 to 1990 m a.s.l. (above sea level). The study
area is classified in the Koeppen’s Climate system as BS1 k.
During 2006, meteorological stations located at 20◦ 430 N,
99◦ 470 W (site A) and 21◦ 130 N, 100◦ 470 W (site B), measured rainfall, temperature (T ), relative humidity (RH) and
wind speed using a WXT510 multi-sensor (Vaisala, Helsinki,
Finland). Net radiation was measured with a Q7.1 net radiometer (Campbell Scientific Ltd., Shepshed UK). These
sensors were connected to a CR1000 datalogger (Campbell
Scientific Ltd., Shepshed UK), averaging at a 1 min time step.
The average annual T and RH prevailing at the study sites
were 14.9 ◦ C and 51.7 %. Figure 1 presents monthly means
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for 24 h for T and RH, rainfall and net radiation. During the
early hours of the morning (04:00 a.m. to 08:00 a.m.), T and
RH were 11.1 ◦ C and 79.7 %; from 08:00 a.m. to 04:00 p.m.,
after the potential fog period, values were 21.4 ◦ C and
36.1 %. Mean wind speed was 0.68 m s−1 and net radiation
was −32 W m−2 during the early hours.
Dew point temperature was calculated using the MagnusTetens formula; the coefficients used were according to Murray (1967). Biomass annual production of T. recurvata reported by Olalde and Aguilera (1998) at site B was used to
scale up laboratory measurements. Plant material used in the
experiments was collected from a site close to the laboratory
facilities (20◦ 430 N, 100◦ 240 W).
2.2
Plant material
Juvenile, vegetative specimens of T. recurvata were chosen
because the trichomes of some bromeliads are reduced in frequency and dimension as the plant reaches the adult form
(Stefano et al., 2008). This kind of specimens would represent the upper bound of water storage. Specimens were
collected during the dry season, at morning hours and on the
programmed day’s experiment.
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
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A. Guevara-Escobar et al.: Ball moss fog interception
The plants used in the experiments were collected from
the field at random but considering that the fresh mass of the
specimen was within an established confidence interval. The
fresh mass of 30 plants was measured and a 99% confidence
interval was constructed (x̄ ± sx̄ tα/2=0.005, gl=29 where: x̄ is
the fresh mass mean, sx̄ is the standard error of the mean and
t the corresponding Student’s t-distribution value for a 99 %
confidence and 29 degrees of freedom).
2.3
Storage capacity: simmulated fog
The storage capacity of T. recurvata was directly determined
in a laboratory where conditions of T and RH were stable,
similar to those present in the study sites A and B during
daylight (21.4 ◦ C and 36.1 %). The plant was suspended by a
0.12 mm nylon line hooked to an electronic balance. A copper wire forming a hook at the end of line secured the specimen. The mass of the line and hook was 24 mg. The plant
mass (W ) was measured in 5 mg steps to the nearest 1 mg.
Data were acquired via the RS232 microcomputer port and
using Bytewedge Pro version 3.3 (Fog software, Inc.). To
control the fog spray a 50 × 55 cm bell-shaped polystyrene
chamber was used (Fig. 2). The scale was located in a platform above the chamber and a 3 mm opening in the top of the
chamber allowed movement of the line without obstruction.
The chamber was placed on a metal base with several connection openings and a fog spray vent. Fog was produced by
an ultrasonic humidifier at a rate of 0.26 l h−1 and 0.0004 mm
mean drop size (Elehum 002, Sunshine, EM, México). A
timer switched on and off the humidifier as required. Inside
the chamber a Hobo Pro sensor (Onset Corp., Bourne, MA)
recorded T and RH every 5 min. A petri dish with water was
placed inside the chamber to satisfy evaporative demand during the draining phase of the experiment. Pypker et al. (2006)
in their experiments of rainfall interception by epiphytes used
a similar conditions (high RH and 22 ◦ C temperature).
Ten plants were individually wetted by simulated fog during 12 h. Constant weight was reached during the last 3 h.
After the wetting period ended another 12 h were allowed for
the draining phase. Maximum storage capacity was calculated as:
Cmax = Wfmax − Wf0 .
(1)
where Wf 0 [mg] was the plant mass before the wetting phase
and Wf max [mg] was the plant mass at saturation. The minimum storage capacity was calculated as:
Cmin = Wfmin − Wf0 .
(2)
where Wf min [mg] was the plant mass at the end of the
draining phase and assuming that evaporation was negligible. At the end of each run the plant dry weight (Ws ) was
obtained by oven-drying at 60 ◦ C for 48 h. Water content
was determined as:
H = Wf0 − Ws .
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
Fig. 2. Diagram of the fog simulation setup (1) computer, (2) electronic scale, (3) nylon line, (4) chamber, (5) Tillandsia recurvata sample, (6) humidity and temperature sensor, (7) petri dish,
(8) metallic base, (9) humidifier, (10) timer.
2.4
Storage capacity: simmulated rain
Ball moss within shrub canopies is mainly found at the mid
and lower branches (Garcı́a-Suárez et al., 2003) and therefore, throughfall interception may be more important than
rainfall interception. Drop size distribution of secondary
drops ranges from 0.5 to 5.5 mm and is characteristic of particular tree species (Calder et al., 1996). Throughfall had
different drop size distribution among canopy species under
conditions of little canopy vibration with low rainfall intensity and wind speed; but throughfall contained smaller drops
under conditions when rainfall intensity was high (Nanko
et al., 2006). Convective high intensity rainfall is typical in
the region; therefore, small, low velocity drops were used.
Thirty specimens were used to estimate the mean minimum interception storage capacity by rain. Spray was horizontally applied over the specimen to produce low velocity
drops, representing throughfall within the tree canopy that
could be intercepted by epiphyte vegetation. Two manual
sprayers were operated at a constant time rate. Rainfall was
measured with a TE-525LL-L tipping bucket (Texas Electronics Inc., Dallas TX) calibrated to record 0.254 mm per
tip. Rainfall intensity was 70 to 79 mm h−1 and drop diameter was 1.8 ± 0.2 mm. The simulation stopped when the
weight of the wetted specimen was constant. The drying
phase was 12 h or until drainage finished.
(3)
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A. Guevara-Escobar et al.: Ball moss fog interception
2.5
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Storage capacity: soaking
Some authors have reported the maximum water holding capacity (S) as the amount of water detained by a material after
soaking for a period of time and a draining phase (Sato et al.,
2004; Pypker et al., 2006). For T. recurvata it would be expected that any air trapped in the surface irregularities was
eliminated and then S could be different from Cmax or Cmin .
Conceptually, S would be similar to Cmin if the wetted surface was smooth, because both represent water storage after
saturation and draining. However, water fluxes must be different for rough surfaces when wetted by soaking, rain or
fog.
A sample of ten plants was used. Values of S were determined by measuring Wf 0 and then suspending the plant as
previously described. A 500 ml container was placed below
the specimen and filled using a venoclysis and a syringe until the plant was immersed. The container was emptied by
gravity after 3 h and the plant immediately weighed to assess
Wf max . The mass after 12 h of draining represented Wf min ,
assuming that evaporation was negligible. Values of S were
determined as:
S = Wfmin − Wf0 .
2.6
2.8
Interception scaling up
Potential storage capacities of T. recurvata in the vegetation
were calculated as a function of Wf 0 (Pitman, 1989):
C0 =
C
.
Wf0
(5)
S0 =
S
.
Wf0
(6)
(4)
Short duration fog event
The experiments described were designed to estimate the
values of Cmax , Cmin and S. However, field conditions in
the thorny scrub are adequate for fog formation only during
few hours of the day and, not every day (Garcı́a-Garcı́a and
Zarraluqui, 2008). In this experiment the evolution of C was
followed during 1 h of wetting with simulated fog and 12 h
drying phase. The C determined at the end of the experiment
would represent the available water on the specimen surface
at the early night when stomata could open. Any water available for evaporation at the plant surface would reduce evaporative demand because stomatal control of transpiration is
strong only when boundary layer conductance is high in relation to stomatal conductance. Also, if no water was left after 12 h, then water absorption through stomata or trichomes
could only take place during the simulated fog event.
Twelve plants were randomy asigned to two treatments:
control and wetted with fog during daylight. Afterwards, the
change in fresh mass (1Wf ) was recorded for seven days
for all plants. Plants were placed in the laboratory where T
was 22 ◦ C and RH was ≤30 %; these were similar to field
conditions from 08:00 a.m. to 04:00 p.m.
2.7
During the day, stomata would be closed and therefore, plant
mass should be lower than that of plants wetted at dark.
We assumed that foliar trichomes absorve fog water and are
functional following light or RH daily cycles. Schmitt et al.
(1989) determined that bromeliad may take up water from
the gas phase of the atmosphere, when the RH increases, by
equilibration of the hygroscopic cell walls of the dead scale
cells in the trichomes. These authors concluded that this gain
in water is lost when RH decreases at the begining of the
following day, so bromeliad do not have a net gain of water from this mechanism. To cast some light on this aspect,
six specimens were fog wetted from 05:00 to 06:00 a.m. at
dark; another six were wetted during daylight, but at the same
time schedule. 1Wf was recorded during the following seven
days.
0 , C0
0
−1
where Cmax
min and S have units mg mg . Afterwards,
mean values were multiplied by the biomass reported by
Olalde and Aguilera (1998) to obtain the intercepted depth
of water [mm].
Based on the rain wetting experiment, the estimation of
annual interception was the result of the number of rain
events greater than 0.19 mm multiplied by the storage capacity, 0.19 mm. Rain events smaller or equal to 0.19 mm were
added up. Rainfall events were identified as storms separated
by periods of time longer than 24 h.
2.9
Statistical analysis
The relationship between storage capacity and biomass was
explored by regression analysis and using Table Curve 2-D
v 5.01 (Sistat Software Inc.). ANOVA and the Tukey test
were performed to estimate the difference among S and C
values. A repeated measures model was used to assess differences regarding 1Wf over time. The level of significance
was fixed at α = 0.05. All tests were performed using the
GLM and MIXED procedures of SAS (SAS Institute, Cary,
NC, USA).
Day and night fog events
All the tests described were performed during daylight.
However, the values of C and S correspond to live specimens of T. recurvata and it was impossible to partition the
water stored on the plant surfaces from that probably absorbed via stomata and trichomes during the wetting phase.
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3
3.1
Results and discussion
Storage capacity
Three wetting methods were applied to determine the water storage capacity of Ball moss and, there were differences
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
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A. Guevara-Escobar et al.: Ball moss fog interception
2.5
Table 1. Interception storage capacity of T. recurvata in laboratory
conditions.
Fog
Mean
SE∗
Rain
Mean
2.0
SE∗
-1
SE∗
mg
Wf 10
Ws2
S3
4
Cmax
4
Cmin
1953
709
1783
297
29
353
C' (mg mg )
Soaking
Mean
1453
482
116
126
2110
873
191
92
2788
1610
484
329
942
444
1.5
1.0
0.5
mg mg−1
S 05
C0 5
max
C0 5
0.89
0.0
0.09
2.04
1.41
min
0.26
0.19
0
0.47
0.02
mm
S
4
Cmax
4
Cmin
4
6
8
10
12
Hours
Fig. 3. Means and standard error of stored water in T. recurvata
during 12 h wetting by simulated fog at ≥90 % RH and 22 ◦ C.
0.36
0.81
0.56
2
0.19
1 :fresh weight, 2 : dry weight, 3 : maximum storage capacity, 4 : the maximum and minimum interception storage capacities, 5 : the capacities per unit of fresh mass, ∗ : standard
error of the mean.
between the methods for the amount of water stored after a
draining phase was allowed. Soaking the specimen was not
representative of rainfall or fog events because the storage
0
parameter S 0 was different from Cmin
under simulated fog or
rain (p = 0.03); therefore, the soaking method would not be
advisable for water relations studies of species like T. recurvata. Table 1 presents the storage coefficients obtained in the
laboratory.
Leaf and trichomes traits could explain the difference be0
tween values of Cmin
in relation to S 0 . Sato et al. (2004)
reported similar S for leaf litter of species with trichomes or
glabrous, but Cmin values were higher for the species with
trichomes, they suggested that drop size and aggregation are
important for storage capacity and drainage. In the present
work, it was possible that coalescence increased drainage in
the immersion tests for S 0 but this phenomena probably was
not as important as in the case of fog simulations, due to
0
the smaller drop diameter; thus, Cmin
was also higher for
fog compared to rain (p ≤ 0.0001). Although the dominant
forces contributing to water storage are gravity and capillary adsorption-tension, viscous forces also are an important component in water retention, as in the case of surfaces
where the flux is slow and with a high roughness coefficient.
Contact angle is an indicator of water repellency of a surface
and this trait has great ecological importance. Droplet contact angle and droplet retention are inversely proportional to
the trichome density (Pandey and Nagar, 2003). In the case
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
of T. recurvata trichome wings movement during the wetting
phase also would have to be considered to explain drainage
fluxes and storage capacities.
According to these results, water was better captured by
T. recurvata when fog forms. However, natural fog rate in
the study site probably was much smaller than the flows generated during the experiments, mainly because fog events are
shorter in duration than the 12 h period used in the experiment. Fog events could last less than three hours in the
study sites according to the temperature records. For example, stored fog during the first hour of simulation was 50 %
of Wf 0 and by the third hour it was 120 % (Fig. 3).
Jarvis (2000) reported the storage capacity of epiphytics
after 30 h of fog simulation at 6.4 l h−1 as 5.94 times their
dry weight whereas in the present work it was 4.53 times after 12 h at 0.26 l h−1 . Applying a lower rate, similar to those
reported in nature, would result in longer test runs. More
importantly, the result suggest that for T. recurvata, fog interception could not be a transfer mechanism, through drainage,
beneficial to terrestrial species and therefore, the ecological
relevance of fog interception by T. recurvata probably is indirect, if any.
Finding an alometic relationship for the storage parameters is important to scale up the storage capacities. Leaf area or number of leaves are cumbersome
to measure, but total biomass was an easily measurable
trait of T. recurvata and in the fog simulation tests Wf 0
was related to Cmax and Cmin (Cmax = −728.77 + 2.11 Wf 0 ,
r 2 = 0.52, p = 0.02; Cmin = −790.91 + 3.14 Wf 0 , r 2 = 0.56,
p = 0.02). In the case of simulated rain the relationship was
Cmin = 129.34 +0.38 Wf 0 , r 2 = 0.79, p ≤ 0.0001. For these
relations the explained variance by Wf 0 was low. The relationship could improve increasing the number of samples
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A. Guevara-Escobar et al.: Ball moss fog interception
0.4
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Table 2. Weight and storage of T. recurvata after a one hour fog
event at daylight or dark in laboratory conditions.
-1
C' (mg mg )
0.3
0.2
0.1
Wf 10
C2
1Wf 3
mg
mg
mg
C2
1Wf 3
mg mg−1
mg mg−1
Sig.∗∗
Daylight
Mean
SE∗
Dark
Mean
SE∗
1677
325
−244
256
56
78
2908
542
−28
541
77
63
n.s.∗∗∗
0.05
0.03
0.215
−0.134
0.04
0.04
0.201
−0.001
0.03
0.02
n.s.
0.01
1 : fresh biomass, 2 : storage, 3 : the change in fresh mass, ∗ : standard error of the mean,
∗∗ : Significance, ∗∗∗ : not significative at the 0.05 level.
0.0
0
2
4
6
8
10
12
Hours of drying
Fig. 4. Means and standard error of stored water in T. recurvata
after 1 h wetting by simulated fog at ≤30 % RH and 22 ◦ C.
or including other other traits, such as number of leaves or
density of trichomes (Martorell and Ezcurra, 2007). Biomass
dry weight (Ws ) was not related to Cmax or Cmin because the
plant water content was variable, despite the relatively constant ambient during the dry season and phenological development of the specimens.
3.2
Short duration fog event
There was no statistical difference in the daily weight
change rate during one week of drying between plants fog
wetted for 1 h (−29.4 ± 6.6 mg d−1 ) and those not wetted
(−15.5 ± 6.0 mg d−1 ). Fifty percent of the intercepted water, during 1 h of fog wetting, was lost after 12 h of drying
at low RH (Fig. 4). Because the wetting occurred during
daylight there was little opportunity for water to be absorbed
through stomata and probably very little of the water remaining on the plant surfaces after 12 h was useful for the plant.
The stored water inside the plant before the experiment was
the most likely source of water for the observed change in
weight.
3.3
Day and night fog events
Specimens wetted for 1 h at dark gained more weight in comparison to wetting during daylight (C, Table 2). However,
both treatmens stored a similar ammount of water per unit of
fresh mass (C 0 , 0.215 vs. 0.201 mg mg−1 ) immediately after
the fog simulation ended. At this point during the experiment both treatments had a similar amount of water “stored”
but probably the partition was different.
Specimens wetted at dark lost less weight after seven days
of drying following the fog simulation. This result was
interpreted as higher water detention inside the plant as a
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consequence of nocturnal stomatic opening during CO2 assimilation since T. recurvata is a CAM species. Proportional
to the initial fresh weight, the weight loss represented −1 %
and −15 %, for dark and light conditions respectively.
These results showed that T. recurvata water relations depended upon recurrent conditions favorable for fog formation
or condensation. The experiment at dark also suggested that
stomata opening is more important than foliar trichomes regarding water relations. Schmitt et al. (1989) showed that
the patterns of water uptake and loss are similar in dead or
alive T. recurvata during day-night cycles with respectively
low and high RH. In studies of leaf water isotopic enrichment Reyes-Garcı́a et al. (2008b) also demonstrated the importance of water vapour exchanges at high humidity for epiphytic bromeliads. In our experiment ambient RH was low,
temperature constant and water condensation was not possible; therefore, the plants could not gain weight from ambient
water during the seven day drying phase.
3.4
Extrapolation to natural condition
Annual rainfall during 2006 was 732 and 770 mm at sites A
and B. González-Sosa et al. (2009) reported 30 and 20 %
rainfall interception by the canopy of A. farnesiana and
27 and 21 % by P. laevigata. Standing biomass of T. recurvata was 4000 ± 480 kg ha−1 (Olalde and Aguilera, 1998).
Scaled up Cmin for rainfall was 0.19 mm for T. recurvata (Table 1) and, 0.93 mm for the host shrubs A. farnesiana and
P. laevigata (Mastachi, 2007).
We derived the annual interception of T. recurvata as 4.8 %
(35.8 mm) of annual rainfall. The biomass used in the calculation was important, as the site was considered infested by
Ball moss, also juvenile speciments are more likely to intercept a higher ammount of rainfall. Five percent interception
loss represented the high bound figure and, still it was relatively small amount in terms of annual rainfall at the sites.
In terms of fog, 57 min were needed to intercept the same
ammount as the storage capacity for rain (0.19 mm). This
value was obtained scaling up to area basis the data presented
in Fig. 3 and using the relationship C = −0.83 + 0.25 ln(t),
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
2516
A. Guevara-Escobar et al.: Ball moss fog interception
Fig. 5. Mean and standar error of (a) air temperature (◦), dew point temperature (•) and (b) relative humidity (RH) for morning hours 04:00
to 08:00 a.m. at the study sites.
r 2 = 0.98, p ≤ 0.0001; where t was time [min] and C was the
stored water [mm].
The relative importance of fog relative to rainfall interception in field conditions needs to be investigated. Using Ball
moss as real time biomonitor would be the best option, because fog gauges produce an overestimate (Villegas et al.,
2008). Our results showed that Ball moss has a high monitoring capacity, as represented by the storage capacities and;
good resolution over time, as showed by the dark experiments and the weight loss follow ups. An apparatus similar
to that presented by Mulligan et al. (2011) for mosses could
be designed for such monitoring with Ball moss.
Interpretation of the present data depends on the real fog
amount and adequate conditions for vapor condensation in
the air surrounding the shrub vegetation. Fog happens when
the temperature and dew point are equal or within a degree
(Gultepe et al., 2007), but during the rainy season the difference between T and Tr was 1.1 to 1.9 ◦ C from July to November at 04:00 a.m. to 08:00 a.m. (Fig. 5). However, it should
be considered that bromeliad leaf temperature is lower than
air temperature at dawn (Andrade, 2003), thus condensation
over surfaces could occur in absence of visible fog. Another
factor important in fog formation is the evaporation from soil
moisture and dew when the surfaces heat up (Gultepe et al.,
2007). Higher T and Tr suggested that rainfall could be more
important than fog for Ball moss water relations at the selected sites. Accordingly, Andrade (2003) concluded that
dew deposition was not adequate to support growth for epiphytic bromeliads during the driest months of the year in a
tropical dry forest. On the other hand, Martorell and Ezcurra
(2002) identified fog belts at intermediate altitudes as a main
determinant of species distribution and diversity in the rosette
scrub in arid mountains (including bromeliads), altitude and
temperature being of lesser importance. In either case, fog
biomonitoring would be desirable, and growth should independently measured.
Hydrol. Earth Syst. Sci., 15, 2509–2518, 2011
3.5
Considerations and implications
Drop dimension of the simulated fog was considered representative of natural fog because the effect of drop size is
very small at these sizes, according to the model of Calder
et al. (1996). Simulated fog has a uniform drop size and
this is different to the drop size distribution of natural fog,
but could be considered an homogeneous fog. If convection
was 0.5 m s−1 and the mean concentration was 0.5 mg m−3
during three hours, then available water would be 0.54 mm
(m−3 ) or 0.12 mm (m−3 ) in the case of radiative fog.
There are few reports regarding fog storage capacity by the
vegetation (Jarvis, 2000), and the present work is the first to
determine the storage capacity for a bromeliad. The results
showed that drainage under natural conditions must be very
difficult because the measured storage capacity is too high
in comparison to natural fog fluxes reported for the region
(Martorell and Ezcurra, 2002). Using stable isotopes Ingram
and Matthews (1988) found that fog drip may be an important source of infiltration and groundwater recharge in an arid
climate.
Photosynthesis and transpiration of shrub species such as
P. laevigata are important during the morning, but under
drought and dark conditions stomata remain closed (Hultine et al., 2003). Although the leaf area of P. leavigata
and others is greater than that of T. recurvata, their interception storage capacity for fog, per unit of leaf area, must be
smaller because the leaves of A. farnesiana and P. leavigata
are glabrous and have a water-repellent waxy coat. In these
conditions the presence of T. recurvata and their fog interception capacity could increase the availability of water for
evaporation and decrease the vapor deficit of their host plants
for longer during the early day.
Tillandsia recurvata could reduce soil erosion at the start
of the rainy season, when most of the host shrubs are leafless
in the semiarid climates. On the other hand, there are some
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A. Guevara-Escobar et al.: Ball moss fog interception
reports of pine forest decay allegedly to T. recurvata infestation. For terrestrial species that depend upon fog interception a water stress situation could develop in the presence of
T. recurvata. While the causes of disordered growth of T. recurvata are multiple, availability or resources, such a s water,
is paramount.
Although these views appear to be contradictory, it is
likely that the habitat of T. recurvata is changing due to
global warming. In a catchment near the study area GómezDı́az et al. (2007) predicted increases of 57 to 62 % in
the arid climate and a reduction of 23 % in the temperate
area. Increased temperatures would allow T. recurvata to expand from lowland semiarid environments to higher altitudes
where temperate forests are dominant. In addition, higher
nocturnal temperatures due to climate change (IPCC, 2002),
would reduce the chance of advective fog formation and increase the preasure on the availabily of water resources.
4
Conclusions
The fog and rain data obtained under simulation showed the
potential impact of T. recurvata on water relations where occult precipitation occurs. The interception storage capacity
for rain was 0.19 mm which translates to 35.8 mm of annual
rainfall. On the other hand, the fog interception storage capacity of 0.56 mm was much higher. However, the fluxes of
natural fog probably are not enough to fill this capacity. Water detention after 1 h of wetting by fog was higher in darkness and therefore, stomata play an important role in water
uptake. Tillandsia recurvata depends on their hosts to intercept fog and thus conserving complete natural ecosystems is
important for water resource management. The benefits of
T. recurvata related to reduced water uptake by shrubs, grass
and herbs and the impact on soil conservation and aquifer
recharge still need to be investigated.
Acknowledgements. The authors would like to thank all their
colleagues and students at Universidad Autónoma de Querétaro
who contributed to this study.
Edited by: H. H. G. Savenije and S. White
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