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The Ecological Role of Biological Soil Crusts in The Rome Sand Plains of Central New York
The Ecological Role of Biological Soil Crusts in The Rome Sand Plains of Central New York
Carlos M. Rymer
Rebecca L. Schneider
Summer, 2006
Cornell Biological Field Station
Cornell University
Table of Contents
Abstract..................................................................................................................... 2
Introduction.............................................................................................................. 3
Results..................................................................................................................... 9
A. In Situ Substrate Moisture Content............................................................... 9
B. Biological Soil Crust Contribution to Water Absorption.............................. 9
C. Effects of Controlled Environmental Conditions on Crust Water
Absorption.................................................................................................... 10
D. Influence of Biological Soil Crust on Soil Microclimate............................. 11
E. Soil Stability................................................................................................. 11
F. Biological Soil Crust Recovery.................................................................... 11
Discussion............................................................................................................... 11
Conclusion.............................................................................................................. 13
Acknowledgements................................................................................................ 14
References.............................................................................................................. 21
1
Abstract
Biological soil crusts are communities of mosses, lichens, cyanobacteria, green algae, and
other organisms living at the soil surface of barren or desert habitats. It has been frequently noted
that biological soil crusts enhance soil moisture retention, increase nutrient availability, stabilize
soils, improve water infiltration, and increase plant nutrient uptake. The influences that
biological soil crusts have on soil physical properties, including moisture retention and soil
stability, in the Rome Sand Plains of Central New York was assessed. Eight to ten replicated
microcosm samples, each 5cm in diameter and 10cm in depth, were collected from the Rome
Sand Plains for bare soil, lichen-covered crust, and moss-covered crust. Replicates were
subjected to four different treatments of temperature and relative humidity levels. Samples were
dried and weighed at regular intervals under each treatment to quantify moisture absorption or
loss. Differences in surface temperature between areas covered with biological soil crust and
those with bare sand were measured continuously using TidBitTM temperature data logging
devices buried approximately 8cm below the surface, which allowed the assessment the crust’s
influence on in situ soil physical conditions. Soil stability was measured in the biological soil
crust and the adjacent, barren sand dunes using a soil penetrometer. Spots disturbed by sampling
were treated with fertilizer, inoculums, a combination of these two treatments, or left untreated to
determine if recovery can be enhanced. Recovery was periodically assessed based on soil
penetrometer measurements.
Our results show that the biological soil crust is important in enhancing soil physical
properties. Water absorption and evaporation were greatest in samples containing lichens only.
Under hyper-arid conditions, the biological soil crust lost its ability to absorb and retain water.
Sand moisture content was greatest in sand beneath the biological soil crust. Biological soil crust
water contribution ranged between 10-20% of total absorbed water. Soil covered with biological
soil crust had significantly greater stability than bare soil. The biological soil crust in the Rome
Sand Plains improves soil physical conditions through water absorption and contribution, soil
stabilization, and soil water retention. These results show that biological soil crusts, even in a
moist, temperate region, are playing a key role in enhancing soil properties.
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Introduction
The recent increase in desertification, particularly when caused by human land
disturbance, has led to a search for methods of restoring the ecosystems that were formerly
disturbed. To this end, ecologists have focused on the chemical and physical needs of vascular
plants and other primary producers, which are essential in restoring a disturbed ecosystem.
Recently, communities of small, photosynthetic, and nutrient-fixing organisms called biological
soil crusts (BSCs) have gained attention as potential components of effective restoration efforts.
Biological soil crusts are found in dry, barren soils in most regions of the world, and are
composed in varying proportions of mosses, fungi, lichens, green algae, and cyanobacteria; they
are commonly known as the primary components of ecological succession. As a whole, these
communities perform extremely important functions, including nitrogen and carbon fixation,
improved water infiltration, erosion control, temperature regulation, and surface moisture
contribution (Belnap et al., 2001). Most of the scientific literature on these communities
describes the physiology of the organisms within the crust. Few studies have actually looked at
how biological soil crusts capture and retain moisture from the atmosphere and how they
influence subsurface processes.
Most studies of biological soil crusts have focused on desert ecosystems, where they can
be widely found. Yet these communities are actually distributed throughout the temperate region
of North America, including Ohio (Veluci et al. 2006), Michigan (Thiet et al. 2005) and
Massachusetts (Smith et al. 2004), although they are only poorly studied. Due to the temperate
climate of the eastern U.S., large vascular plants have occupied much of the territory; but
throughout this region, scattered “barrens” containing stable biological soil crust populations still
exist today. One such place is the Rome Sand Plains of central New York (Figure 1). This unique
ecosystem, a remnant of the eastern, sandy shores of Lake Iroquois, which receded about 10,000
years ago, is composed of mixed hardwood forests, swamps, peat bogs, sand dunes, and
tributaries (DEC, 2003). With its unique physical features, this pine barren is one of twenty
inland Pine Barrens found worldwide. Given an average annual precipitation of ~100 cm, it is
worth examining the role that biological soil crusts play in this environment.
The objective of this study is to investigate the influence the biological soil crust has on
soil physical conditions in the Rome Sand Plains. We measured the effectiveness with which
different crust microhabitats absorb and retain water and determined how the biological soil crust
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influences soil moisture, microclimate, and stability in a temperate region. We studied these
different functions in the field and the laboratory, where we subjected different crust microcosms
to various environmental conditions to observe temporal water absorption trends.
Methods
Study Site
The study was conducted within the Rome Sand Plains, which is located in Oneida
County, New York (Figure 1). All samples were collected within two patches of biological soil
crust, each approximately 150m², surrounded by a mixed deciduous-coniferous forest and
bisected by a ~1.0m-wide foot trail. Additional measurements were taken in a nearby sand dune.
In each of these locations, the soil type was mainly from the Windsor Soil Series, which consists
of deep, well-drained soils (USDA, 2006). Within the study site, five distinct categories were
noted. These were as follows:
• Moss-dominated crusts (Ceratodon purpureus, with occasional Cladonia cristatella);
• Lichen-dominated crusts (Genus Cladonia, including C. arbuscula and C. rangiferina);
• Mixed crusts;
• Sand dunes (loose sand devoid of crusts or macrophytes); and
• The trail (compacted soil devoid of crusts or macrophytes).
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hours. After that period, the dishes were allowed to cool in a dessicator for 30 minutes. Final
weights were recorded and the percent moisture was obtained using the following formula:
% moisture = [(initial weight – final weight)/initial weight] * 100%
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from the soil surface using a spatula. Immediately thereafter, both the soil and the organisms
were weighed separately. Each segment was then oven-dried at 45°C ± 2.0°C for five hours.
Water loss by each segment determined where the absorbed water was located.
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compact fluorescent light tubes were used, which were less intense (~200 Lumens) than the
previous light bulb used. A heater was kept on during the day to maintain the high temperature,
and the humidifier was kept on to maintain the high humidity. The heater was turned off at night,
when the temperature decreased an average of 3.0°C. An initial weight was measured, and
successive weights were measured every 5 to 6 hours, except during overnight hours, which
lasted between 10 to 12 hours. This experiment lasted 213 hours.
Experiment 4: The same absorption experiment was repeated using the same sand and lichen-
dominated crust samples, but with 10 different moss-dominated crust samples. They were also
oven-dried at an average temperature of 33°C ± 2.0°C to eliminate all the moisture contained in
the sand. They were then placed in the chamber with an average temperature of 21°C ± 1.0°C
and relative humidity of 90% ± 4%. The cooler was used to decrease the temperature, as in the
first experiment. As in the previous experiment, the humidifier was left on for the duration of the
experiment. An initial weight was measured, and successive weights were measured every 5 to 6
hours, except during overnight hours, which lasted between 10 to 12 hours. This experiment
lasted 183 hours.
Experiment 5: This experiment was performed under natural conditions. In this experiment, 5
lichen-dominated crust samples, 5 moss-dominated crust samples, and 5 sand samples were used.
It was initiated with soil moisture levels at field capacity. Initial weights were measured, and
successive weights were measured every 3 hours, except during overnight hours, which lasted 10
to 12 hours. The temperature and humidity was measured at each 3-hour interval using the multi-
function meter. The experiment lasted 252 hours.
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A total of 6 TidBitTM devices were used. Each device was placed into a PVC pipe 25cm
long and 3.5cm internal diameter. A string was attached to each device, which was pulled outside
of the tube before the bottom opening was partly closed with a plastic canvas (mesh size 14),
allowing air to circulate into the tube. Plastic bags were tightly placed into the opposite opening
to minimize air transfer between that in the tube and the atmosphere. Duct tape was then used to
tightly close the opposite opening. Each pipe was buried approximately 9cm into the sand at an
angle of approximately 30 degrees with the surface (Figure 3). In each patch, one pipe was
buried underneath sand covered with biological soil crust, while another was buried under
exposed sand. In the exposed sand treatments, an area of 0.25m² of biological soil crust was
removed. The remaining two pipes were each attached to a tree within each patch, approximately
50cm above the ground, to measure the surrounding air temperature in the immediate area. The
temperature, humidity, and precipitation for Rome, New York were obtained from the National
Weather Service for comparison. The pipes were collected 30 days later, and the data from the
TidBitTM devices were transferred onto a computer for analysis.
Soil Stability
The influence of the biological soil crust on soil stability was measured using a dial
pocket soil penetrometer (H-4205), which measures the pressure required to push the soil exactly
0.6cm in. The locations used to compare soil stability were patches of biological soil crust cover,
the compacted sand trail, and the southern sand dune. The measurements were taken on July 6
and 10, 2006, 3 days after a 0.03cm rain event and 4 days after a 0.03cm rain event, respectively
(additional date: we did it twice because we also wanted to take measurements at the sand dune).
The 10mm diameter plunger was used to maximize the sensitivity of the readings. Within each
habitat type, ten random spots were selected. In each spot, the soil penetrometer was pushed into
the sand approximately 0.6cm, and the resulting reading recorded in kg/cm². Due to greater soil
instability in the sand dune, a 20mm plunger was used. The different plunger required conversion
of the measurement to account for the increase in surface area.
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35 holes were created as samples were extracted. These holes were refilled with sand from the
trail and used as the disturbed spaces for this experiment. Three treatments were applied
randomly to three-fourths (n=27) of the recovery holes. Treatment 1 consisted of an inoculum
made from crushed lichens and pieces of mosses; treatment 2 was a combination of inoculum
and a 50mL-sample of fertilizer (Concentration?), which contained zinc, manganese,
phosphorous, and low amounts of nitrogen; and treatment 3 was a 50-mL sample of the same
fertilizer. The remaining one-quarter (n=9) of the recovery holes were not treated, but left
exposed as controls. Flags were labeled with sample codes and placed in the center of each
recovery hole for identification.
Recovery of the biological soil crust was measured using the soil penetrometer and
observation on June 29, July 6, and August 8. Exactly two soil penetrometer readings were taken
during the course of this experiment, and descriptions were recorded periodically. Soil
penetrometer measurements were taken at the center, edge, and outside of each hole. A system
for descriptions was devised using letters for 1) moss growth; 2) lichen elongation; and 3) no
recovery. After the experiment, the recovery holes remained undisturbed for full recovery.
Results
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transfer to the underlying soil, with greater amounts retained and transferred by lichens than by
mosses (Figure 12).
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Influence of Biological Soil Crust on Soil Microclimate
The results from this field experiment show that the biological soil crust buffers soil
temperature (Figure 10). The average daily temperature underneath the biological soil crust was
significantly lower than that underneath bare sand (No Cover Avg: 26.35 ± 4.70ºC; Cover Avg:
25.85 ± 3.73ºC; Paired t-Test: n=289; p=0; df=288).
Soil Stability
Average soil penetration values in areas covered with biological soil crusts was
significantly greater than those measured in the sand dune (ANOVA: n=11; p>0.05; df=2). The
soil penetrometer measurements showed that soil stability in the trail was not statistically
different than that of the biological soil crust, but it was significantly higher than that of the sand
dune (Figure 11).
Discussion
The results of this study clearly showed that biological soil crusts are playing a strong
role in controlling environmental conditions of underlying soil, including properties of moisture
content, temperature regime, and soil stability. These functions have been well documented in
biological soil crusts of desert or dry ecosystems around the world, including the southeastern
United States (Hawkes and Flechtner, 2002). Our study represents one of few studies conducted
in moist temperate regions of northeastern U.S. In this region, where average rainfall is plentiful,
~100 cm annually, the full presence and functions of biological soil crusts need to be examined.
A synthesis of limited literature suggests their occurrence in the eastern, temperate U.S. is
consistently associated with well-drained, sandy soils found in glacial deposits of Pine Barrens,
such as in Ohio (Veluci et al. 2006), Michigan (Thiet et al. 2005), New York, and the coastal
sand dunes of Massachusetts (Smith et al. 2004).
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Biological soil crusts influence the water-holding capacity of soils. In this study, substrate
moisture content was related to biological crust structure. As expected, soil without cover by any
macroscopic species had the least moisture content. This may be due to low soil stability and
organic matter; greater sand stability increases surface area in sand particles, while organic
matter increases the soil’s water-holding capacity.
Biological soil crusts captured more atmospheric moisture and maintained higher levels
of moisture content than bare soil, both in situ at the Rome Sand Plains and in experimental
microcosms left exposed to natural fluctuations in summer environmental conditions. However,
the amount of moisture captured and retained was significantly related to the type of crust cover,
specifically where a few species of moss or lichen dominated. When started with dry tissues,
lichen-dominated microcosms consistently captured and retained more moisture than moss-
dominated microcosms.
It is important to note that the macroscopic species of the biological soil crust have the
ability to rapidly absorb water during the first few hours of availability, just as in other
ecosystems (Tirkey and Adhikary, 2005). Under controlled summer relative humidity and
temperature levels, lichens took up moisture, on average, more rapidly (~40g/hr m²) than moss
(~20g/hr m²), which rate was greater than that of bare soil (~10g/hr m²). At equilibrium, lichen
absorbed the highest amount of moisture at the highest rate (lichen ~2.5-10 g/hr m²; moss ~4.0-
5.0 g/hr m²; sand ~4.0g/hr m²).
Only a few studies report rates of uptake or equilibrium water contents, but our rates
appear to be equal or greater than those reported. One explanation may be greater biomass of
biological soil crust at our site compared to elsewhere. At this study site, biological soil crusts
accounted for 1.02±0.26 kg (oven-dried)/m² of biomass. Other studies do not typically report the
biomass of the crust community, limiting opportunities for comparison. We did not evaluate
individual physiological mechanisms, but other studies indicate that morphology, biochemical
composition, and moisture-capturing ability (Belnap et al. 2001) are important characteristics of
biological soil crusts. These characteristics are usually determined by water limitation, which is
important in determining crust cover and the potential for controlling desertification (Bowker et
al., 2006).
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Our field capacity experiments, which initially saturated the microcosms and then
allowed them to dry out under controlled environmental conditions, showed no significant
difference in the rate of moisture loss between the crust and bare soil.
One ramification of moisture retention by the biological soil crust may be a buffering of
underlying soil temperature. In our experiments, their presence moderated underlying soil
temperatures, reducing high extremes and raising minimum extremes, as experienced during
summer months. This can be explained, in part, due to higher moisture content in the soil due to
crust cover, but other factors not examined, such as albedo, may also be contributing to the
temperature buffer. Biological soil crusts were also found to significantly increase soil stability
as compared with the sand dune; the trail’s stability was not significantly different likely because
of compaction due to footsteps. Clearly, this crust function reduces soil erosion, explaining the
difference in relief between the biological soil crust and the sand dune, which was slightly rolling
and prone to wind erosion (Lange et al., 1994).
Overall, the influences of biological soil crusts on the temperate biome’s soil are creating
suitable habitat for larger plants and may encourage further succession. If this is so, then the
extent of dry periods may be critical in determining successful establishment of vascular plants
and how biological soil crusts may contribute to such establishment. This has significant
implications for vascular plant establishment in disturbed lands, which require improved soil
conditions (Hawkes and Flechtner, 2002).
Conclusion
This study shows that biological soil crusts have a major influence on soil physical
properties even in moist, temperate regions. In the Rome Sand Plains, the biological soil crust is
improving soil physical conditions through the following functions:
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Acknowledgements
I would like to thank Dr. Ed Mills and the entire staff at the Cornell Biological Field
Station, particularly Brian Young and Bill Thelen, for its help in this study. I greatly appreciate
the efforts of my advisor, Rebecca Schneider, in aiding me throughout the study and providing
needed materials and important suggestions. Her help greatly influenced the outcome of this
study. I’m grateful to the New York State Department of Conservation for permission to use two
patches of biological soil crust in the Rome Sand Plains, and to Professor Pfitsch from Hamilton
College for allowing me to work in his research area.
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Reindeer
Moss
Ladder
Lichen
British
Soldier
Burned Ground
Moss 3cm
Cyanobacteria
Table 1. Macroscopic Species of the Biological Soil Crust in the Rome Sand Plains.
Common Name Scientific Name
Common Species
Burned Ground Moss Ceratodon purpureus
Reindeer Moss (lichen) Cladonia rangiferina
Ladder Lichen Cladonia verticillata
British Soldiers Cladonia cristatella
Uncommon Species
Reindeer Lichen Cladonia arbuscula
Powdered Trumpet (lichen) Cladonia fimbriata
Yellow Moss Homalothecium fulgescens
Haircup Moss Polytrichum strictum
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Figure 4. Soil Moisture Content.
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Figure 6. Water Loss by Biological Soil Crust at High Temperature.
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Figure 8. Water Absorption.
18
Rain Period
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Figure 12. Biological Soil Crust Moisture Contribution.
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