-
Main factors determining bioerosion patterns on rocky cliffs in
a drownedvalley estuary in the colombian pacific (eastern tropical
pacific)
Alba Marina Cobo-Viveros, Jaime Ricardo Cantera-Kintz
PII: S0169-555X(15)00329-3DOI: doi:
10.1016/j.geomorph.2015.05.036Reference: GEOMOR 5250
To appear in: Geomorphology
Received date: 6 August 2014Revised date: 8 May 2015Accepted
date: 10 May 2015
Please cite this article as: Cobo-Viveros, Alba Marina,
Cantera-Kintz, Jaime Ricardo,Main factors determining bioerosion
patterns on rocky clis in a drowned valley es-tuary in the
colombian pacic (eastern tropical pacic), Geomorphology (2015),
doi:10.1016/j.geomorph.2015.05.036
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MAIN FACTORS DETERMINING BIOEROSION PATTERNS ON ROCKY
CLIFFS IN A DROWNED VALLEY ESTUARY IN THE COLOMBIAN
PACIFIC (EASTERN TROPICAL PACIFIC)
Alba Marina Cobo-Viverosa and Jaime Ricardo Cantera-Kintz
a,b
a. Research Group in Estuaries and Mangroves - Ecomanglares.
Department of Biology.
Faculty of Natural and Exact Sciences. Universidad del Valle.
Calle 13 #100-00. Cali,
Colombia. AA. 25360. Phone number: (+57) 2 3212100 ext.
2824.
[email protected]
b. Titular Professor. Department of Biology. Faculty of Natural
and Exact Sciences.
Universidad del Valle. Calle 13 #100-00. Cali, Colombia. AA.
25360.
[email protected]
Corresponding author: Alba Marina Cobo Viveros1
1 Present postal address: Instituto de Investigacins Marias. Ra
Eduardo Cabello 6. 36208 Vigo
(Pontevedra). Spain. [email protected] Present cell phone:
(+34) 622 078 341.
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HIGHLIGHTS
We measured perforation volumes on two rocky cliffs in the
Colombian Pacific
coast.
Interior cliffs (not exposed to wave action) presented more
porosity and
perforation volumes.
Exterior cliffs (exposed to wave action) had higher bioeroder
diversity and
abundance.
Lower tidal zones showed higher abundance of bioeroders than the
other zones.
Boring bivalves were less abundant compared to boring
crustaceans.
ABSTRACT
Bioerosion is an important process that destroys coastal rocks
in the tropics. However,
the rates at which this process occurs, the organisms involved,
and the dynamics of
rocky cliffs in tropical latitudes have been less studied than
in temperate and subtropical
latitudes. To contribute to the knowledge of the bioerosion
process in rocky cliffs on the
Pacific coast of Colombia (Eastern Tropical Pacific) we
compared: 1) boring volume, 2)
grain size distribution of the rocks, and 3) rock porosity,
across three tidal zones of two
cliffs with different wave exposure; these factors were related
to the bioeroding
community found. We observed that cliffs that were not exposed
to wave action (IC,
internal cliffs) exhibited high percentages of clays in their
grain size composition, and a
greater porosity (47.62%) and perforation (15.86%) than exposed
cliffs (EC). However,
IC also exhibited less diversity and abundance of bioeroding
species (22 species and
314 individuals, respectively) compared to the values found in
EC (41.11%, 14.34%, 32
and 491, respectively). The most abundant bioeroders were
Petrolisthes zacae in IC and
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Pachygrapsus transversus in EC. Our findings show that the tidal
zone is the common
factor controlling bioerosion on both cliffs; in addition to the
abundance of bioeroders
on IC and the number of bioeroding species on EC. The
integration of geology,
sedimentology, and biology allows us to obtain a more
comprehensive view of the
patterns and trends in the process of bioerosion.
KEYWORDS
Bioerosion, Grain size distribution of rocks, Boring volume,
Rock porosity, Bioeroding
fauna, Rocky cliffs, Eastern Tropical Pacific.
1. INTRODUCTION
Bioerosion is an important process that destroys coastal rocks
in the tropics (Trenhaile,
1987); it occurs through the biological breakdown and removal of
hard substrates by
surface abrasion and boring. During surface abrasion, endolithic
and grazing organisms
(e.g. molluscs, echinoderms, fish, and some crustaceans) rasp,
bite and scrape away a
thin layer of rock (Trudgill, 1985; Trenhaile, 1987), produce
particulate detritus
(Torunski, 1979), and obtain nutrition from endolithic algae.
During boring, perforating
organisms (e.g. endolithic bacteria, algae, fungi, and lichens;
sponges, sipunculans,
polychaetes, bivalves, crustaceans, and echinoderms) directly
remove rock material and
weaken the remaining rock, making it more vulnerable to
mechanical wave erosion and
weathering (Trenhaile, 2005). As a consequence of these two
processes, rocks collapse
and decompose (Hutchings, 1986; Ricaurte et al., 1995; Cantera
et al., 1998), generating
new substrates, changing cliff structure, and enriching the
surrounding ecosystems with
sediments and rocks from the fallen material, thus modifying the
biological community.
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Not all marine organisms destroy the underlying rock; some can
also protect it from
incoming waves and physicochemical attack by forming organic
crusts in the lower
intertidal and upper subtidal zones (rhodophytes: Lithothanium,
Lithophyllum;
chlorophytes: Halimeda; sublittoral brown algae; barnacles:
Chthamalus, Balanus;
limpets: Patella, Lottia, Fissurella, Siphonaria, Crepidula)
(Trenhaile, 1987). However,
determining the scale used to identify the role played by
organisms involved in
bioerosion can be difficult (Forns et al., 2006): sometimes it
is clear that an individual
acts as a bioeroder or as an occupant nestler of a previously
existing hole (while
modifying it), but this is not always easy to ascertain. For
example: encrusting
organisms that have an important role in protecting the rock
surface from physical
erosion (Focke, 1977) sometimes also act as bioeroders; they can
take away some rock
when removed from the cliff (e.g. barnacles), or they can weaken
what they are
supposedly protecting by chemical or other processes (e.g. micro
and macroalgae)
(Naylor and Viles, 2002).
Cliffs are also destroyed by mechanical and chemical means
(McLean, 1974). Wave
erosion is considered the dominant mechanical erosional agent in
many parts of the
world (Trenhaile, 1987); it occurs through steady wave action,
generation of high shock
pressures (Trenhaile and Kanyaya, 2007; Bezerra et al., 2011),
or the abrasion from
sweeping, rolling, or dragging of rocks and sand (Trenhaile,
1987). Chemical
weathering is the result of a series of chemical reactions
(Trenhaile, 1987) that modify
the rock carbonate chemical equilibrium when working together
(Trudgill, 1985). Some
conceptual models of erosion on rocky coasts highlight the
importance of the wave
force/rock resistance relationship and leave aside that of
biological agents (Sunamura,
1994) but the effects of chemical, mechanical, and biological
erosion can be synergistic
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(Hutchings, 1986). The reconceptualization of Naylor et al.
(2012) demonstrated that
most of the geomorphologic processes are affected by organisms
and included
biological agents as important reducers of the resisting force
of the rock.
There has been a growing interest in rock coast geomorphology in
temperate and sub-
tropical latitudes (Naylor et al., 2010), but bioerosion rates
and dynamics of rocky cliffs
in tropical latitudes have been less studied (Moses, 2013). Some
publications integrate
biological, geological and sedimentary variables (Fischer,
1981a, 1981b; Cantera et al.,
1998), and quantify cliff retreat (Ricaurte et al., 1995;
Cantera et al., 1998) and
bioerosion rates of several organisms (Rasmussen and
Frankenberg, 1990; Toro-Farmer
et al., 2004; Herrera-Escalante et al., 2005; Asgaard and
Bromley, 2008; Lozano-Corts
et al., 2011). Cantera et al. (1998) measured erosion rates and
studied the biodiversity,
zonation, and types of cavities made by perforating fauna in two
rocky cliffs in
Buenaventura bay (Pacific coast of Colombia). Additionally,
there are some works that
studied perforations by crustaceans and bivalves (Cantera and
Blanco-Libreros, 1995;
Ricaurte et al., 1995), and that quantified the erosion rate of
sea urchins in rocky cliffs
(Lozano-Corts et al., 2011). However, little work has been done
on bioerosion of rocky
cliffs in Colombia, in spite of the impact it can have on nearby
human settlements living
on top of the cliffs or near them. This process requires further
study (Correa and
Gonzalez, 2000).
The present study contributes to the knowledge of the grain size
distribution and boring
volumes of two rocky cliffs in a drowned valley estuary in the
Colombian Pacific
(Eastern Tropical Pacific). As intertidal bioerosion cannot be
understood without
biological processes (Trudgill, 1985), boring volumes are used
as a quantitative
bioerosion indicator relative to wave exposure, tidal zone, and
the bioeroding
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community found. Combining the effects of several bioerosion
variables can indicate
the areas that erode more quickly, compared to studying the
effects of each variable
separately.
2. MATERIALS AND METHODS
2.1. Study site
Buenaventura Bay is an ancient drowned valley estuary located in
the Pacific coast of
Colombia, between 348N 354N and 7705W 7720W (Fig. 1). It is
located in
one of the most humid places of the world: the average annual
precipitation rate is more
than 7000 mm/year, which makes chemical erosion very high.
Buenaventura Bay has a
tropical hot and humid rainforest climate: the mean annual
temperature is 26.2C, and
the mean relative humidity is 89%. The rainy season occurs
between August and
November. Waves can reach heights of 2 m outside the bay, but
they are rapidly
reduced to 0.9 m near the entrance due to energy dissipation
related to floor friction.
The bay undergoes forcing by semi-diurnal tides with a
meso-macrotidal range of 4 m
(Cantera and Blanco, 2001).
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Fig. 1. Geographical location of the cliffs studied. Right:
South America, showing the
location of the Pacific coast of Colombia (Middle). Left.
Buenaventura Bay, showing
the locations of IC (unexposed cliffs) and EC (exposed
cliffs).
The north side of Buenaventura Bay is characterized by vertical
to sub-vertical cliffs
that range from 10-20 m in height, cut into horizontal to
sub-horizontal Tertiary
sandstones, shales and mudstones (Correa and Morton, 2010). The
tops of the cliffs are
covered with dense vegetation, as occurs in most humid tropical
regions (Trenhaile,
1987). The cliffs located on this side of the bay are composed
by the Raposo and
Mayorqun geological formations (of sedimentary origin) from the
Superior and Median
Tertiary (Galvis and Mojica, 1993; Martnez, 1993); these
formations consist of shale,
mudstones, and dark gray siltstones organized in layers that
vary from a few centimeters
to 2 m thick. Coarse sediments (sandstone, slabs and clusters)
are also present,
randomly arranged between the strata (Cantera et al., 1998). Two
cliffs located on this
side of the bay were chosen (Fig 1): one on the external zone
(EC, located 0.5 km from
the entrance and exposed to wave action) and the other on the
internal zone (IC, located
15.4 km from the entrance of the bay and not exposed to
substantial wave action).
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Sedimentary deposits of continental origin reach Buenaventura
Bay through the rivers
flowing into it; these sediments particularly affect cliffs in
IC so that the rocks forming
these cliffs are softer than the ones forming EC.
2.2. Sampling.
Nine blocks of approximately 15 cm x 15 cm x 15 cm were
extracted from each cliff,
using a chisel: three from the supralittoral, three from the
upper intertidal, and three
from the lower intertidal. These three tidal zones were
distinguished based on the
characteristic algae, perforations, and tidal coverage: the high
zone (Supralittoral or
splash zone) is covered by patches of the algae Cladophora
albida, Cladophora
herpestica (or a mix of both), and Bostrychia tenella; it is
poorly bored and is only
covered by the tide during spring tides, but other than that it
only receives the splash
from waves that break in the inferior tidal levels. The upper
intertidal zone (or Superior
Mesolittoral) is covered by Bostrychia radicans, with patches of
Cladophoropsis sp.
and Boodleopsis verticillata; it is slightly perforated and it
stays submerged for a longer
period of time than the high zone. The lower intertidal zone (or
Inferior Mesolittoral)
can present coverage by B. radicans; it is the most perforated
zone, it stays submerged
for longer periods of time than the other two zones, and it is
sometimes separated from
the upper intertidal zone in this locality by a stratum of
volcanic, hard rock covered by
oysters and barnacles.
After the blocks were extracted from the cliffs, they were
submerged in a mixture of
water, alcohol, and clove oil in order to collect all the
benthic fauna within the rock
(which was preserved in 70% alcohol); partial desalinization
occurred in this process.
The fauna collected inside the blocks were identified using
taxonomic keys for each
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group (Haig, 1960; Olsson, 1961; Keen, 1971; Fauchald, 1977;
Brusca, 1980; Brusca
and Iverson, 1985; Froidefond, 1985; Williams, 1986; Kim and
Abele, 1988; Abele and
Kim, 1989; Ros and Ramos, 1990; Poore, 1994; Hilbig, 1997;
Cantera et al., 1998).
The organisms were marked and preserved in 80% alcohol.
The blocks of rock were cored into smaller blocks of
approximately 10 cm x 10 cm x 10
cm. To determine the perforation volume, the rocks were first
saturated in water
(submersion time depended on the characteristics of each block),
taken out, and the
liquid remaining inside the perforations was extracted.
Afterwards, the rocks were
weighed in air (with an electronic balance of precision 0.1 g)
and in water (Annex B),
calculating their volume from the weight difference in both
environments. The volume
of rock (VR), including perforation volume (due to bioerosion)
and porosity volume (due
to the incipient rock porosity), was found using equation 1:
(1)
where WR is the weight of rock + rock pores in the air, WsR is
the weight of rock + rock
pores in the water, and water is the water density.
The volume of rock without perforations (VT) was found by
filling the boreholes with
modeling clay, sealing the rock with paraffin wax, weighing the
block in air and water,
and applying equation 2.
(2)
WRC is the weight of the sealed rock in air, and WsRC is the
weight of the sealed rock in
water.
The volume of rock due to perforations (VP) was calculated from
the difference between
VT and VR (equation 3).
(3)
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2.3. Specific gravity
The specific gravity (Gs) of any substance is the unitary weight
of the material divided
by the unitary weight of distilled water at 4C. This variable is
used in the hydrometer
analysis (Bowles, 1981a) to obtain the void ratio of a soil or
ground, and to predict its
unitary weight (equation 4). The methodology is described in
Bowles (1981a).
(4)
Wsol is the weight of the solids, Wfw is the weight of the flask
+ water, and Wfws is the
weight of the flask + water + solids.
2.4. Hydrometer analysis
This analysis estimates the particle size distribution of soils
that contain a considerable
amount of particles between 0.075 and 0.001 mm (clay and silt).
The Stokes Law
(equation 5) estimates the falling rate of spheres in a fluid
(v, in cm/s) from the specific
weight of the spheres ( , in
g/cm3), the specific weight of the fluid ( , usually water), the
absolute viscosity or fluid
dynamic (, in dynes x seg/cm2) and the sphere diameter (D, in
cm) (Bowles, 1981a;
Das, 2001).
(5)
This equation is valid for particle diameters between 0.0002 and
0.2 mm. Temperature
was taken into account, since the specific weight and viscosity
of water depend on this
variable.
The hydrometer analysis was performed on the rocks from the
cliffs and on a control
solution prepared with 125 ml of 4% sodium hexametaphosphate
(NaPO3, a dispersing
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agent which neutralizes the charges on the smallest grain sizes
that often have negative
charge) and sufficient distilled water to produce 1000 ml. The
hydrometer was put into
the control cylinder to record zero and meniscus corrections;
the temperature was
measured as well.
To prepare the sample for the hydrometer analysis, the blocks of
rock were crumbled
and dried at 110C for 24 h, after which they were macerated and
passed through the
#200 sieve to obtain a 50 g sample. 125 ml of 4% NaPO3 were
added to the sample and
the resulting mixture was left standing for 24 h. Afterwards the
mix was transferred to a
dispersion (or malt mixer) cup and water was added until the cup
was about two-thirds
full. The contents were mixed for a minute, and then the mixture
was carefully
transferred to a 1000 ml sedimentation cylinder. Any soil left
in the dispersion cup was
rinsed using a plastic squeeze bottle and the remains were
poured into the sedimentation
cylinder. Next, water was added until the 1000 ml level and the
mixture was agitated
again for 1 min to homogenize the material within the column;
this was done by placing
the palm of the hand over the open end and turning the cylinder
upside down and back.
Finally, the sedimentation cylinder was set on a table and a
ASTM 152H hydrometer
was inserted; readings were taken at the time intervals t = 0.5
min, 1 min, 1.5 min, 2
min, 2.5 min, 3 min, 3.5 min, 4 min, 8 min, 16 min, 30 min, 60
min, 120 min, and 240
min, or until the reading became constant. Readings were always
taken at the upper
level of the meniscus because suspended soil water solution
makes the system opaque.
Temperature was also recorded at each time interval.
After all the readings were taken, a series of corrections were
performed due to zero,
meniscus and temperature (Bowles, 1981b). The zero correction
(Cz) is applied to the
actual hydrometer reading depending on the hydrometers zero
reading in the control
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cylinder: if this reading is below the water meniscus, Cz will
be positive; if it is above, it
will be negative; and if the reading is at the meniscus, Cz will
be zero (Bowles, 1981a).
The meniscus correction (Cm) is the difference between the upper
level of meniscus and
water level of the control cylinder (Bowles, 1981a). The
temperature correction (CT) is
done when the temperature of the soil suspension is not 20C
(hydrometers are
generally calibrated at this temperature); if it is above, the
hydrometer reading will be
less and CT will be positive (and vice versa) (Bowles, 1981a).
CT was determined from
Table A1 in the Annex.
The corrected hydrometer reading (Rc) was calculated from the
actual reading (Rreal) as
follows:
(6)
The percent finer (P, percentage of particles that go through
the sieve) was calculated as
follows
(7)
where a is a correction factor used whenever the Gs of soils is
different from 2.65 (the
Gs at which the 152H hydrometer was calibrated). It was
determined from Table A2 in
the Annex using Gs. Ws is the weight of the soil sample (in
grams).
The equivalent particle diameter (D, in mm) was calculated using
the following
formula:
(8)
where the factor K is a function of temperature, Gs and water
viscosity; for the known
Gs of the soil, K was obtained from Table A3 in the Annex. L is
the effective
hydrometer depth L (in cm) obtained from Table A4 in the Annex
for the meniscus
corrected reading; and t is the time interval (in minutes).
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Grain size distribution curves (D) were plotted versus the
percent finer (P) on a semi-
logarithmic plot.
2.5. Statistical analysis
The percentage of perforation and porosity volumes were
determined and compared
between tidal zones of each cliff with a one-way ANOVA, and
between both cliffs with
a two-way ANOVA. Homogeneity of variances was tested for using
Levenes test.
Bioeroder abundance and number of bioeroding species between
tidal zones was
compared with a one-way ANOVA, and between cliffs with a two-way
ANOVA,
because normal distribution and homogeneity of variances are not
critical to perform an
ANOVA when sample sizes are equal (Hammer and Harper, 2008). If
the ANOVA
showed significant inequality of means, the post-hoc
Tukey-Kramer pairwise
comparison was used (Hammer et al., 2001).
The percentage of biodegraded volume was related to the
abundance and richness of
eroding fauna, tidal zone, and percentage of natural porosity of
the rock using a simple
correlation analysis (Zar, 2010). The correlation coefficients
were compared with a one-
way ANOVA.
Finally, to determine if the combined effect of all factors on
the bioerosion process was
higher than the effect of each factor taken separately, a
multiple regression analysis of
perforated volume, richness and abundance of bioeroding fauna,
tidal zone, and volume
of natural porosity of the rock was performed.
3. RESULTS
3.1 Composition of cliff sediments
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Both cliffs are composed mainly of shale. In the hydrometer
analysis, both cliffs
showed a high percentage of fine particles in their grain size
distribution, but unexposed
cliffs (IC) showed finer particles than exposed cliffs (EC)
(Table 1, Fig. 2A). In fact, we
found particles smaller than 2m (clays) in IC but not in EC
(Table 1, Fig. 2B).
Table 1. Grain size composition (percentage) of unexposed (IC)
and exposed cliffs
(EC) in the Pacific coast of Colombia.
Particle Diameter IC EC
Fine sands < 75m 100% 100%
< 50m 80-92% 62-86%
Silts < 20m 40-51% 28-45%
< 10m 11-30% 8-15%
Clays < 2m 9-11% 0%
One curve from the low (L) and another from the middle (M) tidal
zone in EC (blocks
L1 and M1, Fig. 2B) differed from the rest of the cliffs grain
size distribution. They
represent an inclusion of hard rock that occurs along these
kinds of cliffs, with a thicker
composition than the rest of the analyzed samples: only 40% of
the particles exhibited
diameters
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Fig. 2. Grain size distribution of A. Unexposed (IC) and B.
Exposed cliffs (EC) on the
Pacific coast of Colombia. The legend on the right represents
each tidal zone: low (L),
middle (M), and high (H).
3.2. Perforation and Porosity volumes
More perforation volume (produced by bioerosion of the rock) was
found in the low and
high tidal zones of IC, but the middle tidal zone was more
densely perforated in EC
(Table 2). Significant differences were found in the perforation
volume between tidal
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zones in EC (p = 0.001) due to dissimilarities between the high
and low (p = 0.004) and
the high and middle tidal zones (p = 0.002). Significant
differences were also found
when we compared the perforation volume of both cliffs (p =
0.002), between the low
tidal zone in IC and the high tidal zone in EC (p = 0.012), and
between the high and
middle tidal zones in EC (p = 0.017). Significant differences
were not found in
perforations between wave exposures (p = 0.666) or an
interaction between wave
exposure and tidal zones (p = 0.126).
Significant differences in the porosity volume (incipient rock
porosity) were found
between wave exposures (p = 0.016) because IC showed more
porosity than EC in all
tidal zones; these differences were due to dissimilarities
between the high tidal zone in
IC and the low tidal zone in EC (p = 0.039). There were no
significant differences in
porosity between tidal zones (p = 0.146), nor an interaction
between wave exposure and
tidal zone (p = 0.336).
Table 2. Percentage of perforations due to bioerosion (relative
to total volume) and
percentage of porosity (relative to volume of solids + volume of
pores) found for the
three tidal zones of unexposed cliffs (IC) and exposed cliffs
(EC) in the Pacific coast of
Colombia. Richness and abundance of bioeroding species of the
studied cliffs is also
shown.
Cliffs Perforations
(%)
Porosity
(%)
Abundance
(Individuals)
Richness
(Number
of species)
IC
Low 25.08 46.39 183 18
Middle 14.37 47.04 99 13
High 8.14 49.41 32 11
Average 15.86 47.62 Total 314 22
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EC
Low 21.02 34.15 295 25
Middle 23.93 41.20 123 14
High 0.30 45.65 73 4
Average 15.08 40.33 Total 491 32
3.3. Bioeroding fauna
We found 314 individuals belonging to 22 macrobioeroding species
in IC (Table 2).
More bioeroder species and abundance of grazers and borers were
found in the low tidal
zone (58.3%). Petrolisthes zacae was the most abundant species
in this cliff (41.1%;
Fig. 3A) because it appeared in great numbers in the low and
middle tidal zones. The
amphipod Chelorchestia sp. was the most abundant in the high
tidal zone (relative
abundance of 37.5%). Significant differences were found in the
abundance of
bioeroders between the high and low tidal zones (p = 0.004) but
not in the number of
bioeroding species (p = 0.086).
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Fig. 3. Abundance of bioeroding species in the low, middle and
high tidal zones of A.
Unexposed cliffs (IC) and B. Exposed cliffs (EC) in the Pacific
coast of Colombia. The
category Others groups bioeroding fauna with total abundances of
less than 10
individuals.
A total of 491 individuals belonging to 32 bioeroding species
were found in EC,
concentrated in the low zone (25 bioeroder species and 60% of
total cliff abundance).
Pachygrapsus transversus, Alpheus javieri and Upogebia
tenuipollex were the most
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abundant species in this cliff (relative abundances of 18.1%,
14.5%, and 12.4%,
respectively; Fig. 3B). However, Petrolisthes armatus, P.
transversus and Ligia
baudiniana were the most abundant for the low, middle, and high
tidal zones (48, 38,
and 59 individuals, respectively). Significant differences were
found in the number of
bioeroding species between the high and low tidal zones (p =
0.029), but not in the
abundance of bioeroders (p = 0.104).
When both cliffs were compared, significant differences were
found in the abundance of
bioeroders between the high zone in IC and the low zone in EC (p
= 0.018). We also
found significant differences in the number of species between
the low and high tidal
zones in EC (p = 0.011). However, there were no significant
differences in the
abundance or richness of bioeroders between wave exposures (p =
0.149 and p = 0.876,
respectively), nor in the interaction between wave exposure and
tidal zone (p = 0.621
and p = 282, respectively).
3.4. Relationship between cliff composition and bioeroding
fauna
The perforation volume in both cliffs was negatively correlated
to tidal zone (IC: r = -
0.686, p = 0.041, Fig. 4A; EC: r = -0.782, p = 0.022, Fig. 4C)
but positively correlated
with the abundance of bioeroders in IC (r = 0.74, p = 0.023,
Fig. 4B), and with the
richness of bioeroders in EC (r = 0.725 p = 0.042, Fig. 4D).
The combination of factors did not indicate a statistically
significant effect on the
percentage of perforations found for any of the cliffs. However,
the R2 values indicate
that 63.5% (83.9%) of the total variation of perforations in IC
(EC) is explained by the
regression (Table 3). R2 values are higher in EC than in IC,
indicating that all variables
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chosen for the analysis influence the percentage of perforations
found in EC, but not in
IC.
Fig. 4. p-value, correlation (r) and determination coefficients
(r2) found for significant
linear correlations in unexposed cliffs (IC; A-B) and exposed
cliffs (EC; C-D) on the
Pacific coast of Colombia. Biodegraded volume was negatively
correlated with tidal
zone in both cliffs (A and C), and positively correlated with
abundance of bioeroders in
IC (B) and with richness of bioeroders in EC (D).
Table 3. Multiple regression analysis comparing volumes of
perforation with tidal zone,
volume of natural porosity of the rocks, richness and abundance
of bioeroding fauna in
unexposed (IC) and exposed cliffs (EC). Multiple correlation
coefficient (R), multiple
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determination coefficient (R2), and adjusted coefficient of
determination are shown
(Ra2), as well as results of ANOVA for the multiple regression
data.
Statistics IC EC
Multiple R 0.797 0.916
Multiple R2 0.635 0.839
Adjusted Ra2 0.271 0.625
F F(4, 4, 0.95) = 1.746 F(4, 3, 0.95) = 3.922
p 0.301 0.145
4. DISCUSSION
4.1. Sediment composition of the cliffs
Our findings confirm the fine sediment composition of the rocks
forming the cliffs on
the Pacific Coast of Colombia, which is mainly due to
differences in particle
sedimentation rates during cliff formation, and to the energy of
the deposition
environment. The sedimentary rocks forming the cliffs are
composed of ancient mud
and silt and were produced by accumulation of sediments from the
river flow.
Unexposed cliffs (IC) were formed by sediments from rivers
flowing into Buenaventura
Bay, which is why they have clays only in the higher and middle
zones and few hard
substrata inclusions (as seen by the higher percentage of small
grain sizes). The
presence of coarse particles in the rocks between layers of fine
sediments in the low and
middle tidal zones of exposed cliffs (EC; blocks L1 and M1 in
Fig. 3B) is due to the
Raposo Mayorqun formation, characterized by the presence of
coarse sediments
(sandstone, slabs and clusters) randomly arranged between the
strata (Cantera et al.,
1998). These coarse sediments result from the consolidation of
sediments from the river
flow that subsequently changed the grain size composition of the
original Mayorqun
formation.
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It is important to recognize that different agents of bioerosion
may operate in distinct
zones across shore platforms (Naylor et al., 2012): for example,
biological weathering
and erosion-enhancing agents are typically found in
morphologically lower, moister
positions of shore platforms (our lower intertidal zones)
(Naylor et al., 2012), whereas
chemical/physical weathering agents are likely to be relatively
more important in drier,
morphologically high points where wetting/drying and
swelling/contraction are more
common (our supratidal zones) (Gmez-Pujol and Forns, 2009). Wave
action is
weaker in the tropics than in high latitudes, and younger
limestones or shales are
physically much weaker (Trenhaile, 1987). EC receives a more
constant and higher
wave action than IC, which can affect its porosity by removing
smaller particles from
the cliff in the first stages of erosion, causing abrasion on
the bigger particles that are
left. However, even though EC is more exposed to waves and we
expected this cliff to
be more perforated, it seems that porosity volumes play a more
important role in
determining perforation volumes. Trudgill (1985) established
that increased porosity
decreases rock resistance to erosion compared to that of
well-cemented rocks with few
joints; so the increased porosity in IC makes this cliff more
susceptible to erosion, in
spite of it being less exposed to waves.
In addition, although waves perform the erosive work, it is the
tidally modulated
distribution of wave energy that determines where this work is
performed (Trenhaile,
1978). The lower tidal zones of both cliffs stay submerged
longer than the other zones
as a result of the semi-diurnal tidal cycle of Buenaventura bay;
this benefits the
bioerosion community inhabiting the lower zones of both cliffs,
enhancing the higher
perforation volume exhibited by them.
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Bioerosion (in some cases) is greatest in sheltered sites
because wave shock and mobile
sedimentary particles driven by waves and currents can prevent
colonization of exposed
areas by some organisms (Trenhaile, 1987). Cliffs in IC are
exposed to higher river
discharges and a low wave impact; this allows a film to be
formed on the surface of
these cliffs, protecting them from rain and growth of
microbioeroders (e.g. algae).
Hutchings et al. (2005) found a similar effect on coral
bioerosion. In cliffs in IC, this
film also hinders grazer colonization and increases the presence
of bioeroding larvae in
all tidal zones in IC that otherwise would be eaten by grazers
feeding on the algae.
4.2. Bioeroding fauna
The difference in species richness between tidal heights of the
cliffs is exposed sites (as
EC) have better humidity and shelter conditions that allow
higher species richness in the
lower tidal zones, especially of species representative of the
lower tidal zones
(Petrolisthes armatus, A. javieri, U. tenuipollex). On the other
hand, species typical of
the high tidal zone (Chelorchestia sp., Petrolisthes zacae, P.
transversus) dominate
sheltered sites (as IC) (Palmer et al., 2003).
The boring habit is well developed in four pelecypod families:
Pholadidae and
Petricolidae (mainly mechanical borers) and Gastrochaenidae and
Mytilidae (largely
chemical borers that require a calcareous substrate) (Yonge,
1955; Trenhaile, 1987).
Drilling Mytilidae species have been reported as responsible for
the greatest amount of
perforations in hard rocks (Cantera et al., 1998), while species
of Petricolidae (Ansell,
1970) and Pholadidae have been for soft rocks (Warme and
Marshall, 1969; Pinn et al.,
2005, 2008). In this study, we found Cyrtopleura crucigera
(Pholadidae) in both cliffs,
while Sphenia fragilis (Myidae) and Pholadidea tubifera
(Pholadidea) were found in
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EC only. These findings differ from results previously found for
the area, where C.
crucigera, S, fragilis and Pholadidea sp.1 were found in IC
(Cantera et al., 1998). Three
species of bivalves were found in EC that were reported as
borers by Keen (1971):
Cryptomya californica (Myidae), Ensitellops hertleini
(Basterotiidae), and Barnea
subtruncata (Pholadidae). The mytilid species Brachidontes
playasensis was found in
low abundances in the low zone of cliffs in IC, where it could
be playing an important
role in protecting the rock surface from physical erosion.
Crustaceans, on the other hand, have been reported as borers of
wood (Davidson and de
Rivera, 2012), sandstones (Cade et al., 2001), and basalts
(Fischer, 1981b). Upogebia
tenuipollex and Alpheus javieri are boring decapods in cliffs of
the Colombian Pacific
coast (Ricaurte et al., 1995). They were found in great numbers
in EC but not in IC,
where they were outnumbered by Alpheus villus, Upogebia
spinigera, and Upogebia
burkenroadi. These three species may be taking an active part in
the bioerosion process
in cliffs that are less exposed to wave action
Several worms are also active borers in calcareous and
non-calcareous substrates
(Trenhaile, 1987). Hutchings and Peyrot-Clausade (2002)
recognize polychaetes and
sipunculans as dominant groups of macro-boring organisms in
newly available dead
coral substrate, facilitating the subsequent recruitment of
other boring organisms such
as sponges and bivalves. For the Pacific coast of Colombia,
Cantera et al. (1998) found
Polydora sp. in cliffs in EC. However, although we found three
species of polychaetes
inhabiting the rock burrows in EC (Nereis sp., Lysidice sp. and
Syllis sp.) and two in IC
Nereis sp. and Neanthes sp.), we did not find Polydora within
our samples. Of the
genera found, Lysidice has been found to be burrowing inside
Porites colonies
(Hutchings, 2008), so they could also be boring into rocky
cliffs. We only found one
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individual of the sipunculid genus Phascolosoma in a rock burrow
in the lower tidal
zone of EC (data not shown), contrary to previous findings that
highlighted their
importance on the bioerosion process in cliffs of the Pacific
coast of Colombia (Cantera
et al., 1998). Our evidence indicates that molluscs and
crustaceans are more important
than polychaetes and sipunculans in the bioerosion process in
the cliffs that were
studied.
4.3. Contribution of bioeroding fauna to perforation volumes
Previous authors found that a greater biological contribution to
erosion rates occurs on
sheltered shores compared to exposed ones (Trudgill, 1976;
Spencer and Viles, 2002;
Moura et al., 2012). This coincides with our findings, in which
IC presented better
conditions for the establishment of bioeroder communities, which
in turn was reflected
in a higher volume of perforation found compared to EC.
Naylor et al. (2012) highlighted the importance of the
biological role in the removal of
rocky masses and the erosion of rocky coasts (which had been
previously neglected).
The direct erosional role of grazing organisms is of particular
significance on tropical
and warm temperate limestone coasts, where wave attack may be
fairly weak
(Trenhaile, 1987). Grazing organisms always contribute directly
to the erosion of rock
surfaces, and their presence in IC can explain the higher
porosity found here, despite the
high percentage of small particle composition; macro- and
micro-grazers facilitate the
penetration of microflora into the substrate and indirectly
weaken and increase rock
porosity (Trenhaile, 1987; Naylor et al., 2012). Rock borers
play a direct and indirect
role in the disintegration of rocky substrates, particularly in
the lower portions of the
intertidal zone. Boring directly removes some rock material, but
the rest is left
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susceptible to breakdown by wave action and other destructive
mechanisms (Trenhaile,
1987). Borers also enhance the rock environment for algal
colonization, and increase
the area of rock surface exposed to other physical and chemical
processes (McLean,
1974). The indirect role of rock borers may be of greater
significance to the destruction
of coastal rocks than is the direct removal of material.
4.4. Bioerosion and Habitat Heterogeneity
The higher number of perforations (providing refuge) and the
time a cliffs tidal zone
remains submerged, determined the higher richness and abundance
of bioeroders
inhabiting the burrows of the cliffs lower tidal zones. The tide
determines how long a
substrate is underwater or exposed (subject to desiccation)
(Trenhaile, 1987), which
partly depends on the tides being diurnal, semi-diurnal, or
mixed (Johnson and Sparrow,
1961). This permits less desiccation, and changes in temperature
and salinity (Palmer et
al., 2003) in the cliffs lower zones.
Biodiversity, in terms of number of species, is higher when
suitable microhabitats for
vagile species are present in addition to those available for
sessile species. Bivalve and
crustacean burrows provide more shelter for vagile species than
irregularities in the
naturally occurring substratum (such as crevices), and thus
enhance the abundance and
diversity of intertidal species low on the shore (Pinn et al.,
2008). The use of crevices as
shelter was seen in both cliffs by the presence of eight species
of fish during low tide:
Pisodonophis daspilotus, Cerdale ionthas, Gobulus hancocki and
Erotelis armiger in
IC; and Clarkichthys bilineatus, Cerdale paludicola, Microdesmus
dipus and
Pythonichthys asodes in EC. The fact that both cliffs are
located near mangrove zones,
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which are known to act as nurseries, explains why Grapsidae,
Porcellanidae and
Ocypodidae crustacean megalops were found in these sites.
4.5. Further studies on tropical rocky cliffs
Sea cliff erosion in the tropics is an understudied subject and
there is a dearth of
information on erosion rates and dynamics. Although it is a
difficult task, the
understanding of the relative contribution of wave impact and
abrasion to total erosion
rates through field measurements requires further study (Moses,
2013). Furthermore, the
effects of climate change on erosion rates need to be more
thoroughly studied, because
the predicted increase of storm activity and/or intensity,
sea-level rise and the
interaction of both could contribute significantly to erosion
(Phillips and Jones, 2006).
To assess and predict the impacts of climate change, the
understanding of bioerosion
dynamics needs to be expanded to harder igneous and sedimentary
rocks because
studies have been largely limited to recent and relatively weak
beach rock and reef
limestone (Moses, 2013; Moses et al., 2014).
Another item that could have important consequences particularly
for rocky cliffs in
the Colombian Pacific is the modification of seawater chemistry
by organisms
inhabiting the burrows. pH reduction during night hours (as an
imbalance between
photosynthesis during the day and respiration during the night)
would increase the
solubility of calcium carbonate in the rocks and facilitate
their degradation. This process
needs to be further studied.
5. CONCLUSIONS
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Bioerosion is a process in which biological, geological, and
geomorphological factors
interact. For the Pacific coast of Colombia, the abundance of
bioeroders (biological
factor) and tidal zone (physical factor) were the most
influential eroding factors on cliffs
sheltered from wave action. On the other hand, tidal zone and
richness of bioeroders
(also a biological factor) were the most important in
determining erosive volumes in
cliffs exposed to wave action. Rock composition in IC presented
smaller grain sizes
than EC, resulting in more porous and perforated rocks. The
highest abundance of
bioeroding organisms was found in the lower tidal zones of both
cliffs because this zone
stays under water for a longer period of time, providing vital
conditions for the fauna
that takes refuge inside the cliffs during low tide. Boring
bivalves were less abundant in
this study compared to that of boring crustaceans. We suggest
that the importance of
crustaceans in the bioerosion process needs to be highlighted
because it has always been
given a secondary role. In addition to Alpheus javieri and
Upogebia tenuipollex
(previously reported as borers of the cliffs on the Colombian
Pacific coast), we
recommend that Alpheus villus, Upogebia spinigera and Upogebia
burkenroadi should
be considered as active boring species in cliffs of IC. We also
include Cryptomya
californica, Ensitellops hertleini, and Barnea subtruncata as
active borers for cliffs in
EC.
6. ACKNOWLEDGMENTS
This project was supported and funded by the Universidad del
Valle, Biology
Department, Marine Biology Section, and by internal funding of
the Research
Vicerectory of the Universidad del Valle. We thank Philip A.
Silverstone-Sopkin and
Amparo Viveros for correcting the manuscript, Humberto Maya from
the Biological
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Station of Universidad del Valle in Buenaventura, M. Cuellar, S.
Cobo-Viveros, A.I.
Vsquez, V. Izquierdo, F. Vejarano and the locals from Piangita
who helped during
different times on the extraction of blocks from the cliffs.
Biologists J.F. Lazarus, L.A.
Lpez de Mesa, E. Rubio, L. Herrera from Ecomanglares research
group, and B.
Valencia helped during the flora and fauna identification
process. C. Manrique and N.
Durn from the Civil Engineering School at Universidad del Valle
helped on the
sedimentology processing of the rocks. E. Londoo was very
helpful on statistical
inquiries. Finally, thanks to R. Neira who helped in the
organization of field trips and
experimental design.
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FIGURE LEGENDS
Fig. 1. Geographical location of the cliffs studied. Right:
South America, showing the
location of the Pacific coast of Colombia (Middle). Left.
Buenaventura Bay, showing
the locations of IC (unexposed cliffs) and EC (exposed
cliffs).
Fig. 2. Grain size distribution of A. Unexposed (IC) and B.
Exposed cliffs (EC) on the
Pacific coast of Colombia. The legend on the right represents
each tidal zone: low (L),
middle (M), and high (H).
Fig. 3. Abundance of bioeroding species in the low, middle and
high tidal zones of A.
Unexposed cliffs (IC) and B. Exposed cliffs (EC) in the Pacific
coast of Colombia. The
category Others groups bioeroding fauna with total abundances of
less than 10
individuals.
Fig. 4. p-value, correlation (r) and determination coefficients
(r2) found for significant
linear correlations in unexposed cliffs (IC; A-B) and exposed
cliffs (EC; C-D) on the
Pacific coast of Colombia. Biodegraded volume was negatively
correlated with tidal
zone in both cliffs (A and C), and positively correlated with
abundance of bioeroders in
IC (B) and with richness of bioeroders in EC (D).
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TABLE HEADINGS
Table 1. Grain size composition (percentage) of unexposed (IC)
and exposed cliffs
(EC) in the Pacific coast of Colombia.
Table 2. Percentage of perforations due to bioerosion (relative
to total volume) and
percentage of porosity (relative to volume of solids + volume of
pores) found for the
three tidal zones of unexposed cliffs (IC) and exposed cliffs
(EC) in the Pacific coast of
Colombia. Richness and abundance of bioeroding species of the
studied cliffs is also
shown.
Table 3. Multiple regression analysis comparing volumes of
perforation with tidal zone,
volume of natural porosity of the rocks, richness and abundance
of bioeroding fauna in
unexposed (IC) and exposed cliffs (EC). Multiple correlation
coefficient (R), multiple
determination coefficient (R2), and adjusted coefficient of
determination are shown
(Ra2), as well as results of ANOVA for the multiple regression
data.
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Table A1. Temperature Correction Factor (CT) applied to the
actual hydrometer
readings. Source: Bowles (1981a).
Temperature
C CT
15 -1.10
16 -0.90
17 -0.70
18 -0.50
19 -0.30
20 0.00
21 +0.20
22 +0.40
23 +0.70
24 +1.00
25 +1.30
26 +1.65
27 +2.00
28 +2.50
29 +3.05
30 +3.80
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Table A2. Correction factor a determined for unit weight of
solids (specific gravity).
Source: Bowles (1981a).
Unitary weight soil solids
(g/cm3)
Correction factor
a
2.85 0.96
2.80 0.97
2.75 0.98
2.70 0.99
2.65 1.00
2.60 1.01
2.55 1.02
2.50 1.04
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Table A3. Values of K used in Equation 7 for several
combinations of unitary weights and temperatures to compute the
particle diameter in
the Hydrometer Analysis. Source: Bowles (1981a).
Temperature
(C)
Gs
2.45 2.50 2.55 2.60 2.65 2.70 2.75 2.80 2.85
16 0.01510 0.01505 0.01481 0.01457 0.01435 0.01414 0.03940
0.01374 0.01356
17 0.01511 0.01490 0.01460 0.01440 0.01420 0.01400 0.01380
0.01360 0.01338
18 0.01492 0.01470 0.01440 0.01420 0.01400 0.01380 0.01360
0.01340 0.01321
19 0.01474 0.01450 0.01430 0.01400 0.01380 0.01360 0.01360
0.01340 0.01305
20 0.01456 0.01430 0.01410 0.01390 0.01370 0.01340 0.01330
0.01310 0.01289
21 0.01438 0.01410 0.01390 0.01370 0.01350 0.01330 0.01310
0.01290 0.01273
22 0.01421 0.01400 0.01370 0.01350 0.01330 0.01310 0.01290
0.01280 0.01258
23 0.01404 0.01380 0.01360 0.01340 0.01320 0.01300 0.01280
0.01260 0.01243
24 0.01388 0.01370 0.01340 0.01320 0.01300 0.01280 0.01260
0.01250 0.01229
25 0.01372 0.01350 0.01330 0.01310 0.01290 0.01270 0.01250
0.01230 0.01215
26 0.01357 0.01330 0.01310 0.01290 0.01270 0.01250 0.01240
0.01220 0.01201
27 0.01342 0.01320 0.01300 0.01280 0.01260 0.01240 0.01220
0.01200 0.01188
28 0.01327 0.01300 0.01280 0.01260 0.01240 0.01230 0.01210
0.01190 0.01175
29 0.01312 0.01290 0.01270 0.01250 0.01230 0.01210 0.01200
0.01180 0.01162
30 0.01298 0.01280 0.01260 0.01240 0.01220 0.01200 0.01180
0.01170 0.01149
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Table A4. Values of L (effective hydrometer depth, in cm) for
use in Stokes formula to
determine particle diameters using an ATSM 152H hydrometer.
Source: Bowles
(1981a).
Original hydrometer reading
(only corrected for meniscus)
Effective depth
L (cm)
Original hydrometer reading
(only corrected for meniscus)
Effective depth
L (cm)
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
16.3
16.1
16.0
15.8
15.6
15.5
15.3
15.2
15.0
14.8
14.7
14.5
14.3
14.2
14.0
13.8
13.7
13.5
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
11.2
11.1
10.9
10.7
10.6
10.4
10.2
10.1
9.9
9.7
9.6
9.4
9.2
9.1
8.9
8.8
8.6
8.4
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18
19
20
21
22
23
24
25
26
27
28
29
30
13.6
13.2
13.0
12.9
12.7
12.5
12.4
12.2
12.0
11.9
11.7
11.5
11.4
49
50
51
52
53
54
55
56
57
58
59
60
8.3
8.1
7.9
7.8
7.6
7.4
7.3
7.1
7.0
6.8
6.6
6.5
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Annex A
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Annex B