Computer modeling AUTHORS

Andrew D. La Croix  Applied Research in

bioturbation: The creation Ichnology and Sedimentology (ARISE) Group,
Department of Earth Sciences, Simon Fraser
of porous and permeable University, Burnaby, British Columbia, Canada,
V5A 1S6; alacroix@sfu.ca
fluid-flow pathways Andrew La Croix received his M.Sc. degree in
geology from the University of Alberta and
Andrew D. La Croix, Murray K. Gingras, is currently a Ph.D. student at Simon Fraser Uni-
Shahin E. Dashtgard, and S. George Pemberton versity in Burnaby, British Columbia. Andrew’s
research interests include ichnology, sedimen-
tology, stratigraphy, petroleum geology, and
basin analysis. The focus of his Ph.D. dissertation
ABSTRACT is the sedimentological and ichnological charac-
teristics of the tidal-fluvial transition in rivers.
Computer modeling of trace fossils (Skolithos, Thalassinoides,
Planolites, Zoophycos, and Phycosiphon) and ichnofacies (Sko- Murray K. Gingras  Ichnology Research
Group (IRG), Department of Earth and Atmo-
lithos, Cruziana, and Zoophycos ichnofacies) is undertaken to
spheric Sciences, University of Alberta, Edmonton,
assess the impact of bioturbation on porosity and perme- Alberta, Canada, T6G 2E3;
ability trends in sedimentary media. Model volumes are mgingras@ualberta.ca
randomly populated with the digitally modeled trace fossils Murray Gingras received his diploma in mechanical
to test for connectivity between burrows. The probability of ver- engineering technology from the Northern
tical and lateral interconnections is compared with bioturbation Alberta Institute of Technology in 1987, his B.Sc.
intensity. degree from the University of Alberta in 1995,
The results of the simulations indicate that biogenic flow and his Ph.D. from the University of Alberta in
1999. Gingras has worked professionally in the
networks develop at low bioturbation intensity, between 10
hydrocarbon industry at the Northern Alberta
and 27.5% bioturbation (BI-2). However, the efficiency of Institute of Technology and as an assistant
connectivity is controlled by the architecture of the burrows. professor at the University of New Brunswick.
For all trace-fossil and ichnofacies models, regardless of trace- His research focuses on applying sedimentology
fossil orientation, continuous horizontal and vertical con- and ichnology to sedimentary rock successions
nectivity across the sediment volume is achieved within a 0 to as a paleoecological tool, a reservoir-development
tool, and in process-driven sedimentology.
10% range in bioturbation.
In subsurface aquifers and petroleum reservoirs, the pres- Shahin E. Dashtgard  Applied Research in
ence of bioturbation can significantly influence fluid flow. In Ichnology and Sedimentology (ARISE) Group,
particular, for marine sedimentary rocks, where burrows are Department of Earth Sciences, Simon Fraser
University, Burnaby, British Columbia, Canada,
more permeable than the surrounding matrix, a greater degree
V5A 1S6; sdashtga@sfu.ca
of three-dimensional burrow connectivity can produce pre-
Shahin Dashtgard is an assistant professor in
ferred fluid-flow pathways through the rock. Recognizing
the Department of Earth Sciences at Simon
these flow conduits may enable optimization of resource ex- Fraser University. His research focuses on char-
ploitation or may contribute to increasing reserve estimates acterizing modern depositional environments
from previously interpreted nonreservoir rock. as analogs for the rock record. Before his Ph.D.,
Shahin worked as a petroleum geologist in
western Canada.

Copyright ©2012. The American Association of Petroleum Geologists. All rights reserved.
Manuscript received March 8, 2011; provisional acceptance June 16, 2011; revised manuscript received
July 11, 2011; final acceptance July 14, 2011.
DOI:10.1306/07141111038

AAPG Bulletin, v. 96, no. 3 (March 2012), pp. 545–556 545

 S. George Pemberton  Ichnology Research
Group (IRG), Department of Earth and Atmo-
spheric Sciences, University of Alberta, Edmonton,
Alberta, Canada, T6G 2E3;
george.pemberton@ualberta.ca
S. George Pemberton is a professor in the
Department of Earth and Atmospheric Sciences
at the University of Alberta. He is a Fellow
of the Royal Society of Canada and holds a
Canada Research Chair in Petroleum Geology
(Natural Sciences and Engineering Research
Council). George’s fields of research and
expertise are in ichnology and the investigation
of animal-sediment interactions in both recent
and ancient environments. Current research
activities include the application of trace-fossil
studies in sequence stratigraphy and the
exploration and exploitation of hydrocarbons.
Recent research activities involve emphasis on
the Cardium and Viking formations, the
Athabasca and the Cold Lake oil sands of
Alberta, as well as the offshore Hibernia, Ben
Nevis, Terra Nova, and Venture fields.
ACKNOWLEDGEMENTS
We thank Nexen Inc., ConocoPhillips Canada
Ltd., and Total E&P Canada Ltd. for their
generous financial support of the project. Allen
Shen was instrumental to this study, with his
computer modeling expertise. Carl Mendoza
greatly improved the concepts and methods
used herein with thoughtful discussions. Finally,
we thank AAPG Editor Stephen Laubach,
Luis Buatois, and one anonymous reviewer for
their helpful comments.
The AAPG Editor thanks the following reviewers
for their work on this paper: Luis A. Buatois
and an anonymous reviewer.
546 Computer Modeling Bioturbation
INTRODUCTION
Burrows filled with sand can act as porous and permeable
flow pathways through otherwise low-permeability sedimen-
tary media. Provided that burrows are connected, fluid trans-
mission through the permeable biogenic conduits can be
considerable (e.g., Arab-D reservoir, Pemberton and Gingras,
2005; Biscayne aquifer, Cunningham et al., 2009). To better
establish the relationship between burrow type, bioturbation
intensity (i.e., percentage of the sediment volume occupied
by burrows), and trace-fossil connectivity, computer mod-
eling of several trace fossils and trace-fossil assemblages is
undertaken. The models use various three-dimensional (3-D)
shapes designed to emulate burrows and combinations of
shapes to emulate the natural variability expected in bio-
turbated media. The results, at least conceptually, show the
minimum threshold of burrowing intensity required to pro-
duce porous and permeable flow networks. The effects of trace-
fossil shape, size, and orientation on connections are also
evaluated.
Trace fossils have been shown to significantly affect the
porosity and permeability trends in aquifers and petroleum
reservoirs (Gingras et al., 1999, 2002, 2004, 2007; Pemberton
and Gingras, 2005; Cunningham et al., 2009; Tonkin et al.,
2010). Burrowing organisms affect the primary depositional
fabric and diagenesis of sediment and, thereby, the porosity
and permeability distribution (Meadows and Tait, 1989;
Gingras et al., 2002; Bastardie et al., 2003; Katrak and Bird,
2003; Gingras et al., 2004; Pemberton and Gingras, 2005). In
sufficiently burrowed horizons, networks of burrows have the
potential to provide a flow conduit through which subsurface
fluids can be extracted.
In one of the first studies examining the effects of trace
fossils on fluid flow, Gingras et al. (1999) used computer sim-
ulations combined with laboratory and field testing to identify
three factors that have the strongest effect on fluid flow through
burrowed sediments; these are (1) the permeability contrast
between the burrows and surrounding matrix, (2) the degree
of interconnection between burrows, and (3) the overall den-
sity of burrowing. However, the relationship between density
of burrowing and interconnections between burrows was not
defined, and the minimum intensity of burrowing to form
discrete networks was not established. It is essential to un-
derstand how connectivity relates to burrowing density to re-
cognize and model biogenic flow pathways through biotur-
bated reservoirs. This study addresses this issue by simulating
connectivity in computer-generated burrowed volumes.
Figure 1. The five digital trace fossil types used in the simulations to test for vertical and lateral burrow connectivity: (A) a vertical
cylinder representing Skolithos, (B) a horizontal sinusoidal cylinder representing Planolites, (C) three cylinders connected at a central axis
to represent Thalassinoides, (D) a descending beaver-tail shape to represent Zoophycos, and (E) a meandering shape with cylindrical
cross section representing Phycosiphon. All the digital traces are shown inside the model’s 1-m3 (35 ft3) volume.

METHODS The DTs were further grouped into broad cate-

The model design is composed of two parts: (1) the
digital construction of five geometric shapes (digital
trace fossils [DTs]) intended to represent common
burrow morphologies and (2) the random popula-
tion of a sediment volume with various combina-
tions of the DTs and the subsequent assessment of
interconnectivity.
Digital Trace Fossil Construction
Five 3-D shapes were created to compare the spec-
trum of morphologies from simple to complex trace-
fossil forms. The ichnogenera—Skolithos, Planolites,
Thalassinoides, Zoophycos, and Phycosiphon—modeled
as DTs, are some of the most commonly observed
trace fossils in marine siliciclastic environments.
gories intended to represent the Skolithos ichno-
facies (Skolithos, Thalassinoides), Cruziana ichno-
facies (Planolites, Skolithos, Thalassinoides), and
Zoophycos ichnofacies (Zoophycos, Phycosiphon).
Notably, the construction of the digital trace fossils
and their grouping into ichnofacies is inherently
simplistic and, as a result, the combination of DTs
used to represent each ichnofacies does not reflect
the true diversity encountered in modern or rock
record burrowed sedimentary media. Therefore,
the proportions of the various DTs are the main
difference between ichnofacies, and especially the
Skolithos and Cruziana ichnofacies.
The Skolithos DT is a vertical cylinder, 10 cm
(4 in.) high × 1 cm (0.4 in.) diameter (Figure 1A).
The cylindrical shape was chosen because the in-
ferred filter-feeding behavior represented by Sko-
lithos is typically manifest as a simple vertical or

La Croix et al. 547

inclined tube that is often filled with sediment
identical with the surrounding matrix.
The Planolites DT is a horizontal sinusoidal
tube with a length of 10 cm (4 in.) and a diameter
of 1 cm (0.4 in.) (Figure 1B). Although Planolites
represents the activity of selective deposit feeders,
where individual burrows can be continuous for
an indefinite length, the length of the DT is spec-
ified, again, to allow for the stepwise increase in
bioturbation intensity within the simulations.
The Thalassinoides DT is a composite structure
produced by joining the central axis of a vertical
cylinder (1 cm [0.4 in.] diameter, 10 cm [4 in.]
high) to two horizontal cylinders with the same di-
mensions. The two horizontal cylinders are oriented
in the same plane, with 120° angles between each
of the limbs (Figure 1C). Thalassinoides commonly
occurs in box-work patterns where individual bur-
rows are difficult to discern from one another. The
shape of the DT was chosen as a way to discretize
the burrow and to allow for the simulations to use
stepwise increases in bioturbation intensity.
The Zoophycos DT is modeled as a beaver-tail
shape. In plan view, a half circle with a 5-cm (2-in.)
radius is affixed to a Bézier surface (a parametric
curve frequently used in computer graphics and
design) that tapers to 1 cm (0.4 in.). The hori-
zontal distance from the center of the half circle to
the tapered tip is 25 cm (10 in.) and the vertical
height, from the base to the tip, is 15 cm (6 in.).
The cross sectional thickness of the Zoophycos DT
is 1 cm (0.4 in.) (Figure 1D). This geometry was
chosen because it provides a simple way to model
the fan-shaped excavations descending from a
central tube, which is the characteristic Zoophycos
morphology.
Finally, the Phycosiphon DT is randomly gen-
erated in 60 steps, where the length of each indi-
vidual DT is unique. In each step, an extra seg-
ment of a tube (diameter 1 cm [0.4 in.]) extends
in a random direction (in 3-D space), a randomly
determined distance between 0 and 4 cm (1.6 in.).
This is a simple way of modeling meandering trace
fossils (Figure 1E), but it falls short of encapsulating
the subtle and intricate details that define individ-
ual ichnogenera. In this particular situation, the
digital trace was over scaled for modeling purposes.
548 Computer Modeling Bioturbation
However, these geometric shapes suffice for a first
pass at numerically modeling bioturbation.
Model Generation and Simulation
The development of burrow networks is evaluated
using a function that tests for vertical and lateral
connections in a volume that is populated with DTs
(Figure 2). The function operates on the principles
of percolation theory (Stauffer and Aharony, 1994).
The critical threshold of percolation (connectivity
from one side of a lattice to another) is dependent
on the probability of DTs occupying sites in the
lattice (i.e., the bioturbation intensity). In theory,
for an infinite lattice, a threshold bioturbation in-
tensity must exist above which percolation always
occurs and below which percolation does not oc-
cur. However, in practice, this is not the case, and
percolation develops within some range in biotur-
bation intensity. This range in the development of
percolation is caused by the model lattice having a
finite structure and the variation between model
configurations and simulations (discussed below).
The percolation function essentially divides the lattice
into two-dimensional sheets and identifies sites that
are occupied by the DTs. On identifying the occupied
sites, it calculates if adjacent sites in the same sheet or
adjacent sheets line up to create a continuous path-
way from one side of the lattice to the other.
The model volume comprises a cube-shape
lattice, with each side of the lattice containing 256
cells. The cell sizes are scaled in relation to the
digital traces, so that the total sediment volume is 1 m3
(35 ft3). The DTs are populated into the lattice with
random placement and random orientations about
the vertical axis and are arranged such that they can
overlap or be only partially contained within the lattice.
Eight models are tested in the simulations, one
for each of the five DT morphologies (Skolithos,
Planolites, Thalassinoides, Zoophycos, and Phycosi-
phon) and one model to represent each of the three
ichnofacies (Skolithos, Cruziana, and Zoophycos).
The model representing the Skolithos ichnofacies is
a combination of Skolithos DTs and Thalassinoides
DTs, occurring in a 3:1 ratio. The Cruziana ichno-
facies is modeled as a combination of Planolites DTs,
Skolithos DTs, and Thalassinoides DTs, set up in
Figure 2. The stepwise population of a model volume with various numbers of the Planolites digital trace: (A) 1, Planolites; (B) 2,
Planolites; (C) 5, Planolites; (D) 10, Planolites; (E) 25, Planolites; and (F) 50, Planolites. Every model configuration populates the digital
traces in random locations and orientations about the vertical axis.
the ratio 2:1:1, respectively. Finally, the Zoophycos
ichnofacies comprises Zoophycos DTs and Phyco-
siphon DTs in a 1:1 ratio. These models do not cover
the full complexity of the trace-fossil assemblages
observed in the various ichnofacies; however, they
provide a good first pass for establishing the effects
of burrow morphology and trace associations on
burrow connectivity.
The connectivity simulations are conducted on
several configurations for each of the eight models.
Nine to 22 configurations were required for each
model. Every configuration contains a different
number of DTs that are populated into the vol-
ume. Subsequent configurations progressively in-
crease the number of DTs from one to several thou-
sands. The variation between configurations of the
same model allows for a range of bioturbation in-
tensities (percent volume occupied by DTs) to be
tested. Burrow connectivity is simulated on each
model configuration 1000 times (i.e., a Monte Carlo
Test). In each simulation, the distribution of DTs
is randomly generated and the probability of con-
nectivity is calculated. New configurations are added
until the probability of vertical and lateral con-
nectivity unity (i.e., almost surely a continuous con-
nection between burrows across the entire cube
volume exists).
RESULTS
The results of the simulation experiments are pre-
sented graphically in Figures 3 and 4 and described
below. Three statistical values are tracked for each
model simulation: the bioturbation intensity (percent
bioturbation) at 0.1 probability of interconnections
(P10), the bioturbation intensity at 0.5 probability of
interconnections (P50), and the bioturbation intensity
at 0.9 probability of interconnections (P90). The
P10 is considered to represent the value at which
La Croix et al. 549
550
Computer Modeling Bioturbation

Figure 3. Graphs of the probability of connectivity versus intensity of bioturbation for (A) the Skolithos model, (B) the Planolites model, (C) the Thalassinoides model, (D) the
Zoophycos model, and (E) the Phycosiphon model. The solid lines represent lateral connections between burrows, whereas the dashed lines represent vertical connectivity. Ac-
companying the graphs is a table that displays the P10, P50, and P90 values of bioturbation intensity for vertical and lateral connectivity in the various models.
Figure 4. Graphs of the probability of connectivity versus intensity of bioturbation for (A) the Skolithos ichnofacies model, (B) the Cruziana ichnofacies model, and (C) the Zoophycos
ichnofacies model. The solid lines represent lateral connections between burrows, whereas the dashed lines represent vertical connectivity. Accompanying the graphs is a table that
preferential percolation of fluids through the bur-
row network (assuming the burrows are more
permeable than the surrounding matrix) occurs
with low certainty and, therefore, represents the
initiation of connectivity. The P50 is the midpoint
value for percolation, where a moderate certainty
exists in the connectivity of burrows. The P90 is
the value at which a high degree of certainty exists
that a continuous burrow network has developed
(i.e., a complete biogenic conduit that can trans-
mit flow). However, these values are chosen as
arbitrary cutoffs for the purposes of this study and
do not hold any significance in percolation theory.
The values for P10, P50, and P90 are tabulated
and accompany the graphs of bioturbation in-
tensity versus probability of percolation and/or
displays the P10, P50, and P90 values of bioturbation intensity for vertical and lateral connectivity in the various models. interconnections (Figures 3, 4).

Digital Trace Fossil Models

Twenty-two configurations are implemented in

the simulations of the Skolithos model. The number
of DTs is systematically varied from 0 to 61,000.
The configuration consisting of 61,000 DTs corre-
sponds to 33% bioturbation. Although the sum of
61,000 individual Skolithos DT volumes is 47.6% of
the model volume (volume, 1 m3 [35 ft3]), in the
simulations, they occupy just 33% because many
DTs overlap and/or some DTs are only partially
contained within the modeled area. The differ-
ence between burrow volumes and percent bio-
turbation was repeated in all models.
The results of the Skolithos model simulations
(Figure 3A) indicate that vertical connectivity—
connections between burrows spanning from the
bottom to the top of the volume—initiates at ap-
proximately 17.5% bioturbation (P10). With in-
creasing bioturbation intensity, and assuming that
the burrows are more permeable that the surround-
ing matrix, the probability of percolation through
the burrows rises abruptly, such that the P50 occurs
at 18% bioturbation. By 20% bioturbation (P90),
the probability of interconnections between bur-
rows is high. Lateral connectivity begins at 22.5%
bioturbation (P10) and P50 occurs at 24% bur-
rowing. At 26.5% bioturbation (P90), lateral bur-
row connections are essentially fully developed.

La Croix et al. 551

Vertical and lateral connectivity become fully estab-
lished within a 5% bioturbation range of each other.
The Planolites model simulations necessitate
the use of 15 configurations that vary the number
of DTs in the sediment volume from 0 to 40,000.
In the configuration with the greatest bioturbation
intensity, corresponding to 40,000 Planolites DTs,
32% of the model’s volume is occupied by DTs.
The results of the Planolites model simulations
(Figure 3B) show that lateral connectivity is es-
tablished first, beginning at approximately 10%
bioturbation (P10). With increasing bioturbation
intensity, the probability of interconnections in-
creases abruptly (i.e., P50 = 12% bioturbation). By
13% bioturbation (P90), lateral burrow networks
are fully established. The onset of vertical connec-
tivity is initiated at approximately 18% bioturbation
(P10). The development of vertical interconnec-
tions increases with increasing burrowing inten-
sity, such that the P50 occurs at 20% bioturbation.
At 22% bioturbation (P90), vertical connections
between burrows are fully in place. Interconnec-
tivity between DTs, both vertically and laterally, is
established within a range in bioturbation inten-
sity that is less than 10%.
Nine configurations are implemented in the
Thalassinoides model simulations, where the num-
ber of DTs is incrementally increased from 0 to 4750.
At its most populous configuration, 12% of the
model’s volume consists of DTs. The simulation re-
sults (Figure 3C) show that vertical interconnections
begin to develop at 8% bioturbation (P10). With in-
creasing bioturbation intensity, vertical connectivity
improves, such that P50 occurs at 9.5% bioturbation.
Continuous vertical burrow networks are fully con-
structed at 10% bioturbation (P90). Lateral con-
nectivity begins at approximately 9.5% burrowing
(P10) and increases to P50 at 11% bioturbation. Full
establishment of lateral burrow networks exists by
11.5% burrowing (P90). In this model, network
development in both directions occurs within a
3% range in bioturbation intensity.
Sixteen configurations are used for the simu-
lations of the Zoophycos model, wherein the vol-
ume is populated with varying numbers of DTs
between 0 and 26,000. At the maximum biotur-
bation intensity (26,000 DTs), 18% of the volume
552 Computer Modeling Bioturbation
is occupied by DTs. The results of the Zoophycos
model simulations (Figure 3D) indicate that lateral
connections begin to form at 7.5% bioturbation
(P10) and the P50 occurs at 9% bioturbation. By
10% bioturbation (P90), lateral connections are
pervasive. Vertical connectivity ensues at higher
bioturbation intensity, at approximately 11% bur-
rowing (P10). The P50 occurs at 13% bioturbation,
and vertical connectivity is fully established by 14.5%
bioturbation (P90). Connected vertical and lateral
burrow networks develop within a range of only 5%
bioturbation.
The final digital trace fossil model the Phyco-
siphon model, is composed of 13 configurations that
vary the number of DTs from 0 to 14,000. In the most
populated configuration, the DTs occupy 15% of the
total volume. The results of the simulations using the
Phycosiphon model (Figure 3E) show that vertical and
lateral connectivity occurs within the same range of
bioturbation intensity. Connections begin to form at
approximately 7% burrowing (P10), and increase in
probability within a short range: P50 occurs at 9%
bioturbation. At approximately 10% bioturbation
(P90), vertical and lateral burrow networks are fully
established. The burrow connectivity increases rapidly
within a range of approximately 3% bioturbation.
Ichnofacies Models
Fourteen configurations are required in the Skolithos
ichnofacies model simulations. The number of DTs
is incrementally increased in each configuration
until the model contains 24,000 DTs, correspond-
ing to 27% of the total volume. The results of the
Skolithos ichnofacies model simulations (Figure 4A)
indicate that vertical connectivity between burrows
begins at approximately 13% bioturbation (P10).
With increasing bioturbation intensity, the prob-
ability of interconnections increases rapidly, such
that P50 occurs at 16% burrowing. At 18% bio-
turbation (P90), connections are fully established.
Lateral connectivity ensues at 17% bioturbation (P10)
and increases to P50 at 19% bioturbation. By 21%
burrowing (P90), lateral burrow networks always
develop. In the vertical and lateral directions,
burrow network creation occurs within a 10%
range in bioturbation.
 The simulations of the Cruziana ichnofacies
model require nine configurations. At the maxi-
mum bioturbation intensity of 27%, 12,000 DTs
are populated into the model volume. The sim-
ulation results from the Cruziana ichnofacies model
(Figure 4B) illustrate that vertical connections begin
to form at approximately 9% bioturbation (P10).
The P50 occurs at 12% burrowing and P90 at 14%
bioturbation. By P90, vertical burrow networks are
commonplace. Lateral connectivity initiates at 10%
bioturbation (P10) and increases abruptly to P50
at 12% bioturbation. By 14% bioturbation (P90),
lateral interconnections are fully established. The
graphs of vertical and lateral connectivity track one
another very closely and indicate that burrow net-
works develop within a small range in bioturbation
intensity, somewhere between 5 and 10%.
Nineteen configurations are used in the simu-
lations of the Zoophycos ichnofacies model. The
bioturbation intensity ranged from 0 to 20%, cor-
responding to 12,000 DTs occurring in the model
volume. The results of the Zoophycos assemblage
simulations (Figure 4C) show that interconnec-
tivity in the lateral direction begins when the bio-
turbation intensity is 8% (P10). With an increasing
number of DTs, the likelihood of connections
increases, such that the P50 is reached at 10% bio-
turbation. Burrow networks are completely estab-
lished at 11% bioturbation (P90). Vertical con-
nectivity begins at 8% burrowed (P10) but increases
in probability at a slightly slower rate (the slope
of the curve is lower) than lateral connectivity. The
P50 occurs at 12% burrowing. By 17% bioturba-
tion (P90), lateral connectivity is fully developed.
In both instances, burrow connections form within
a small range in bioturbation intensity, within 10%.
DISCUSSION
Burrow Connectivity at Low
Bioturbation Intensity
The simulations show that connected burrows
form continuous networks at low bioturbation
intensity. In the models where connectivity was
established most rapidly (i.e., began with the
lowest number of DTs), the P90 of connectivity
occurred at 10% bioturbation. This was developed
in the Thalassinoides and Phycosiphon models for
connections in the vertical direction, and in the
Zoophycos and Phycosiphon models for lateral con-
nectivity. Where connectivity occurred least rapidly
(i.e., connections occurred at higher bioturbation
intensity), the P90 of connectivity was at 27.5%
burrowing. The Skolithos model was the only model
to develop lateral connectivity at these levels of
bioturbation; all the other models developed both
lateral and vertical connectivity at lower biotur-
bation intensities. The low (10%) bioturbation in-
tensity and high (27.5%) bioturbation intensity end
members illustrate the conceptual range in biotur-
bation intensity in which connected burrow net-
works are produced (Figure 5). The range falls
within ichnofabric index 3 (II-3) and bioturbation
index 2 (BI-2) (Droser and Bottjer, 1986; Taylor
and Goldring, 1993).
Burrow Architecture Affects Connectivity
In addition to the bioturbation intensity, the shape
and orientation of trace fossils impart an important
control on the development of burrow intercon-
nections. Many trace fossils common in marine
siliciclastic environments consist of narrow (commonly
<1 cm [<0.4 in] diameter) shafts, tubes, and tunnels
with vertical, inclined, or horizontal orientations (e.g.,
Skolithos, Planolites, Palaeophycus, Arenicolites, Tha-
lassinoides, Ophiomorpha). Other ichnogenera are
composed of more complex morphologies that are
characterized by variously meandering, spiraling, or
lobate geometries that incorporate horizontal and
vertical elements in their structures (e.g., Phycosiphon,
Zoophycos, Gyrolithes, Psilonichnus, Asterosoma).
The extent that burrows extend or branch in
any particular direction determines the potential
for connecting with other burrows in that direction.
Ichnogenera with architectures having significant
vertical components most efficiently connect ver-
tically, whereas ichnogenera with significant hori-
zontal components are more capable of connect-
ing laterally. Burrows that have similar proportions
of horizontal and vertical components are capable
of connecting vertically and laterally with similar
La Croix et al. 553
Figure 5. A chart comparing the zone of burrow network development (based on P90 values) with two popular classification schemes
for bioturbation intensity: the ichnofabric index (II) (Droser and Bottjer, 1986) and the bioturbation index (BI) (Taylor and Goldring,
1993). The comparison reveals that II-3 and BI-2 characterize the range of bioturbation in which burrows become well connected.

efficiency. The potential for establishing connections
with neighboring burrows is heightened when ani-
mal movement has occurred in all three dimensions.
The relationship between burrow architecture
and percolation (assuming permeable burrows and an
impermeable matrix) is evident in the simulation re-
sults, where vertical connectivity developed in the
models in the following order (lowest to highest
bioturbation intensity): Phycosiphon, Thalassinoides,
Cruziana ichnofacies, Zoophycos ichnofacies, Zoo-
phycos, Planolites, Skolithos ichnofacies, and Skolithos.
The controls of shape and orientation can also be
observed in the order in which lateral connectivity
developed (lowest to highest bioturbation intensity):
Zoophycos and Phycosiphon, Zoophycos ichnofacies,
Thalassinoides, Planolites, Cruziana ichnofacies, Sko-
lithos ichnofacies, and Skolithos. This order reflects
the positive effects of increasing burrow branching
or meander on connectivity. Percolation progresses
first in the models consisting of forms that sub-
stantially extend in the vertical and lateral directions,
followed by forms confined to one direction. The
Skolithos ichnofacies and Skolithos DT models were
the last to exhibit burrow interconnectivity because
the computer-generated Skolithos is narrow and or-
iented vertically. As a result, the potential for lateral
connections with nearby DTs is minimal, and ver-
tical percolation only results when Skolithos DTs are
stacked atop one another.
Unconnected Burrows Still Affect Permeability
Although interconnections between burrows forming
porous and permeable networks are critical for fluid
flow, the presence of non-interconnected burrows
can still have marked effects on bulk permeability. In a
study investigating coupled fracture-matrix fluid
flow, Philip et al. (2005) showed that in reservoirs
with weakly connected fractures, fluid flow was very
sensitive to the lengths of fractures, whereas the
aperture of fractures (i.e., fracture permeability) was a
less important consideration. The fracture length is
important because it ultimately determines the
proportion of fluid flow that occurs through the

554 Computer Modeling Bioturbation

matrix; the longer that fractures are, the more fluid
that can bypass a less permeable matrix. Similarly, in
bioturbated sedimentary media, where burrows are
more permeable than the surrounding matrix, burrows
may provide fluid-flow pathways through the matrix,
even if they are poorly connected. In these cases, the
length or height distribution of the burrows would be
the major factor controlling bulk permeability.
Comparison of Burrow-Associated Percolation
and Examples of Permeable Bioturbated
Sedimentary Strata
Permeable burrow networks may be more com-
mon in bioturbated sediments than many petro-
leum geologist recognize, mainly because connec-
tions between burrows occur at low bioturbation
intensities (between 10 and 30% burrowed; BI-2 or
II-3). In comparison, burrowed marine sediments
commonly exhibit moderate bioturbation inten-
sities (as much as 60%; BI-3 or II-4). Provided that
a burrow’s fill is more permeable than the sur-
rounding matrix (e.g., sand-filled burrows in mud),
porous and permeable burrow networks can de-
velop. In the following two paragraphs, the results of
the computer models are compared with two ex-
amples of porosity- and permeability-enhanced bio-
turbated media.
At Willapa Bay, Washington, Gingras et al. (1999)
explored how Glossifungites surfaces, with well-
developed sand-filled Thalassinoides networks, affect
the permeability of muddy substrates. Through slug
and laboratory flow tests supplemented with com-
puter modeling, Gingras et al. (1999) showed that
the bulk horizontal permeability (Kh) was greatly
affected by the connectivity between burrows. When
connections existed, Kh approached vertical perme-
ability through the burrows (Kv), leading to isotropic
bulk permeability. However, when insignificant bur-
row overlap occured, Kh was substantially lower than
Kv. Overall, permeability enhancement was minimal
until the density of burrowing reached 10% (Gingras
et al., 1999), which is a similar value to that de-
termined from the Thalassinoides connectivity simu-
lations in this study.
Another example of the effects of burrow net-
works on fluid transmission is the Arab-D reservoir
in the Ghawar field of Saudi Arabia (Pemberton
and Gingras, 2005). In the Arab-D reservoir, deeply
penetrating (up to 2 m [6.6 ft]) Thalassinoides are
filled with permeable sucrosic dolomite that acts
as a vertical fluid conduit through a relatively im-
permeable micrite matrix. The burrows are con-
centrated along stratigraphic surfaces, interpreted
as firmgrounds and, therefore, represent the Glos-
sifungites ichnofacies. The intensity of bioturbation
is in the range of 50% (BI-3 or II-4), well above the
intensity required to produce interconnected bur-
row networks. The burrowed Glossifungites sur-
faces are stratiform “Super-K” zones through which
as much as 70% of the oil is produced (Pemberton
and Gingras, 2005).
CONCLUSIONS
Computer simulations using digitally modeled trace
fossils and trace-fossil assemblages were used to as-
sess the bioturbation intensity at the onset of burrow
network development. The simulation experiments
show that burrow connections begin to form a 3-D
system at bioturbation intensities as low as 10% (BI-2).
Connectivity is enhanced with increasing burrowing
intensity, with horizontal and vertical burrow net-
works being established in all experiments by 27.5%
burrowed (BI-2). For any particular trace-fossil as-
semblage, burrow connectivity develops within a
range in bioturbation intensity of less than 10%.
Burrow architecture is a contributing factor that
helps control the direction of connectivity. Trace-
fossil assemblages that consist of vertical or inclined
forms result in preferential vertical percolation.
Assemblages comprising horizontal ichnogenera
construct lateral connections before connecting in
the vertical direction. Therefore, burrow suites of
mixed morphology (vertical, inclined, and hori-
zontal ichnogenera) such as those characterized by
the Cruziana ichnofacies are subject to isotropic
percolation.
Connections between burrows (to form net-
works) commonly occur without mention or are
overlooked in some ichnological studies. However,
because burrows begin to form connections at a low
bioturbation intensity, their presence in bioturbated
La Croix et al. 555
strata is probably underappreciated. Perhaps the
greatest potential of porous and permeable burrow
networks is to connect otherwise hydraulically iso-
lated strata and to increase isotropic permeability in
aquifers and hydrocarbon reservoirs.
REFERENCES CITED
Bastardie, F., Y. Capowiez, J. R. de Dreuzy, and D. Cluzeau,
2003, X-ray tomographic and hydraulic characterization
of burrowing by three earthworm species in repacked
soil cores: Applied Soil Ecology, v. 24, p. 3–16, doi:10
.1016/S0929-1393(03)00071-4.
Cunningham, K. J., M. C. Sukop, H. Huang, P. F. Alvarez, H. A.
Curran, R. A. Renken, and J. F. Dixon, 2009, Prominence
of ichnologically influenced macroporosity in the karst
Biscayne aquifer: Stratiform “super-K” zones: Geologi-
cal Society of America Bulletin, v. 121, p. 164–180.
Droser, M. L., and D. J. Bottjer, 1986, A semiquantitative
field classification of ichnofabric: Journal of Sedimentary
Research, v. 56, p. 558–559.
Gingras, M. K., S. G. Pemberton, C. A. Mendoza, and F.
Henk, 1999, Assessing the anisotropic permeability of
Glossifungites surfaces: Petroleum Geoscience, v. 5,
p. 349–357, doi:10.1144/petgeo.5.4.349.
Gingras, M. K., B. MacMillan, B. J. Balcom, T. Saunders, and
S. G. Pemberton, 2002, Using magnetic resonance imag-
ing and petrographic techniques to understand the textur-
al attributes and porosity distribution of Macaronichnus-
burrowed sandstone: Journal of Sedimentary Research,
v. 72, p. 552–558, doi:10.1306/122901720552.
Gingras, M. K., C. A. Mendoza, and S. G. and Pemberton,
2004, Fossilized worm burrows influence the resource
quality of porous media: AAPG Bulletin, v. 88, p. 875–
883, doi:10.1306/01260403065.
556 Computer Modeling Bioturbation
Gingras, M. K., S. G. Pemberton, F. Henk, J. A. MacEachern,
C. A. Mendoza, B. Rostron, R. O’Hare, M. Spila, and
K. Konhauser, 2007, Applications of ichnology to fluid
and gas production in hydrocarbon reservoirs, in J. A.
MacEachern, K. L. Bann, M. K. Gingras, and S. G.
Pemberton, eds., Applied ichnology: SEPM Short Course
Notes 52, p. 129–143.
Katrak, G., and F. L. Bird, 2003, Comparative effects of the
large bioturbators, Trypaea australiensis and Heloecius
cordiformis, on intertidal sediments of Western Port, Vic-
toria, Australia: Marine and Freshwater Research, v. 54,
p. 701–708, doi:10.1071/MF03015.
Meadows, P. S., and J. Tait, 1989, Modification of sediment
permeability and shear strength by two burrowing inver-
tebrates: Marine Biology, v. 101, p. 75–82, doi:10.1007
/BF00393480.
Pemberton, S. G., and M. K. Gingras, 2005, Classification
and characterizations of biogenically enhanced perme-
ability: AAPG Bulletin, v. 89, p. 1493–1517, doi:10
.1306/07050504121.
Philip, Z. G., J. W. Jennings Jr., J. Olson, S. E. Laubach, and
J. Holder, 2005, Modeling coupled fracture-matrix fluid
flow in geomechanically simulated fracture networks: So-
ciety of Petroleum Engineers Reservoir Evaluation and
Engineering, v. 8, p. 300–309.
Stauffer, D., and A. Aharony, 1994, Introduction to percolation
theory: Revised 2d ed.: Philadelphia, Taylor and Francis,
192 p.
Taylor, A. M., and R. Goldring, 1993, Description and anal-
ysis of bioturbation and ichnofabric: Journal of the Geolog-
ical Society (London), v. 150, p. 141–148, doi:10.1144
/gsjgs.150.1.0141.
Tonkin, N. S., D. McIlroy, R. Meyer, and A. Moore-Turpin,
2010, Bioturbation influence on reservoir quality: A
case study from the Cretaceous Ben Nevis Formation,
Jeanne d’Arc Basin, offshore Newfoundland, Canada:
AAPG Bulletin, v. 94, p. 1059–1078, doi:10.1306
/12090909064.