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Eggshell taphonomy: environmental effects on fragment orientation

James L. Hayward
a
*, Kristin M. Dickson
b
, Susan R. Gamble
c
, Adam W. Owen
d
and Kimberly C. Owen
e{
a
Department of Biology, Andrews University, Berrien Springs, MI 49104, USA;
b
Emory University Psychiatry and Behavioral Sciences,
Tufts House, Suite 218, 2004 Ridgewood Dr, Atlanta, GA 30322, USA;
c
Department of Internal Medicine, Yale School of Medicine,
Residency Training Programs, P.O. Box 208030, New Haven, CT 06520, USA;
d
Central Maine Pain and Headache Center, 10 Minot
Avenue, Auburn, ME 04210, USA;
e
Central Maine Medical Center, 300 Main St., Lewiston, ME 04240, USA
(Received 27 May 2010; nal version received 3 June 2010)
Dinosaur eggshell is abundant in the fossil record but only during the last several decades has this reproductive product been
considered more than a novelty. Recent work has provided evidence that both whole eggs and fragmented eggshell represent a
rich source of information related to paleoecology and dinosaur reproductive biology. In this paper, we report the effects of
environmental variables on modern eggshell fragment orientation. Non-transported eggshell fragments at hatching and
predation sites favoured concave-up over concave-down orientations. Trampled fragments and fragments transported by
wind and water favoured concave-down over concave-up orientations. Although differences in orientation between non-
transported and transported eggshell fragments were usually obvious and signicant under the chosen experimental
conditions, paleontologists are cautioned to interpret the taphonomy at fossil eggshell sites with care and within their
sedimentological context.
Keywords: dinosaur; eggshell; experiments; taphonomy; transport
Introduction
Dinosaur eggshell is abundant in the fossil record but only
recently has this reproductive product been considered a
valuable source of information. Whole fossil eggs and egg
clutches are the most exciting discoveries of this type and
have yielded spectacular insights into the lives of the
animals that produced them (Carpenter et al. 1994;
Carpenter 1999). Recent experimental work has demon-
strated, however, that even fragmented eggshell contains a
wealth of paleoecological information.
Dinosaurs, like modern birds, laid eggs with brittle
calcitic shells. The shells of theropod dinosaurs consisted
of at least two structural layers (Mikhailov et al. 1996;
Mikhailov 1997; Zelenitsky et al. 2002). Some theropods,
like some birds, appeared to nest in colonies (Carpenter
1999), laid their eggs in rimmed nests located on the ground
(Horner 1982; Varricchio et al. 1997; Chiappe et al. 2004)
and apparently brooded or protected their eggs by covering
them with their bodies (Norell et al. 1995; Varricchio et al.
1997; Clark et al. 1999). Dinosaur eggshell was subject to
fragmentation by hatching and predation, transport by wind
and water and alteration by trampling and weathering just
like modern avian eggshell. Thus, taphonomic processes
affecting modern avian eggshell can serve as useful
analogues to processes inuencing ancient dinosaur
eggshell (Hayward et al. 2000).
The bowl-like shape of eggshell fragments, like that of
bivalve shells (Clifton and Boggs 1970), is subject to
physical forces that inuence resting orientation patterns.
An understanding of these patterns holds promise for
increasing our understanding of eggshell paleoenviron-
ments and transport processes (Hayward et al. 1997, 2000).
Here, we report a series of observations and experiments
designed to provide insight into what eggshell fragment
orientation concave surface up or down can provide
regarding taphonomic factors impinging on dinosaur
eggshell. Specically, we test the null hypothesis that
hatching, post-hatching trampling, predation and transport
in a wind tunnel, ume and natural stream lead to 1:1 ratios
of concave surface up and down orientations of eggshell
fragments.
Previous work
To place our study in context, we rst review the small
number of previously published studies in experimental
eggshell taphonomy.
ISSN 0891-2963 print/ISSN 1029-2381 online
q 2011 Taylor & Francis
DOI: 10.1080/08912963.2010.499170
http://www.informaworld.com
*Corresponding author. Email: hayward@andrews.edu

Email: kristin.dickson@gmail.com

Email: susan.gamble@yale.edu

Email: adamowen33@hotmail.com
{
Email: kimcso@hotmail.com
Historical Biology
Vol. 23, No. 1, March 2011, 513
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Early motivation for experimental eggshell taphonomy
was provided by the 18 May 1980 eruption of Mount St.
Helens, Washington. This event buried an entire nesting
colony of ring-billed and California gulls (Larus
delawarensis and L. californicus) 1 day before the onset
of hatching (Hayward et al. 1982). One year later, entire
clutches of desiccated ring-billed gull eggs were found
buried beneath the ash (Hayward et al. 1989). Structural
features of fresh eggshell and eggshell buried beneath the
ash for 1 year and 7 years were compared and the results
demonstrated rapid corrosion by the acidic ash (Hayward
et al. 1991).
Carpenter (1982) discussed the effect of local pHon the
preservation potential of dinosaur eggshell and found that
in 60 days acids produced by decaying vegetation
dissolved the broken edges of chicken eggshell fragments.
Clayburn et al. (2004) examined this effect further, along
with that of temperature, in controlled experiments with
glaucous-winged gull (L. glaucescens) eggshell fragments.
Treatment in buffered solutions of varying pH and
temperature for up to 42 days resulted in corrosion and
pitting of the outer surface and corrosion on the
mammillary layer of the inner surface. Fragment mass,
surface area and thickness decreased in response to
decreased pH and increased temperature and exposure
time. Similarly, Smith and Hayward (2010) inoculated
sterile glaucous-winged gull eggshell fragments in sterile
soil with bacteria isolated from eggshell on the surface of
the nesting colony. Over a 10-week period, eggshell protein
concentration declined, soil calcium concentration
increased and eggshell corrosion patterns similar to those
seen in naturally weathered eggshell appeared.
Horner (1994) reported a study of bone and eggshell
distributions at colonies of American white pelicans
(Pelecanus erythrorhynchos), double-crested cormorants
(Phalacrocorax auritus) and various species of gulls. He
noted that colony surfaces were littered with juvenile
bones and carcasses but that none of the remains were
from adults. Similarly, Hayward et al. (2000) analysed
bone, mollusc shell and eggshell fragment distributions on
the surface of a glaucous-winged gull (L. glaucescens)
colony. Approximately 4 weeks after the time of hatching,
eggshell fragment densities were signicantly higher close
to nests than further away, but this was not true for bones
and mollusc shells. Weathering of gull eggshells, placed
on the colony surface in an exclosure which prohibited
disturbance by colony residents, was followed for 2 years.
Within 1 week, shrinking membranes detached from the
inner surface of the shells and precipitated fragmentation.
After 1 year, some of the membranes were still present but
had changed colouration, as had the outer shell surfaces.
After 2 years, membranes and shell inner surfaces were
coated with unicellular green algae. Hatched and predated
eggshell could be distinguished on the basis of the fracture
patterns.
Soja et al. (2004, 2005) and Soja (2008) carried out
burial experiments with infertile ostrich, emu (Dromaius
novaehollandiae), chicken (Gallus gallus) and alligator
(Alligator mississippiensis) eggs. Extensive fractures
developed across the surface of many eggs, the shells of
which were held together by the inner shell membrane.
Sediment ltered into eggs with fractured shells and
replaced the soft contents after it had drained away. The
sediment prevented further collapse of the shell, thus
increasing its fossilisation potential. Holes and depressed
features in some buried specimens were similar to
holes interpreted as hatching windows in some dinosaur
eggs, suggesting the importance of caution when interpret-
ingthese openings infossil specimens. Scavengers removed
over two-thirds of the eggs at one of the burial sites.
The impact of high temperatures on eggshell, as might
have been experienced during wildres (Wolbach et al.
1988; Scott et al. 2000), was explored by Janssen and
Hayward (2006). Ostrich (Struthio camelus) and glaucous-
winged gull eggshell fragments were heated to temperatures
up to 8508C for various lengths of time. A series of dramatic
colour changes, similar to those reported for conodonts
(Epstein et al. 1977), began to occur at about 2008C.
Reverse curling of eggshell was observed at temperatures
above 6008C. Thermogravimetric analysis showed a
negligible decrease in mass below 2008C, attributable to
water loss, but sharp decreases in mass occurred at about
7108C for both types of eggshell. The remaining CaO
residue represented 55% of the original mass.
Tokaryk and Storer (1991) demonstrated that eggshell
could be transported great distances in simulated uvial
systems without signicant damage. Hayward et al. (1997)
showed that whole chicken eggs were readily recruited
into the sea by rising tides. Fresh eggs rolled along the
benthic surface; simulated incubated eggs, which were less
dense, oated out to sea (see also Schafer 1972). Eggs
placed on the subtidal benthic surface accumulated
epibiont loads and by 44 days were partially buried by
sediment. When lowered to a depth of 626 m in the Pacic
Ocean, fresh chicken eggs withstood hyperbaric pressures
of 62 atmospheres without cracking or breakage.
Methods
In each section below, eggshell fragment orientation
refers to whether the concave surfaces of horizontally
positioned fragments were facing up concave up, or facing
down, concave down. Following each treatment, most
fragments assumed an unambiguously horizontal position;
thus, only horizontally positioned fragments were counted.
Means are reported with standard deviations (^SD), and all
statistical tests were carried out at the 0.05 level of
signicance. Table 1 summarises the observations and
experiments reported, and Figure 1 illustrates the range of
eggshell fragment sizes used in the transport experiments.
J.L. Hayward et al. 6
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Fragment orientation at hatching sites
On 7 July 2004 on the Protection Island National Wildlife
Refuge, Jefferson County, Washington, orientations of
eggshell fragments in 12 glaucous-winged gull (L.
glaucescens) nest cups in which all chicks had hatched
were determined and compared against the null prediction
of a 1:1 ratio using a chi-square test.
Twelve fertile domestic chicken eggs were obtained
from Town Line Poultry Farm, Zeeland, Michigan. Eggs
(mean length 55.5 ^ 1.53 mm; mean width 43.0 ^
1.04 mm) were incubated at 398C in an incubator with an
automatic egg turner. Just prior to the estimated time of
hatching (21 days post-laying), each egg was singly placed
in the centre of a circular cardboard arena, 33 cm in
diameter, 28 cm tall and with a sand-covered base. As soon
as hatching was complete, the chick was removed from the
arena without disturbing the eggshell fragments, and
fragment orientations for all arenas were determined and
compared against the null prediction of a 1:1 ratio using a
chi-square test.
Fragment orientation at egg predation sites
On 7 July 2004, orientations of glaucous-winged gull
eggshell fragments were determined at four egg cannibal
(Hayward et al. 2000) territories on the Protection Island
colony. Stolen eggs at such territories were transported
there whole by the cannibals, and then broken and the
contents devoured. Likewise, on 2 July 2004, orientations
of glaucous-winged gull eggshell fragments were deter-
mined at 10 sites where bald eagles (Haliaeetus
leucocephalus) had preyed on glaucous-winged gull eggs
on the Protection Island colony. These sites occurred in the
immediate vicinity (,2 m) of the predated gull nests
where the eggs had been broken into and the contents
devoured. In each case, orientations were determined and
compared against the null prediction of a 1:1 ratio using a
chi-square test.
Fragment orientation after trampling by chicks
Four domestic chicks from the hatching experiment were
placed in a single arena, shaped as a half circle,
radius 50 cm, with a sand-covered base. One hundred
and twenty crushed chicken eggshell fragments of mixed
diameters were placed concave up on the surface of the
Figure 1. Range of chicken eggshell fragment sizes used in the
transport experiments. A United States 25-cent coin is shown for
scale. Dimensions refer to diameters.
Table 1. Summary of observations and experiments. Type of observation or experiment, total number of eggshell fragments counted in
the observation or included in the experiment and total number of fragments included in the data analysis for that observation or
experiment (i.e. fragments in an unambiguously horizontal position and therefore countable) are listed. Gull refers to glaucous-winged
gull (L. glaucescens) and Chicken and chick refer to domestic fowl (Gallus gallus).
Observation or experiment Fragments included Fragments counted
Hatching sites (gull and chicken eggshell)
Gull (12 nest sites) 145 145
Chicken (9 hatch sites) 220 220
Predation sites (gull eggshell)
Gull egg cannibal (4 sites) 629 629
Eagle predation on gull eggs (10 sites) 557 557
Chick trampling (chicken eggshell)
Initially concave up 120 99
Initially concave down 120 84
Wind tunnel transport (chicken eggshell)
Small fragments
Initially concave up 50 50
Initially concave down 50 50
Large fragments
Initially concave up 50 50
Initially concave down 50 50
Flume transport (chicken eggshell)
Initially concave up 30 30
Initially concave down 30 30
Natural stream transport (chicken eggshell)
Medium fragments (initial orientation random) 355 338
Half shells (initial orientation random) 50 23
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sandy base. Chicks were provided with food and water and
allowed to move freely about the arena for 36 h, after
which they were removed. After chick removal, the
orientations of fragments were determined. The exper-
iment was repeated, except that 120 eggshell fragments
were initially placed concave down. A McNemar test for
signicance of change (Zar 1999; Berenson et al. 2005)
was used to test the null prediction that fragments in both
experiments showed no change in orientation as a result of
treatment.
Fragment transport in a wind tunnel
Transport experiments were conducted using a wind
tunnel 5.7 mm long and 18 cm wide with 26 cm high sides.
The top of the tunnel was covered with glass and the oor
was covered with dry, levelled sand. A squirrel cage fan
served as a wind source. A bafe made of stacked plastic
tubes 15.5 cm long and 3.2 cm in diameter was placed
between the fan and the tunnel to minimise turbulence.
The tunnel was marked at 10-cm intervals. Wind velocity
was measured with a Turbo Meter
TM
electronic wind
speed indicator (Davis Instruments Corp.) which averaged
to 8.2 m/s at the proximal end and 3.8 m/s at the distal end
of the tunnel.
Fifty small (diameter < 2.0 cm) and 50 large
(diameter < 3.7 cm) eggshell fragments were placed
concave up 2 cm in front of the fan bafe, two to four
at a time to avoid fragment collisions. After smoothing
the sand with a straight edge, the fan was turned on and
run for 15 s, after which distance from the starting
point was determined. The experiment was repeated
with each fragment initially placed concave down.
McNemar tests were used to test the null prediction that
fragments showed no change in orientation as a result of
treatment.
Fragment transport in a laboratory ume
Chicken eggshell fragments, 2.55.0 cm in diameter,
were experimentally transported in a Hydraulic Demon-
stration Channel, Model A-8 (Engineering Laboratory
Design, Inc.). In this device, water stored in a reservoir
could be pumped through a ume with transparent sides,
2.20 m long, 0.15 m wide and 0.3 m deep. To minimise
turbulence, water entered the channel through a cross-
section of tubular bafes constructed from stacked 1.27-
diameter PVC pipes 10 cm long. A segment of the channel
with the least turbulence, 90 cm long, was selected as the
working distance for the experiment. The eggshell release
point was located at the proximal end of the working
distance. During each experiment, water depth and tank
slope were maintained at 20 cm and 08, respectively.
Experiments were conducted at two water velocities
determined at the release point as 25 and 30 cm/s using a
Global Flow Probe
TM
FP101-FP-201 (Global Water
Instrumentation, Inc.).
Thirty fragments each underwent two experimental
runs, one initially concave up and the other initially
concave down. The smooth plastic substrate of the ume
channel was made rough by completely covering it with an
asphalt shingle. Before release, each fragment was
submerged and placed entirely on the rough substrate
with its front edge at the release point. Final fragment
orientation was determined once the fragment reached the
end of the working distance. The shingle was then
removed and the experiment was repeated with only the
smooth substrate of the ume channel. McNemar tests
were used to test the null prediction that fragments in
both experiments showed no change in orientation as a
result of treatment.
Fragment transport in a natural stream
Pieces of crushed chicken eggshells were divided into two
categories: fragments .11 mm, n 355 and half shells,
n 50. On 12 September 1996, these samples were
released in mass beneath the waters surface in Lemon
Creek, Berrien County, Michigan, 0.5 km upstream from
where the creek enters the St. Joseph River. Water depth
ranged from 5 to 30 cm; ow rates (measured with a
Global Flow Probe
TM
) ranged from 15 to 45 cm/s.
The nal distributions of eggshell fragments and half
shells were determined at least 6 days later by
positioning a 1-m
2
steel grid with sixteen 0.0625-m
2
quadrants on the benthic surface at increasing distances
downstream from the release point until no eggshell was
observed. Eggshell fragments within each quadrant were
viewed through the bottom of an empty 1.5-litre beaker
pushed below the waters surface. Buried fragments were
exposed by gently disturbing the surface sediment.
Fragment and half shell orientations were determined and
compared against the null prediction of a 1:1 ratio using
chi-square tests.
Results
Fragment orientation at hatching sites
In the 12 glaucous-winged gull nests, 145 eggshell
fragments (

X 12.08 ^ 6.79 fragments/nest) were found,


signicantly more of which were oriented concave up
(n 95) than down (n 50; x
2
13.97, 1 d.f.,
p 0.0002; Figure 2(A)). Nine of the 12 domestic chicken
eggs hatched and yielded 220 eggshell fragments
(

X 24.44 ^ 9.03 fragments/hatched egg), signicantly


more of which were oriented concave up (n 137) than
down (n 83; x
2
13.25, 1 d.f., p 0.0003; Figure 2(B)).
Differences in orientation by fragment size were non-
signicant (x
2
4.97, 2 d.f., p . 0.05).
J.L. Hayward et al. 8
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Figure 2. Eggshell fragment orientations at/in (A) glaucous-winged gull nest cups; (B) domestic chick nest sites; (C) sites where
glaucous-winged gulls preyed on eggs of other gulls; (D) sites where bald eagles preyed on glaucous-winged gull eggs; (E) site of
dinosaur egg clutch at Devils Coulee, Alberta (from Hayward et al. 2000); (F) site of dinosaur eggshell near Augusta, Montana
(from Hayward et al. 1997); (G) domestic chick arena; (H) wind tunnel (small fragments); (I) wind tunnel (large fragments); (J)
ume, rough substrate, 25 cm/s; (K) ume, rough substrate, 30 cm/s; (L) ume, smooth substrate, 25 cm/s; (M) ume, smooth
substrate, 30 cm/s; (N) natural stream (fragments); (O) natural stream (half shells) and (P) experimental turbidites (from Hayward
et al. 1997).
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Fragment orientation at egg predation sites
At the four glaucous-winged gull egg cannibal sites, 629
eggshell fragments were randomly sampled, signicantly
more of which were oriented concave up (n 435) than
down (n 194; x
2
92.3, 1 d.f., p , 0.00001; Figure
2(C)). At the 10 bald eagle predation sites, 557 eggshell
fragments were randomly sampled, signicantly more of
which were oriented concave up (n 401) than down
(n 156; x
2
107.8, 1 d.f., p , 0.00001; Figure 2(D)).
Fragment orientation after trampling by chicks
Of 120 fragments placed concave up, 117 were located
after 36 h exposure to trampling by chicks, 99 of which
could be classied by orientation; of these classied
fragments, 57 were concave up and 42 down. Of 120
fragments placed concave down, 114 were located after
36 h, 84 of which could be classied by orientation; of
these classied fragments, 19 were concave up and 65
down. Thus, signicantly more fragments retained or
assumed a concave-down orientation after exposure to
trampling (n 107) than before (n 84; x
2
7.93, 1
d.f., p 0.0049; Figure 2(G)).
Fragment transport in a wind tunnel
After 15 s in a wind tunnel, small eggshell fragments
initially positioned concave up travelled a mean
distance of 1.07 ^ 0.534 m, whereas the same fragments
initially positioned concave down travelled a mean
distance of 0.26 ^ 0.442 m (paired-t 9.42, d.f. 49,
p , 0.0000001; Figure 3). Similarly, large eggshell
fragments initially positioned concave up travelled a
mean distance of 2.32 ^ 0.763 m, whereas the same
fragments initially positioned concave down travelled a
mean distance of 0.334 ^ 0.716 m (paired-t 9.42,
d.f. 49, p , 0.0000001; Figure 3). For small fragments,
the number of fragments oriented down following wind
treatment (n 75) was signicantly higher than the
number of fragments oriented down before (n 50;
x
2
16.46, 1 d.f., p 0.00005; Figure 2(H)); the same
was true for large fragments following treatment (n 81)
compared with those oriented down before (n 50;
x
2
27.27, 1 d.f., p , 0.000001; Figure 2(I)).
Fragment transport in a laboratory ume
Final orientations of eggshell fragments transported in a
laboratory ume varied by initial orientation and substrate
type; a slightly higher frequency of fragment orientation
change occurred at the higher velocity of water ow on both
smooth and rough substrates. On the rough substrate, the
number of fragments oriented concave down following
treatment at the 25-cm/s owrate (n 51) was signicantly
higher than the number of fragments oriented concave down
before (n 30; x
2
19.05, 1 d.f., p 0.00001; Figure
2(J)); the same was true for fragments following treatment at
the 30-cm/s ow rate (n 53) in comparison with those
oriented concave down before (n 30; x
2
19.17, 1 d.f.,
p 0.00001; Figure 2(K)). On the smooth substrate, the
number of fragments oriented concave down following
treatment at the 25-cm/s ow rate (n 37) was not
signicantly higher than the number of fragments oriented
concave down before (n 30; x
2
3.27, 1 d.f., p 0.071;
Figure 2(L)); the same was true for fragments following
treatment at the 30-cm/s ow rate (n 34) in comparison
with those oriented concave down before (n 30;
x
2
0.45, 1 d.f., p 0.502; Figure 2(M)). Notably, no
fragments transported at either of the water velocities on the
rough surface changed orientation fromconcave down to up
over the entire working distance.
Fragment transport in a natural stream
Of the 355 fragments released into Lemon Creek, 340
fragments, all within 4 m of the release point
Figure 3. Distances travelled by small and large chicken eggshell fragments in a wind tunnel.
J.L. Hayward et al. 10
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(

X 1.2 ^ 0.81 m), were recovered. Of these recovered


fragments, 338 were oriented horizontally: 50 were
concave up and 288 down (x
2
167.6, 1 d.f.,
p , 0.0000001; Figure 2(N)). Of the 50 half shells
released, 38 were recovered up to a distance of 302 m
(

X 55.4 ^ 98.8 m). Unlike fragments in this and other


experiments, most recovered half shells were oriented
concave up (n 16) or sideways (n 15); only seven half
shells were oriented concave down. The comparison
between the number of concave up half shells to that of
concave down half shells was non-signicant (x
2
3.52,
1 d.f., p . 0.05; Figure 2(O)).
Discussion
Eggshell fragmentation occurred at hatch and predation
sites with little or no subsequent transport. Concave-up
orientations were favoured at these sites. Egg surface
curvature ensures that the most likely orientation for an
eggshell fragment to assume when it reaches the ground is
concave up. Only fragments that reach the ground concave
down are likely to retain that orientation without the
impact of a horizontal force. When a fragment reaches the
ground in an unstable vertical orientation, its centre of
gravity tends to pull it down into a stable concave-up
horizontal orientation. Fragments that reach to the ground
concave up following hatching or predation likely will
retain that orientation (Figure 4(A)).
Trampled fragments were more likely to assume a
concave-down orientation than a concave-up orientation.
The raised edges of concave-up fragments were easily
kicked by chicks walking over the substrate or scratching
for food, likely explaining why 42% of fragments initially
placed concave up were turned over by chick activity. By
contrast, only 23% of fragments initially placed concave
surface down were turned over; chick activity was less
likely to overturn these fragments because their edges
were resting on or pressed into the sand.
The most stable orientation for an eggshell fragment
subjected to horizontal uid ow is concave down,
because this orientation affords a streamlined surface over
which ow occurs (Figure 4(B)). By contrast, horizontal
uid ow tends to overturn concave-up fragments, thus
bringing them to a more stable orientation (Figure 4(C)).
Fluid ow encounters larger and more complicated shapes
in half eggshells; thus the preference for a concave-down
orientation by eggshell fragments subjected to this type of
force is not typical for half shells.
Large fragments travelled further than small fragments
in the wind tunnel experiment, and half eggshells travelled
farther than smaller fragments in the natural stream
experiment. The greater surface area of the larger
specimens allowed them to be pushed along more readily
by current ow. The ume experiments demonstrated the
impact of substrate texture on fragment travel. Fragments
were more likely to assume a concave-down orientation
than a concave-up orientation on both rough and smooth
substrates. The proportion of fragments oriented concave
down, however, was higher when transported over rough
substrate. On the smooth substrate, fragments were more
likely to slide along the surface, without turning over.
Thus, the results of transport experiments may differ
depending on the rate of ow and substrate type, and on an
interaction of the size, shape and density of the transported
objects (Clifton and Boggs 1970).
To summarise, hatching and predation sites favoured
concave-up over concave-down fragment orientations.
Trampling and transport by wind and water favoured
concave-down over concave-up orientations. Previous
studies showed that dinosaur eggshell fragments at two
sites were more commonly oriented concave up than down
(Hayward et al. 1997, 2000; Figure 2(E) and (F)). This
nding was consistent with the hypothesis that these sites
represent nesting or predation sites that experienced little
or no trampling or transport prior to burial.
Our ndings comport with those of Emery (1968) who
found that empty bivalve shells favour a concave-up
orientation in quiet water found in deep areas of the
continental shelf, whereas concave-down orientation is
favoured in wave wash zones found in shallow water close
to shore. In experiments reported by Hayward et al.
(1997), however, more fragments transported in simulated
Figure 4. Forces acting on eggshell fragments. (A) When a
fragment reaches the ground in an unstable vertical orientation,
its centre of gravity tends to pull it down into a stable concave-up
horizontal orientation; (B) fragments oriented concave surface
down tend to retain that stable orientation in uid (air or water)
ow and (C) fragments oriented concave surface up are unstable
in uid ow and tend to ip over to a stable concave surface
down orientation.
Historical Biology 11
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turbidity currents settled in a concave-up (n 141)
orientation than a concave-down (n 125) orientation,
although this difference was not signicant (Figure 2(P)).
This result is similar to descriptions by Compton (1962)
and Crowell et al. (1966) of marine shell assemblages in
turbidite deposits, and it is consistent with results obtained
by Allen (1984) with bivalve shells in experimental
turbidites. Clifton and Boggs (1970) noted that a
predominantly concave-up orientation can also occur as
a result of wave action in the uppermost swash zone,
deposition in small sediment traps and migration of small
subaqueous sand ripples. Thus, not all transported
eggshells can be expected to exhibit predominantly
concave-down orientations. Features of the surrounding
sedimentary matrix must be examined in concert with
eggshell taphonomic features when interpreting fossil
eggshell deposits as either transported or non-transported.
In situ and post-transport fragment orientation occurs
within a larger taphonomic context that extends temporally
from egg laying to post-burial (Figure 5). Once laid, a bird
egg is either hatched, transported whole, is broken by
predators or is buried by sedimentation processes. Hatched,
transported and predated eggshell may be buried, broken
into smaller fragments before either further transport or
burial, or dissolve away before burial. Once buried, whole
eggs or nearly complete portions typically fracture, ll with
sediment and often are compressed due to sediment
loading. Sediment pH, Eh and moisture affect the quality
and length of preservation. Each step of this complicated
process deserves further evaluation by paleoecologists and
biogeochemists in the context of the myriad physical,
chemical and biological interactions involved. Some
dinosaur eggs, it should be noted, may have been buried
within soil or vegetation (Cousin et al. 1994; Carpenter
1999) and thus experienced somewhat different tapho-
nomic histories than most modern bird eggs.
Experimental eggshell taphonomy is still in its infancy,
andonlya handful of researchers have done workinthis area.
Transport and burial experiments with a wide variety of eggs
and eggshell fragment sizes, masses and shapes should be
undertaken; correlations between eggshell taphonomic
signatures and the reproductive behaviours of parents and
offspring should be examined and comparisons between
experimental results and eggshell fossil sites should be
made. Eggshell taphonomy provides a broad opportunity for
undergraduates, along with graduate students and senior
researchers, to make signicant contributions to our
understanding of paleoecology and dinosaur reproductive
biology.
Acknowledgements
We are grateful for the encouragement to study eggshell
taphonomy by our late friend Karl F. Hirsch (19211996). We
thank David Varricchio and Frankie Jackson for their helpful
critiques of an earlier draft of this paper and, who along with Jack
Horner, hosted the Fourth International Symposium on Dinosaur
Eggs and Babies where this paper was presented. Research funds
were provided by Andrews University Faculty Grants program,
Andrews University Department of Biology, and the McGregor
Figure 5. Major taphonomic processes (arrows) that inuence eggs and eggshell prior to fossilisation.
J.L. Hayward et al. 12
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Foundation. Logistic support was provided by Rosario Beach
Marine Laboratory, Anacortes, Washington. The last four authors
participated in this research as undergraduate members of the
Andrews University Honors Program.
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