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 SPECIAL PUBLICATION OF THE PALAEONTOLOGICAL SOCIETY OF INDIA
No. 5; February, 2014; ISBN: 978-81-926033-2-2; pp. 205-213

PALAEOBIOGEOGRAPHIC CONSTRAINTS ON DRILLING

GASTROPOD PREDATION: A CASE STUDY FROM THE MIOCENE
KHARI NADI FORMATION IN KUTCH, GUJARAT
SUBHENDU BARDHAN1*, SUMANTA MALLICK1 and SHILADRI S. DAS2
1
DEPARTMENT OF GEOLOGICAL SCIENCES, JADAVPUR UNIVERSITY. KOLKATA – 700 032,
2
GEOLOGICAL STUDIES UNIT, INDIAN STATISTICAL INSTITUTE, 203 BARRACKPORE TRUNK
ROAD, KOLKATA 700 035, INDIA
*
Corresponding author’s e-mail: sbardhan12@gmail.com

ABSTRACT
Predator-prey interaction is an important aspect of biological system. Gastropod drilling predation can be traced in the fossil
record and provide many reliable information about drilling activities e.g., drilling intensity, efficiency of predators through size and site
stereotypy.
Drilling frequency (DF) varies widely in space and time. Temporal changes, although complex, depicts a pattern. Drilling frequency
was low during the Cretaceous and cyclically increased during the Eocene and Miocene. We, however have found low drilling frequencies
on molluscs in the Early Miocene (Aquitanian) Khari Nadi Formation in Kutch, Gujarat.
India experienced varying degree of isolation during the Oligocene – Miocene times and was completely cut off from the western
Tethys during the Aquitanian. Escalated predators from Europe and other provinces were barred from invading the Indian waters which
resulted in significant lowering of the drilling frequencies on molluscan assemblages.
Keywords: Drilling predation, Kutch, Miocene, palaeobiogeographic constraints

INTRODUCTION Palaeozoic drillholes were caused by many

Predator–prey interaction has a long geological
history since the Neoproterozoic times (Bengston and
Zhao, 1992; Hua et al., 2003). However, many such
interactions, e.g., swallowing the prey whole or crushing
of prey shells, hardly leave any readily identifiable
features in the fossil record. Drilling predation is one
such interaction, which has preservable fossil record and
plays significant role in predator – prey system which
resulted in evolutionary consequences like co-evolution
which describes cases where two (or more) species
reciprocally affect each other’s evolution and escalation
which can be defined as changes occurring within any
species due to the changes taking place within their
enemies (Vermeij, 1987; Dietl and Alexander, 2000).
Intensity of predation varies with time. Mesozoic Era is
known as the time of major change in the nature of life
on Earth. One of the major aspects of this change was
the rise of many predatory groups. During the Middle
to Late Mesozoic the predation pressure reached a peak,
which is aptly called as Mesozoic marine revolution
(Vermeij, 1977).
predatory/parasitic groups (e.g., see Fürsich and
Jablonski, 1984; Baumiller, 1990, 1996; Leighton, 2001,
2003), while Mesozoic drillholes may have been made
by some unknown gastropod predators (e.g., Newton,
1983; Smith et al., 1985; Newton et al., 1987; Harper et
al., 1998, Harper, 2003; Heidelberger and Amler, 2002;
Bardhan et al., 2012). Reports of unambiguous
gastropod drilling predation came from the late Early
Cretaceous, when two predatory gastropod families, the
Muricidae and Naticidae, first appeared in the fossil
record (Sohl, 1969; Taylor et al., 1983). Tertiary rock
record is full of holes.
Drillhole morphology, (e.g., shape and size of
drillholes) provide many insights about the identity,
behaviour, and size of the driller. Generally, naticids
make tapered holes (‘truncated paraboloid’, see Carriker
and Yochelson, 1968), whereas muricid drillholes are
‘straight – walled’ (but see Herbert and Dietl, 2002).
Predatory drillholes provide many reliable information
about various significant aspects of predation, including
predation intensity, prey size, predator size, and
206
behaviour of predators (for reviews and references, see
Kitchell et al., 1981; Vermeij, 1983, 1987; Kitchell,
1986; Kabat, 1990; Kowalewski, 2002; Kowalewski and
Kelley, 2002; Leighton, 2002; Kelley et al., 2003).
Naticid outer drillhole diameter is well correlated with
predator size (Kitchell et al., 1981).
Gastropod drilling predation shows
spatiotemporal variation. In general, for any time plane,
drilling intensity increases towards the equator (Vermeij,
1987). Temporally the pattern is complex and probably
cyclic (Kelley and Hansen, 1993, 1996, 2006). All
evidences indicate that the Miocene drilling frequencies
on molluscan assemblages reached a new high. We here,
have reported drilling frequencies on some bivalve and
gastropod species systematically collected from the
Khari Nadi Formation (Aquitanian) of Kutch, Gujarat.
The DF values are significantly low in comparison to
the global data. We have explored the reasons and
considered the palaeobiogeographic constraints might
have played a role.
MATERIAL AND METHODS
The present fossil specimens have been
collected from the rocks of two localities (Figure 1),
which belong to the Khari Nadi Formation of the Early
Miocene (Biswas, 1992). This formation is dominantly
arenaceous and overlies the rocks of the Maniyara Fort
Formation (Figure 2). The thickness of the formation
varies considerably and in the type section it attains a
thickness of 65 meters. Biswas (1992) interpreted the
depositional environment in littoral to shallow inner
shelf areas.
Fig. 1. Map showing the fossil bearing localities (+) within the
Khari Nadi Formation.
SUBHENDU BARDHAN and OTHERS
Fossils are present in var ying mode of
occurrences. In one section on the river bed of the Khari
Nadi fossils are strongly enclosed in a coarse sandstone
bed (Figure 3A) and were very hard to retrieve. We
therefore, collected specimens following grid sampling
protocol of Bardhan et al. (2012). We have made several
grids of 4 square meter each. All the samples within
these grids were photographed and if possible collected
after detailed study. Besides, we have studied drilled
samples outside the grid. All bivalve samples belonged
predominantly to two species, Chlamys sp. and Ostrea
sp. (Figure 3B, C).
Most specimens are disarticulated valves and
show little taphonomic influence. We have analysed both
complete and broken shell separately. In another section
where the fossils are overwhelmingly rich in gastropods
especially turritelline species (Figure 4), the sediment
is particularly friable and fossils are easy to retrieve.
We therefor e employed bulk sampling protocol
following Kowalewski (2002) and Mallick et al. (2013).
Fig. 2. Stratigraphic position of the studied Khari Nadi Formation
(modified after Biswas, 1992).
PALAEOBIOGEOGRAPHIC CONSTRAINTS ON DRILLING GASTROPOD PREDATION: A CASE
STUDY FROM THE MIOCENE KHARI NADI FORMATION IN KUTCH, GUJARAT
We have studied drillhole morphology in detail
and found that drill holes are different in different
taxonomic groups. While the bivalves bears
characteristic cylindrical drill holes (Oichnus simplex
Bromley, 1981) allegedly made by muricids, majority
of turritelline gastropods bear naticids drill holes ‘which
are truncated spher ical paraboloid’ (Oichnus
paraboloides Bromley, 1981; Carriker, 1981; Carriker
and Yochelson, 1968). The different prey preference of
muricids and naticids may be explained by the facts that
Fig. 3. A. Coarse sandstone of the Khari Nadi Formation exposed along the Khari Nadi river near Lakshimpur, B-C. Drillholes on the shells
of Chlamys sp. and Ostrea sp. respectively.
For bivalves frequency of drilling (DF)
predation is calculated by dividing number of bored
valves by total number of individuals in the collection
(equation one of Bambach and Kowalewski, 2000).
i.e., N = [(RV + LV) / 2] + A
where LV, RV, A and N are number of left, right,
articulated valves and total number of individuals
respectively.
So, the drilling frequency (DF) is:
DF = DV/N
207
the present bivalves are epifaunal (Chlamys sp. is
epibyssated while Ostrea sp. is cemented in life mode).
In other examples from modern as well as fossil case
studies it is well known that muricids target epifaunal
prey (Carriker, 1981). In contrast, the gastropods
especially turritellines bear typical naticid drill holes.
Naticids are known to target infaunal prey (Sawyer and
Zuschin, 2010) and turritellines are dominantly infaunal
(see Paul et al., 2013 and references therein).
where DV is the number of drilled valves.
For gastropods, drilling frequency (DF) is expressed as
the percentage of shells with complete drillholes divided
by the total number of shells (Allmon et al., 1990).
So, the drilling frequency (DF) is:
DF = DV/N
where DV is the number of drilled individuals and N
represents total number of samples.
208 SUBHENDU BARDHAN and OTHERS

Frequencies of both incomplete and multiple
drillholes were measured. Unsuccessful drillholes are
an indication of prey effectiveness (PE), which has been
defined as a ratio between total incomplete drillholes
and total number of attempted holes (see Vermeij, 1987
and Kelley et al., 2001). Multiple drillhole frequency is
defined as the number of drillholes in multiply drilled
specimens divided by the total number of attempted
holes (Vermeij, 1987 and Kelley et al., 2001). According
to Kelley et al. (2001) both incomplete and multiple
drillholes indicate relative efficiency of predator and
prey. Incomplete drillholes indicate failure of the attack
and multiple holes indicate repeated attacks. If prey are
more vulnerable, then the value of incomplete and
multiple drillhole frequencies should decrease.

Fig. 4. Three species of turritelline gastropod from the Khari Nadi Formation. A-B. Zaria angulata, abapertural
and apertural views respectively, C-D. Turritella assimilis, apertural and abapertural views respectively, E-G.
Haustator tauroperrites, abapertural and apertural views respectively, G. broken specimen. Note, complete gastropod
drillholes in A, D and G (see arrows).

RESULTS DF is 5.77% (N=1472; drilled samples=85). In species

For Chlamys sp. the number of disarticulated
valves is 202, which are mostly dominated by left valves.
We have found only six articulated specimens of
Chlamys sp. Therefore DF (following the above
equation) is 7.48% (Drilled shell=8). Similarly in Ostrea
sp. we found 168 disarticulated valves and DF is 5.95%
(Drilled shell=5).
In turritelline gastropods, the subfamily level
level, the data range from 3.24% in Haustator
tauroperturrites (N=185) and 8.04% in Turritella
assimilis (N=373).
No incomplete or multiple drillholes were found
in the bivalves, however, relatively high value of
incomplete drilling frequency (PE=18.25%; N=126) and
multiple drilling frequency (PE=14.29%; N=126) were
recorded for the turritelline gastropods.
PALAEOBIOGEOGRAPHIC CONSTRAINTS ON DRILLING GASTROPOD PREDATION: A CASE 209
STUDY FROM THE MIOCENE KHARI NADI FORMATION IN KUTCH, GUJARAT

Table 1: Miocene drilling predation data on turritelline gastropods from all over the world (N=total number of shells examined;
D=number of drilled shells; UD=number of undrilled shells; DF=drilling frequency).

Formation/Place Age Turritellidae gastropod Source

N D UD DF%

Bulgaria Miocene 2642 1012 1630 38.3 Kojumdjieva, 1974

Poland Miocene 1461 270 1191 18.48 Hoffman et al., 1974
Maryland Miocene 101 28 73 27.72 Dudley & Vermeij, 1978
Venezuela Miocene 35 3 32 9 Dudley & Vermeij, 1978
Maryland Miocene 416 87 329 21 Kelley, 1982
Venezuela Miocene 73 12 61 16.44 Allmon et al., 1990
Maryland Miocene 63 14 49 22.22 Allmon et al., 1990
Florida Miocene 239 52 187 21.76 Hagadorn & Boyajian, 1997
Maryland Miocene 4186 323 3863 7.716 Hagadorn & Boyajian, 1997
Virginia Miocene 113 2 111 1.77 Hagadorn & Boyajian, 1997
Calvert Miocene 254 60 194 23.6 Kelley & Hansen, 2006
Choptank Miocene 1581 446 1135 28.2 Kelley & Hansen, 2006
St. Marys Miocene 2172 745 1427 34.3 Kelley & Hansen, 2006
Eastover Miocene 7 2 5 28.6 Kelley & Hansen, 2006
Panama Late Middle Miocene 370 162 208 43.78 Fortunato, 2007
Central Paratethys Lower and Middle Miocene 1757 109 1648 6.2 Sawyer & Zuschin, 2011

DISCUSSION showed low values of predation intensity. High DF

Gastropod predation on molluscan assemblages
has been now well studied and the temporal pattern has
been established. For example, Kelley and Hansen
(1993, 1996, 2006) described a cyclicity of drilling
intensity for both bivalves and gastropods. For
gastropods they reported low DF during the Cretaceous
and rapid rise in the aftermath of K-T extinction which
continued up to the late Eocene; then there was a decline
during the Oligocene in the DF followed by rapid rise
again during the Miocene. This pattern was valid for
various taxonomic ranks ranging from assemblage to
the level of species. For example, in turritelline
assemblage the Miocene drilling frequencies recorded
from other parts of the world were shown in Table 1.
Most of the DFs are quite high in comparison to Kutch
turritellines. Similar Miocene highs were also observed
in bivalve assemblages (Table 2). Average DF on
bivalves was again significantly higher than the Kutch
bivalve species (chi-square test of independence, 2 =
39.109, df = 1, p<0.01). Bardhan et al. (2012) recorded
high frequency of drilling (30.53%) from a Jurassic
bivalve (Neocrassina subdepressa) of Kutch. Kelley and
Hansen (2006) showed fluctuating values of drilling
predation from Cretaceous onwards. Except Brightseat
Formation (32.7%) all other Palaeocene formations
values were recorded again in the Oligocene and
Miocene, while in the Plio-Pleistocene time predation
rate again droped.
Why was Kutch Miocene drilling intensity so
low in comparison to other areas? Most of Miocene
(Kelley and Hansen, 1993, 1996, 2006; Hoffman et al.,
1974; Hoffmeister and Kowalewski, 2001; Sawyer &
Zuschin, 2011) data were from higher latitudes
especially from western Europe and North America.
Kutch belonged to the northern subtropic during the
Miocene (see Smith et al., 1981). In many instances,
predation intensity had latitudinal gradients where DFs
increased with decreasing latitudes (Vermeij, 1987; Paul
et al., 2013). The temporal increase in DF and especially
high values during the Miocene support the hypothesis
of escalation (Vermeij, 1987). It suggests that predators
learned from experience and became progressively more
efficient in handling the prey. The low DF in Kutch
turritelline gastropods may be explained by the fact that
prey were rather escalated and adapted well to resist
drilling predation and this was reflected in relatively
high value of incomplete drilling frequency
(PE=18.25%; N=126) and multiple drilling frequency
(PE=14.29%; N=126). For bivalves, however, no
incomplete or multiple drillhole was found.
210 SUBHENDU BARDHAN and OTHERS

Table 2: Miocene drilling predation data on bivalve assemblages from all parts of the world (N=total number of shells examined;
D=number of drilled shells; UD=number of undrilled shells; DF=drilling frequency).

Formation/Place Age Bivalve assemblage Source

N D UD DF (%)

Central Europe (Boreal) Miocene 563 92 471 16.3 Hoffmeister & Kowalewski, 2001
Central Europe (Atlantic) Miocene 47.5 5 42.5 10.5 Hoffmeister & Kowalewski, 2001
Central Europe (Paratethys) Miocene 425 72 353 16.9 Hoffmeister & Kowalewski, 2001
Calvert Miocene 1758 614 1144 34.9 Kelley & Hansen, 2006
Choptank Miocene 6755 2722 4033 40.3 Kelley & Hansen, 2006
St. Marys Miocene 8307 2367 5940 28.5 Kelley & Hansen, 2006
Eastover Miocene 4987 1730 3257 34.7 Kelley & Hansen, 2006
Central Paratethys Lr. & MiddleMiocene 8473 726 7747 8.6 Sawyer & Zuschin, 2011
Kutch, India Miocene (Burdigalian) 137.5 27 110.5 19.64 Chattopadhyay & Dutta, 2013

The other explanation for the low DF on the
Miocene molluscs of Kutch may be
palaeobiogeographic constraints. The Miocene Epoch
was an eventful time interval during which much of the
configuration and topography of the modern world
began to take shape (Theodor, 2005). The drifting and
reassembly of continents triggered a sudden increase of
biodiversity both in marine and terrestrial ecosystems.
Faunal diversification and modernity especially in
marine taxa also took place during that time (Theodor,
2005; Mondal et al., 2009). Miocene also experienced
global marine transgression and many parts of several
continents were flooded (Vail et al., 1977). Miocene
rocks are found in many coastal parts of the world
including India (see Sahni and Mehrotra, 1981; Mondal
et al., 2009) and from adjoining areas (see Vredenburg,
1928). Harzhauser et al (2009) studied Oligocene,
Miocene gastropod assemblages of Kutch. They found
that faunal similarities between Kutch and western
Tethys during Oligocene were good indicating the
existence of a Tethyan sea way for shallow water
molluscs. But there was a drastic decrease of similarities
between western Tethys and Indian faunas during the
Early Miocene time. This was also noticed by
Vredenburg in 1928 and others (Harzhauser et al., 2002,
2009; Martin, 1931; Piccoli et al., 1991). Mondal et al.
(2009) while studying Miocene shark teeth also came
to the same conclusion about the final closure of Tethyan
connection with India during the Miocene.
They observed poor faunal similarities among
India, Portugal (Western Tethys), North and South
America. This may be explained by the “final closure
of the eastern strait located in the Middle East, and
separating the proto-Mediterranean (Western Tethys)
and the new Indo-Pacific ocean (Eastern Tethys) during
the late-Early Miocene (Adnet et al., 2007, Popov et
al., 1993; Rögl, 1998, 1999; Adams et al., 1999)”.
As we have indicated that high DFs during the
Miocene were restricted mainly in the western Tethys
and predators particularly naticids were highly efficient
as evident from other criteria like size and site stereotypy
(Anderson, 1992; Hagadorn and Boyajian, 1997; Kelley,
1988). These highly escalated predators were widely
distributed but could not enter the Indian waters during
the Miocene due to reasons stated above. This
biogeographic constraints may be responsible for the
low DF in Indian molluscan assemblage. This is also
supported by the fact that the only naticid species, Natica
obscura (Harzhauser et al., 2009; fig. 2 q, r) found in
the Miocene beds is endemic to Kutch. We here did not
consider Globularia and Ampullinopsis as naticids (for
detailed discussion see Bardhan et al., 2012). Only
muricid genus reported so far is Athleta (Volutospina)
dentata (Harzhauser et al., 2009; fig. 5 d, e). Again the
species is endemic. We conclude that biological
interaction is also constrained by physical environmental
changes and evolutionary trends issued from biological
system may be influenced or affected by changes in the
physical system.
Our hypothesis that loss of faunal exchange
between Western Tethys and Indian waters decreased
the DFs on the Early Miocene Kutch molluscan
assemblages was further supported by the fact that in
the next Burdigalian Age, sea routes were further
established between Kutch and Western Tethys and there
was an increase in faunal similarities (Harzhauser, 2007;
Harzhauser et al., 2009). Consequently there was an
increase in drilling frequency on molluscs especially
PALAEOBIOGEOGRAPHIC CONSTRAINTS ON DRILLING GASTROPOD PREDATION: A CASE
STUDY FROM THE MIOCENE KHARI NADI FORMATION IN KUTCH, GUJARAT
bivalves gastropods in the subsequent Chhasra
Formation. Chattopadhyay and Dutta (2013) recorded
19.64% DF in bivalve assemblage and in particular, 13%
in Chlamys and 21.05% in Ostrea.
ACKNOWLEDGEMENTS
Thanks are due to the local people of
Lakshimpur and Bhadra, Gujarat and P. Roy, Durgapur
Government College, Durgapur and R. Dutta, Jadavpur
University, for help in the field, S. Bauri and S.
Chowdhury for help in the laboratory. The first author
(S.B.) acknowledges the partial funds provided by DST
(Department of Science and Technology) and CAS
(Centre of Advanced Study), Department of Geological
Sciences; UPE-II (University with Potential for
Excellence) and DST-PURSE (Department of Science
and Technology - Promotion of University Research and
Scientific Excellence) provided by central project,
Jadavpur University. S.M. also acknowledges the grant
for fieldwork funded by UGC (University Grants
Commission). S.S.D. also acknowledges Indian
Statistical Institute, Kolkata, for providing a grant to
pursue the research work.
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