Accepted Manuscript
Predation in the marine fossil record: Studies, data, recognition,
environmental factors, and behavior
Adiël A. Klompmaker, Patricia H. Kelley, Devapriya
Chattopadhyay, Jeff C. Clements, John W. Huntley, Michal
Kowalewski
PII: S0012-8252(18)30504-X
DOI: https://doi.org/10.1016/j.earscirev.2019.02.020
Reference: EARTH 2803
To appear in: Earth-Science Reviews
Received date: 30 August 2018
Revised date: 17 February 2019
Accepted date: 18 February 2019
Please cite this article as: A.A. Klompmaker, P.H. Kelley, D. Chattopadhyay, et al.,
Predation in the marine fossil record: Studies, data, recognition, environmental factors,
and behavior, Earth-Science Reviews, https://doi.org/10.1016/j.earscirev.2019.02.020
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Predation in the marine fossil record: studies, data, recognition, environmental factors, and
behavior
Adiël A. Klompmaker
a,
*, Patricia H. Kelley
b
, Devapriya Chattopadhyay
c
, Jeff C. Clements
d,e
,
John W. Huntley
f
, Michal Kowalewski
g
a
Department of Integrative Biology & Museum of Paleontology, University of California,
Berkeley, 1005 Valley Life Sciences Building #3140, Berkeley, CA 94720,
USA
b
Department of Earth and Ocean Sciences, University of North Carolina Wilmington,
Wilmington, NC 28403-5944, USA
c
Department of Earth Sciences, Indian Institute of Science Education and Research (IISER)
Kolkata, Mohanpur WB-741246, India
d
Department of Biology, Norwegian University of Science and Technology, 7491 Trondheim,
Norway
e
Department of Biological and Environmental Sciences, University of Gothenburg, Sven Lovén
Centre for Marine Sciences – Kristineberg, Fiskebäckskil 45178, Sweden
f
Department of Geological Sciences, University of Missouri, 101 Geology Building, Columbia,
MO 65211, USA
g
Florida Museum of Natural History, University of Florida, 1659 Museum Road, Gainesville, FL
32611, USA
*
Corresponding author. E-mail address: adielklompmaker@gmail.com (A.A. Klompmaker)
ABSTRACT
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The fossil record is the primary source of data used to study predator-prey interactions in deep
time and to evaluate key questions regarding the evolutionary and ecological importance of
predation. Here, we review the types of paleontological data used to infer predation in the marine
fossil record, discuss strengths and limitations of paleontological lines of evidence used to
recognize and evaluate predatory activity, assess the influence of environmental gradients on
predation patterns, and review fossil evidence for predator behavior and prey defense. We
assembled a predation database from the literature that documents a steady increase in the
number of papers on predation since the 1960s. These studies have become increasingly
quantitative and have expanded in focus from reporting cases of predation documented by fossils
to using the fossil record of predation to test ecological and evolutionary hypotheses. The data on
the fossil record of predation amassed so far in the literature primarily come from trace fossils,
mostly drill holes and, to a lesser extent, repair scars, derived predominantly from the Cenozoic
of Europe and North America. Mollusks are the clade most often studied as prey and inferred
predators. We discuss how to distinguish biotic from abiotic damage and predatory from
parasitic traces, and how to recognize failed predation. Our data show that identifying the
predator is easiest when predator and prey are preserved in the act of predation or when predators
were fossilized with their gut contents preserved. However, determining the culprits responsible
for bite traces, drill holes, and other types of predation traces can be more problematic.
Taphonomic and other factors can distort patterns of predation, but their potential effects can be
minimized by careful study design. With the correct identification and quantification of
fossilized traces of predation, ecological trends in predator-prey interactions may be discerned
along environmental gradients in water depth, habitat, and oxygen and nutrient availability.
However, so far, these trends have not been explored adequately for the fossil record. We also
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review the effects of climate change and ocean acidification on predator-prey interactions, but,
again, few studies consider those factors from a deep-time perspective. Finally, fossils have been
used successfully to infer the behavior of ancient predators and identify and interpret active and
passive defense strategies employed by their prey. The marine fossil record of predation has
become a major field of research over the last 50 years, but many critical gaps remain in our
understanding of the evolutionary history of predator-prey interactions.
Keywords: behavior, fossil record, mollusks, parasitism, predation, prey
Contents
1. Introduction
2. Predation database, history and types of studies, and types of data
2.1. Predation database
2.2. History and types of studies
2.3. Types of data
3. Recognizing and identifying predation
3.1. Biotic and abiotic shell damage
3.1.1. Circular and irregular holes
3.1.2. Shell fragments
3.1.3. Non-lethal damage
3.1.4. Correcting for abiotic damage
3.2. Predatory versus parasitic traces
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3.3. Identifying the predator
3.4. Recognizing and interpreting failed predation
3.5. Taphonomic and other factors affecting evidence of predation
4. Environmental factors
4.1. Depth gradients
4.2. Different habitats and ecology
4.3. Oxygen
4.4. Nutrients and food scarcity
4.5. Temperature
4.6. Ocean acidification
5. Behavior of predators
6. Antipredatory features and other defense strategies in deep time
6.1. Active defenses
6.2. Passive defenses
7. Conclusions and future directions
1. Introduction
Predation is viewed by many ecologists as one of the most important factors shaping
marine ecosystems today (e.g., Langerhans, 2007; Paine, 1966). Similarly, many paleontologists
consider predation to have been a major force throughout the evolutionary history of marine
ecosystems (e.g., Signor and Brett, 1984; Stanley, 2008; Vermeij, 1977), and ample evidence
suggests that predator-prey interactions were common from the Neoproterozoic onward (e.g.,
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