Paleobtology, 11(1), 1985, pp.

105-119

Taphonomy's contributions to paleobiology

Anna K. Behrensmeyer and Susan M. Kidwell

Abstract.—Taphonomy established itself in paleontology primarily as a subdiscipline of paleoecology, but

it has evolved into a much broader study of the ways in which preservation affects the fossil record. The
past decade has seen a change in emphasis from descriptive taphonomic studies of fossil assemblages to
more experimental, process-oriented investigations of necrolysis, stratification, and diagenesis of organic
remains in modern environments. These actualistic studies are increasing the sophistication of taphonomic
analysis in the fossil record by sharpening the diagnosis of bias in paleontological data and by providing
a baseline for quantitative modeling of preservational patterns. The analysis of bias is also expanding into
the evaluation of temporal resolution in the fossil record (sample acuity, stratigraphic completeness), and
taphonomic research is thus contributing to broad-scale problems in evolution, biogeography, and bio-
stratigraphy. In addition, taphonomic studies are providing new insights into paleoenvironmental recon-
struction and into the direct paleobiological significance of post mortem processes such as the behavior of
scavengers and the role of dead hardparts in structuring benthic communities. One of taphonomy's most
promising new frontiers is comparative analysis applied to different taxonomic groups within assemblages
and across environments, tectonic settings, and climatic regimes. All of this currendy active research is
contributing to a better understanding of the fossil record as the result of a dynamic, evolving, integrated
system of biological and sedimentological processes that have both limited and enhanced knowledge of
Earth history.

Anna K. Behrensmeyer. Department of Paleobiology, NHB-E207 M.S. 121, Smithsonian Institution, Wash-
ington, D.C. 20560

Susan M. Kidwell. Department of Geosciences, University of Arizona, Tucson, Arizona 85721

Accepted: January 15, 1985

Introduction gies and focal problems, taphonomy is devel-

Taphonomy is concerned with how organic oping a clearer identity as a discipline within the
remains are incorporated into the rock record natural sciences. Most paleontologists, however,
and the fate of these materials after burial. It would still define the field as the study of post
was originally described as the "study of the mortem information loss, stressing the tradition-
transition of organic remains from the biosphere al cautionary role that taphonomic analysis has
into the lithosphere" (Efremov 1940) and has played particularly in paleoecological research.
stressed the recognition and evaluation of the This characterization of taphonomy fails to com-
extent to which fossil assemblages are biased rec- municate its broader concern with how fossil as-
ords of ancient life (Lawrence 1968, 1979). The semblages are changed during preservation by
analysis of bias has remained preeminent in the the addition and alteration of information as well
last decade and has become more quantitative as by its loss. We thus propose a new working
and accurate, largely through inaeased emphasis definition for the field as the study of processes of
on actualistic, process-oriented research. The na- preservation and how they affect information in
ture of taphonomy has expanded beyond this the fossil record. This encompasses not only
focus on bias, however, because of the increasing information loss and bias, but also the more
number of paleontological and geological disci- positive contributions that taphonomy is now
plines that utilize information on the post mor- making to the study of organisms and environ-
tem history of organic remains. Our understand- ments through time.
ing of the fossilization process is rapidly
accelerating owing in large part to constructive A Brief History of Taphonomy
interactions among these fields (Fig. 1). Although the field was named only 45 yr ago
As the primary data concerning preservation (Efremov 1940), the study of fossilization and
become organized around stronger methodolo- the fidelity of the fossil record has a long history.
© 1 9 8 5 The Paleontological Society. All rights reserved. 0094-8373/85/1101-0009/11.00
Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 106 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

SOURCES OF FIRST ORDER ANALYSIS OF PALEOBIOLOGICAL

INFORMATION PRESERVATIONAL EFFECTS PROBLEMS
AND BIASES
SEDIMENTOLOGY PALEOECOLOGY
STRATIGRAPHY SPECIATION PATTERNS
ENVIRONMENTAL
GEOCHEMISTRY CONTEXT NORMAL AND
MASS E X T I N C T I O N
TECTONICS T I M E FACTORS
PALEOCOMMUNITIES
PALEOCLIMATOLOGY ^ TAPHONOMY CONSEQUENCES
OF ANATOMY LIVE:DEAD I N T E R A C T I O N S
ECOLOGY s' / /
B I O L O G I C A L AND PALEOBIOGEOGRAPHY
ICHNOLOGY / /
PHYSICAL DIVERSITY
GEOMORPHOLOGY, PROCESSES
BIOSTRATIGRAPHY
STATISTICS

FIGURE 1. The present structure of taphonomic research, in terms of its interdisciplinary connections, primary first-order
research objectives, and some of the paleobiological problems to which taphonomic information contributes.

The German workers Abel (1912), Wasmund marine invertebrates, Lawrence (1968) on in-
(1926), Weigelt (1927), and Richter (1928) formation loss" in fossil marine communities,
laid the foundations of the field in the first three Clark et al. (1967) on Oligocene vertebrate pa-
decades of the twentieth century, interpreting leoecology, and Voorhies (1969) on bone trans-
both common and unusual fossil deposits in port experiments and Pliocene vertebrate assem-
terms of post mortem processes that operate in blages. Taphonomy became known in the United
modern environments. The strong reliance on States as a prerequisite for paleoecologic research
actualism is traditionally a part of much ta- (Lawrence 1968), and was so firmly linked with
phonomic research, although the pros and cons paleoecology that its relevance to the question of
of analogic reasoning and the validity of as- bias in biostratigraphy and evolution was largely
sumptions concerning the present as a key to the unappreciated. When the importance of study-
past are rarely discussed (but see GifFord 1981). ing preservational biases in such contexts was
Efremov's (1940, 1953) concept of taphon- noted (e.g., Simpson I960), it was not referred
omy was the earliest truly comprehensive view to as taphonomy.
by modern standards, encompassing diagenetic The early 1970s saw an expansion of tapho-
as well as biostratinomic and necrolytic biases in nomic research in Germany, spurred largely by
vertebrate perservation (Fig. 2), and recognizing the Sonderforschungsbereichs Project 53 on Pa-
different scales of bias ranging from those af- leoecology centered on Seilacher's group at the
fecting taxa in a single locality to those imposed University of Tubingen (Seilacher 1976). The
by factors of continental scale. Although Efre- Fossil-Lagerstatten and Fossil-Diagenese pro-
mov's clearly articulated concept of taphonomy grams funded a range of investigations into un-
soon had significant influence in vertebrate pa- usual fossil deposits and modes of preservation
leontology (Olson 1952, 1958, 1980), it failed as a means of better understanding more normal
to unify the diverse studies of preservation into types of occurrences (Seilacher 1970). These
one field. Interest in taphonomic problems has programs alone have produced approximately
grown more or less independently within the 100 papers to date (see listings in Zentralbl. fur
traditional divisions of paleontology and, more Geol. und Palaont., Teil II, 1976; Neues Jahrb.
recently, in archaeology. Geol. und Palaont., Abh. 157, 1979, and Abh.
During the 1950s and 1960s, the most influ- 164, 1982). The acceleration of interest in ta-
ential taphonomic papers in the United States phonomy in the United States during this same
addressed post mortem bias in paleoecologic in- period and its broadening to include stratigraph-
formation. These included the work of Olson ic, sedimentologic, and actualistic approaches are
(1952, 1958) on Permian vertebrates, Johnson due in part to the influence of this German work
(1957, I960, 1962) on fossil and recent shallow and a rediscovery of older pathfinding European
Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY A N D PALEOBIOLOGY 107

SEQUENCE OF PROCESSES SUB-DISCIPLINE

AFFECTING PRESERVATION OF TAPHONOMY

O R G A N I S M D I E S OR BIOSPHERE
SHEDS BODY PARTS

O R G A N I C ( S O F T ) PARTS DECAY
NECROLOGY > ♦ ♦♦
S E D I M E N T A R Y PROCESSES 1 o
y BIOSTRATINOMY z
I N T E R A C T W I T H R E M A I N I N G PARTS
o
BURIAL -> z
CHEMICAL ALTERATION DIAGENESIS <
AND L I T H I F I C A T I O N

COLLECTION
1
1
FIGURE 2. The subdisciplines of taphonomy address fossilization from the perspective of three phases of post mortem
change: decay of softparts and organic matrix, other alteration of remains before burial, and post burial changes. The
influence of biological processes wanes and the importance of physical (including chemical) processes increases through these
phases.

studies, fostered by the translation of Schafer's Cadee 1968; Lawrence 1971; Warme 1969,
actuo-paleontologic opus into English (Schafer 1971; Levinton and Bambach 1975; Scott and
1972). West 1976; MacDonald 1976; Peterson 1976;
In the 1970s, taphonomy also expanded into Schopf 1978). During this same period, actu-
paleoanthropology and archaeology with the alistic comparisons between life and death as-
work of Brain (1958, 1967, 1969), Behrens- semblages also became important in palynology
meyer(1975, 1976a,b), and Hill (1975, 1978), (e.g., Havinga 1967; Davis 1969; Davis and
which emphasized the paleoecological context of Webb 1975; Bonnefille 1979), vertebrate pa-
human evolution and human versus nonhuman leontology (e.g., Zangerl and Richardson 1963;
agencies of bone modification. Taphonomy has Schafer 1972; Hill 1975; Behrensmeyer et al.
been so enthusiastically incorporated into both 1979) and micropaleontology (e.g., Sliter 1975;
the theory and the practice of archaeology (Gif- Berger 1976).
ford 1981) that many recent students in the field The early to mid-1970s also witnessed the
are unaware of its history and origins in paleon- increased use of statistical techniques in taphon-
tology. The application of taphonomy to ar- omy as workers sought to remedy "information
chaeological problems has stimulated a great deal loss," particularly in species diversity and relative
of rethinking of previous interpretations regard- abundance data. Rarefaction techniques devel-
ing early human behavior and resource utiliza- oped by Sanders (1968) for living marine com-
tion. munities we applied to fossil assemblages. "Min-
An early and continuing goal of taphonomic imum numbers" estimates originated by Shotwell
analysis has been to derive accurate estimates of (1955) for vertebrate assemblages have been
species' relative abundances for paleocommunity subject to reanalysis using statistical techniques
reconstruction. In vertebrate paleontology, there (Grayson 1978; Holtzman 1979). The prob-
were a number of attempts to develop meth- lems of large-scale biases in the fossil record have
odologies for this, beginning with the work of also been subjeaed to a wide range of innovative
Shotwell (1955, 1958) and Clark et al. (1967). mathematical modeling techniques (e.g., Sep-
In marine invertebrate faunas, pioneering work koski 1975, 1976; Raup 1976, 1979).
by a number of taphonomists led to understand-
ing of differential preservation, time-averging ef-
fects, and the difficulties in obtaining accurate Current Status and Scope of Taphonomy
information concerning species abundances in the From a traditional viewpoint, taphonomic
original communities (e.g., Johnson 1962, 1965; processes begin when an organism loses control
Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 108 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

PALEOBIOLOQICAL
RECONSTRUCTION

BIOSPHERE
PALEONTOLOGICAL SAMPLE

/ /f
/LITHOSPHERE 'V
/ FOSSILS

X IMMEDIATE BURIAL
^1 ^J^&r
DEAD REMAINS A \ ^%^v"^3^ \ 1 BURIED REMAINS
DISCARDED PARTS 1

Fr^^^^^N.
EXPOSED
\Jm
REMAINS

FIGURE 3. The progression of organic remains through distinct stages from death to final discovery, with intervening
processes acting as filters on paleontological information, has become an accepted paradigm for the biosphere-lithosphere
transition. Taphonomy is the study of how the different components in the system relate to one another and of the processes
that effect the successive transformations.

over the organic components of its body, either taphonomic (rather than paleoecological or sta-
through death or discard, and extend through tistical) is usually determined by the perspective
the preburial interval of exposure and concen- of the researcher. For example, the strength of a
tration through final burial and diagenesis (Fig. cephalopod shell clearly has ecological signifi-
3). Preservation is strongly affected by predeath cance for the animal but is also a critical factor
circumstances such as habitat (e.g., nonaquatic in its potential for preservation and burial. In
vs. aquatic), however, and these can easily be- our survey of the last 10 years of research in
come part of a taphonomic study. Biases in pa- taphonomy, we have included works with ob-
leontological data also inevitably result from vious application to taphonomic problems as well
outcrop patterns, collecting strategies, prepara- as those actually reporting taphonomic analyses.
tion, and even museum curation, hence these Taphonomy varies in its importance to major
also may be included in taphonomic analysis. In topics in paleontological research. Many tradi-
practice, therefore, preservational effects extend tional problems relating to taxonomy and phy-
well beyond the scope of the "biosphere to litho- togeny either bypass obvious taphonomic biases
sphere" transition, and the perspective and or are chosen (often intuitively) to niinirnize such
methodologies of taphonomy can be applied biases. Systematic description of species and
throughout this expanded range of preservation- functional morphology are usually based on well-
al effects. preserved specimens whose structures were not
Whether or not a problem is regarded as affected significantly by taphonomic processes. In

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY AND PALEOBIOLOGY 109

studies of populations, however, processes such taphonomy is represented in about 30%, inver-
as hardpart size sorting and time averaging can tebrate taphonomy in another 30%, and the re-
affect morphometric parameters. Differential maining 40% includes all other fields (e.g., pa-
preservation of species has important and well- leobotany and palynology, micropaleontology,
recognized effects on paleocommunity recon- anthropology, sedimentology, ichnology). The
structions but also relates to problems in bio- vast majority of these publications are descriptive
stratigraphy. Phenomena such as the "Lazarus studies of specific fossil occurrences or strati-
effect," in which species appear to become ex- graphic intervals. Sedimentological papers per-
tinct but then reappear farther up a stratigraphic taining to taphonomy are underrepresented in
column (Jablonski 1983) can reflect taphonomic the Georef listing primarily because key words
processes that therefore bias biostratigraphic cor- (e.g., taphonomy, fossilization, bias) often are
relations and patterns of speciation and extinc- not included in taphonomic articles published in
tion. geological journals. A similar situation exists with
In the past decade, emphasis has changed from regard to biological literature on skeletal pro-
documenting biases in the records of ancient duction, bioerosion, and ecological significance
communities and "stripping off the taphonomic of organic remains (specifically for marine inver-
overprint," to focusing on biologically meaning- tebrates). As a consequence, a great deal of in-
ful information that is contained in time-aver- formation relevant to taphonomy is not easily
aged, ecologically mixed assemblages, and to accessible through presently available key word
sampling designs aimed at these new goals. Bias- listings and library search procedures.
es themselves are now recognized as problem The Georef search suggests a significant ex-
specific, and with the increase in knowledge con- pansion in taphonomic research in the United
cerning biasing processes, taphonomists can re- States relative to other countries since 1975. Of
fine and simplify their studies by isolating and more than 20 other nations represented in the
analyzing biases with respect to carefully defined search, the Soviet Union appears to be the most
problems. Moreover, processes that bias some active. Taphonomic publications in Europeran
types of evidence in fossil assemblages are now university journals and doctoral dissertations
better appreciated as sources of other kinds of outside the United States are underrepresented
information. For example, bone samples with in Georef, however, which obscures some of the
carnivore damage may have taxonomic biases research activity in other countries, particularly
owing to prey seleaion but at the same time Germany. Even with its limitations, the Georef
reveal behavioral traits of the bone-processing listing reflects generally available published work
agent. This positive trend is gradually replacing that influences the overall impact of taphonomy
a commonly held view of taphonomic processes in paleontology, and there is little doubt of the
as obstructive rather than enlightening. increasing interest and involvement of U.S. re-
A Georef search of the past 10 yr of tapho- searchers during the past decade.
nomic literature (Appendix 1) reveals an average The largest proportion of taphonomic articles
publication rate in taphonomy of nearly 50 ar- in Paleobiology (Fig. 4; Appendix 2) are con-
ticles per year since 1975. This is only a mini- cerned with preservational and sampling biases,
mum estimate of the actual number of publi- and these cover a variety of problems ranging
cations relating directly to the field and may be from the effects of differential preservation of
off by a factor of 2 or more since the Georef taxa (e.g., Lasker 1976; Koch and Sohl 1983)
listing of 441 tides included only 19 of 40 ar- to representation of body size classes in mammal
ticles in the first 9 volumes of Paleobiology which assemblages (Damuth 1982) and stratigraphic
have taphonomy as a primary theme (Appendix and area effects on local and Phanerozoic marine
2). Taphonomy was mentioned by name in only diversity (Sepkoski 1975; Flessa and Sepkoski
14 of the 150 Paleobiology articles of taphonom- 1978; Jablonski 1980). Theoretical as well as
ic significance, explaining in part the incomplete- actualistic approaches have focussed on meth-
ness of the Georef listing, which used "taphon- odologies for analyzing bias in fossil assemblages
omy" as the primary key word. (e.g., Schopf 1978; Behrensmeyer 1982a; Buzas
Of the articles found by Georef, vertebrate et al. 1982).

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 110 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

current direction and regime (fossil orientation,

6O-1
geometry and alignment of fossil concentra-
A «p tions), substratum mass properties (bioclastic
50- A ** fabric, hardpart residence time on the seafloor),
Jr and hydraulic energy and sediment transport di-
40- rections (fossil allochthony). While such data are
PERCENT

often collected with sedimentological rather than

30-
paleobiological goals in mind, they are also a
s source of important information about tapho-
M
o nomic processes that affect paleoecological and
20- u. I
—K®^ evolutionary data.

10-

New Directions in Taphonomy

CA8E 8TUDY STRUCTURE
The scope of taphonomy is increasingly broad,
PRESERV./BIAS but it is also becoming more clearly focused on
FIGURE 4. Types of taphonomic information contained in several major new areas of research.
articles in Paleobiology. An article focusing on preservation^ Time resolution. The way in which time is
patterns or biases was catagorized as 'Preserv./Bias.' Other represented in fossil assemblages determines how
central themes were classed as 'Process,' 'Structure' (e.g.,
affecting preservation) or 'General.' (See Appendix 2 for they can be used to address paleoecologic, bio-
further explanation.) stratigraphic, and evolutionary questions. The
temporal significance of paleontological samples
can be examined on several scales. (1) What is
the time represented in a single sample—how
Other themes in Paleobiology and in tapho- many years were required for its formation? This
nomic literature in general include the analysis has been characterized as the problem of time
of skeletal architecture, growth habit, hydraulic averaging or temporal acuity (e.g., Schindel
behavior, and substrate attachment (see "Struc- 1980, 1982). (2) How complete is a fossilifer-
ture" in Appendix 2) and studies of biological ous sequence—what percentage of increments of
and physical processes that leave identifiable a given duration are represented in the rock rec-
traces on organic remains ("Processes" in Ap- ord, and how does lithostratigraphic complete-
pendix 2). Some of these articles are written as ness relate to biostratigraphic completeness? (3)
studies in autecology but have important tapho- How are fossiliferous time increments distribut-
nomic implications (e.g., Schopf et al. 1980; ed through the sequence—are beds (and paleon-
Vermeij et al. 1980; Westermann and Ward tological samples) distributed evenly, randomly,
1980; Cheetham and Thomsen 1981). Publi- or in other patterns that affect the record of pa-
cations focusing on vertebrate taphonomy have leontological change? Methodologies for study-
stressed processes (predators, scavengers, fluvial ing acuity and completeness are now being de-
transport, weathering) that afFea bone assem- veloped for a variety of organisms and
blages and leave recognizable evidence for their depositional settings (see review by Behrensmey-
influence on the assemblages (e.g., Behrensmey- er and Schindel [1983}), but the analysis of se-
er 1978; Andrews and Nesbit Evans, 1983; quence patterns (point 3) remains a frontier.
Haynes 1983). Although time averaging can obscure short-
Not surprisingly, research in sedimentological term paleobiological processes, it also has advan-
aspects of biostratinomy and diagenesis has been tages for paleoecological and neontological sur-
underrepresented in Paleobiology. Analysis of post veys of community composition (Peterson 1976;
mortem histories of fossils is being used to reveal Warme et al. 1976; Scott and Medioli 1980).
the dynamics of sediment deposition and erosion Time averaging integrates "noise" from seasonal
(evidence for cycles of exhumation and burial), and other short-term spatial and temporal vari-
porewater chemistry and timing of lithification ations in species abundance (e.g. Johnson 1971),
(stage of decay, corrosion, mold formation, min- thereby enhancing longer-term patterns that may
eral replacement, and recrystallization), paleo- be of greater significance for the community as
Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY AND PALEOBIOLOGY 111

a whole. Badgley (1982a) used modern ana- samples of comparable temporal acuity by dis-
logues to model time-averaging effects in the tinguishing essentially instantaneous (cata-
vertebrate fossil record. Comparison of two dif- strophic) accumulations from longer-term ones.
ferent types of samples from a recent vertebrate Jones (1980) demonstrated one method for doing
community in East Africa, representing 1 yr and this using molluscan growth lines to determine
100 yr of time averaging of mammal popula- season of death, and catastrophic versus attri-
tions, revealed how each might represent actual tional histories for other types of assemblages can
live populations in the fossil record. A similar be deduced from other kinds of life-history pa-
approach to other types of communities would rameters and taphonomic evidence.
help define the limits to paleocommunity recon- The completeness of the fossil record is now
struction using short-term time-averaged sam- being evaluated using the probabilistic method
ples. of Sadler (1981), which estimates the underlying
In the fossil record, time averaging has been stratigraphic completeness by comparing long-
evaluated on the basis of {a) the ecological com- and short-term sedimentation rates. This meth-
parability of the assemblage to a living com- od has been applied to questions such as how
munity, using arguments based either on recent finely microevolution can be resolved in a par-
analogues or functional morphology, and (b) the ticular stratigraphic section and how sedimentary
taphonomic uniformity of the assemblage— increments are distributed in a given section with
whether features of the specimens themselves respect to an absolute time scale (Gingerich
show comparable post mortem histories. More 1982; Sadler and Dingus 1982; see also NAPC
recently, the mode of formation of the sedimen- Symposium, reviewed in Behrensmeyer and
tary deposit containing a fossil assemblage has Schindel [1983]). The relation between bio- and
also been used to estimate the time it represents lithostratigraphic completeness has been argued
(Behrensmeyer 1982a, b; Kidwell 1982; Retal- several ways: biostratigraphic completeness may
lack 1984; Wing 1984). These estimates are be less (for a given time interval) because of
based on analogy with modern processes that unfossiliferous beds in the sequence (e.g., Din-
concentrate organic remains (e.g., storms, floods, gus and Sadler 1982; Sadler and Dingus 1982),
predation, hydroseres) or on the stratigraphic but the biostratigraphic record may also be more
context of the deposit (e.g., condensed marine complete because of stratigraphic condensation
transgressive lags, paleosol frequency, and dis- of fossils during intervals of low sedimentation
tribution through a section). (Behrensmeyer 1982a,b; Kidwell 1982). In such
Estimates of time averaging based on tapho- instances, the intervals of time that are well rep-
nomic history and other paleoenvironmental cri- resented by fossils are poorly represented by sed-
teria may be at odds with estimates of acuity iments, and these may alternate with other non-
based on statistical treatment of sedimentation condensed increments in which biostratigraphic
rates (e.g., Schindel 1980, 1982). Velbel (1984) completeness is more a function of increased rates
has pointed out that discrepancies of as much as of sedimentation.
6 orders of magnitude can arise because esti- Megabiases.—Analysis of bias in large-scale
mates of acuity based on average sedimentation paleobiologic patterns is another focal area for
rates do not take into account sedimentary pro- growth in taphonomy. "Megabiases" in relation
cesses responsible for the formation of individual to Phanerozoic diversity have already received
beds. For example, Velbel's paleontological considerable attention in a series of recent papers
samples from a 1-cm-thick interval in a sequence (Raup 1976; Sepkoski 1976; Sheehan and Raup
of platform limestones have estimated acuities 1977; Flessa and Sepkoski 1978; Lasker 1978;
of 1,200 yr each if continuous sedimentation is Signor 1978; Sepkoski et al. 1981). These con-
assumed, yet actually may represent a single day's front many of the biases that affect the record
event (e.g., a storm) averaging remains from a of taxa through time, including the effect of
few months or years of attritional death. Ta- exposure area and preserved thickness of strata
phonomic analysis that draws upon knowledge of different ages, proportion of marine versus
of sedimentologic and ecologic processes can re- continental rocks, number of systematic studies
solve time components more precisely than and workers on groups and systems, and the
probabilistic methods and can also pinpoint biostratigraphic bias due to our more complete
Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 112 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

knowledge of Recent faunas relative to ancient affect paleoecological interpretations. The genus
faunas ("The Pull of the Recent" [Raup 1979]). Hipparion has been regarded as a dominant
Further taphonomic work could contribute to member of the mammal community in the Mio-
the analysis of diversity patterns by (1) testing cene of Pakistan after its abrupt appearance about
the effects of time averaging and other tapho- 9.5 ma B.P. Yet through careful sampling and
nomic processes on resolving immigration and taphonomic analysis, Badgley (1982b) found
extinction events, (2) providing a basis for as- that its apparent abundance was more likely due
sessing body size biases in the record, (3) esti- to the differential preservation and identifiability
mating diversity biases due to loss of soft-bodied of its teeth. The reduced importance of Hippar-
organisms in specified environmental settings, (4) ion, a supposed grazer, in the paleocommunity
examining the differences in large-scale preser- changes interpretations of the habitats as open
vation potential for stenotopic versus eurytopic woodland or savanna and also affects hypotheses
organisms. Investigations into other types of concerning its impact on the community struc-
large-scale, environmentally imposed biases could ture and faunal turnover.
also prove fruitful. For example, land vertebrate Differential preservation has also become an
deposits accumulate and are preserved under important consideration in distinguishing real
fairly specific combinations of tectonic, climatic, from apparent extinction events and in placing
and sedimentologic conditions, and the best rec- confidence limits on paleontologic patterns near
ords may result from rapid infilling of continen- mass extinction boundaries (e.g., Lazarus effect
tal basins in relatively dry or highly seasonal ijablonski 1983, 1985]; unconformity trunction
climates. If true, this would imply that much of effect [Birkelund and Hankasson 1982]). Al-
land vertebrate paleoecology and evolution as we though this applies directly to such currently de-
understand it is represented within a rather nar- bated extinaions as the end-Cretaceous event
row, environmentally defined window through (Raup 1982; Van Valen 1984), there has been
time. The same type of phenomenon could also relatively little taphonomic evaluation of com-
apply to macrofloral and pollen records. peting hypotheses. Ecological catastrophes that
Taphonomy may play a larger role in struc- accomplish mass global extinction on the scale
turing diversity patterns than is generally rec- of days, weeks, and months (e.g., extreme ver-
ognized, both in terms of differing post mortem sions of the asteroid impact hypothesis of Al-
biases among assemblages and in the changing varez et al. [1980], such as Ahrens and O'Keefe
role of dead hardparts on community dynamics. [1983] and Pollack et al. [1983]) might leave
For example, to what extent do Bambach's a qualitatively different taphonomic record than
(1977) estimates of within-habitat diversity re- more prolonged gradual or stepped extinaions.
flea qualitatively different patterns of tapho- A change in the preserved record might be ex-
nomic bias in his samples? The Neogene data peaed to result from a sudden inaease in rate
are derived largely from condensed shell beds of of hardpart supply to the record, enhanced by
the Atlantic coastal plain, where fossil molluscan the swamping (or inaaivity) of scavengers and
assemblages record prolonged time averaging and decomposers.
a mixture of habitat types within the nearshore What would be the taphonomic consequence
setting. The condensed nature of these deposits of other faaors in the final Cretaceous event,
would tend to increase apparent taxonomic di- such as reduction in shallow marine sample area
versity for the habitat. Moreover, because shell and possibly accelerated terrestrial erosion asso-
gravels provide opportunities for taphonomic ciated with deforestation, regression, and base-
feedback that are not available in communities level fall? Additionally, is the stratigraphic record
from soft-bottom habitats of identical bathym- of the aitical interval too time averaged or in-
etry, the two types of fossil assemblages are complete to resolve the short-term dynamics of
probably not comparable for biological reasons. a mass extinction? And what might be the effeas
Another important taphonomic effect on of small sample size (e.g., for dinosaur remains)
community structure and evolution lies in dif- on interpretations of the observed record? The
ferential preservation of taxa, which can strongly taphonomy of mass extinaions deserves further

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY AND PALEOBIOLOGY 113

consideration, both in terms of the fidelity of the nodules have greater preservation potential than
fossil record and as a potential source of direct dispersed bone elements (A. Hill, pers. comm.).
information on the biological nature of the events. There is also growing evidence that rapid burial
Feedback systems.—Since taphonomic phe- of soft tissues promotes geochemical changes that
nomena straddle the boundary between the bio- enhance pyrite and carbonate nodule formation
sphere and lithosphere, information on the post and thus fossil preservation. It thus appears that
mortem fate of organic remains provides insights early diagenesis may accentuate primary patterns
concerning interactions between biological and in hardpart abundance created by physical and
geological systems. Positive and negative feed- biological processes: the diagenetic filter on pa-
back between these systems has become a pro- leontologic information is not random, nor is it
ductive line of new research. For example, al- in opposition to the preservation of primary pat-
though living organisms influence the terns in hardpart accumulation.
composition of the death assemblage, the dead The role of organic remains in feedback sys-
hardparts also have potential influence on the tems that influence not only living organisms but
composition and dynamics of living benthic also preservation potential offers fertile ground
communities by altering the physical habitat. for future taphonomic research. One conse-
Dead hardparts provide attachment sites for or- quence of such feedback may be a bias toward
ganisms and, by changing the mass properties massed organic remains and the ecological and
(firmness, texture, topography) of the substrate, sedimentary situations favoring these throughout
facilitate colonization and survival for firm-bot- the fossil record.
tom and shell-gravel taxa while inhibiting the Comparative taphonomy.—Documentation of
success of soft-bottom, primarily infaunal groups. taphonomic features in fossil and recent case
The entire spectrum of interrelationships be- studies is building an essential data base for the
tween living and dead hardparts has been termed field, and there is no question that such research
taphonomic feedback (Kidwell and Jablonski will and should continue. There are also signs,
1983), which serves as a driving mechanism of however, that taphonomy is entering a more syn-
community change that is highly dependent upon thetic and theoretical phase in which the vast
the physical sedimentary dynamics of hardpart amount of information on preservation can be
accumulation (e.g., Kidwell and Aigner, 1985). organized into a more coherent approach to the
Although recognition of the effects of hardparts fossil record. In particular, there is fertile ground
on the marine benthos is not new (e.g., studies for comparisons of similar taphonomic phenom-
of encrusting and boring taxa in shell substrate ena that operate differently on major taxonomic
[Walker and Parker 1976; Watkins and Hurst groups both within assemblages and across dif-
1977]), a perspective stressing live:dead inter- ferent paleoenvironmental settings. For example,
actions as a dynamic feedback system provides why are plants and vertebrates so seldom pre-
the basis for a new look at benthonic marine served in the same sedimentary bed? Are assem-
ecosystems. blages from biostratinomically identical storm
In addition to the ecological consequences of lags in different latitudes and climates compa-
live: dead interactions, positive feedback prob- rable in their early diagenesis? Have taphonomic
ably operates as well between processes of hard- pathways changed qualitatively or only quanti-
part concentration and their preservation poten- tatively through the Phanerozoic as a conse-
tial. For example, buried hardparts in shallow quence of biological evolution and Earth history?
marine settings are more likely to create favor- A more integrated, comparative taphonomic
able chemical microenvironments and thus sur- approach to the fossil record will also help iden-
vive early diagenesis when associated with other tify areas needing further research. For example,
hardparts (e.g., MacCarthy 1977; Sepkoski the taphonomy of microorganisms and macro-
1978; Aller 1982; Fursich 1982). Vertebrate floral remains has received far less attention than
remains may reflect similar feedback systems in that of vertebrates and macroinvertebrates. Fur-
fluvial deposits, where dense primary bone con- thermore, vertebrate taphonomy has focused pri-
centrations and those associated with calcareous marily on preservation in terrestrial settings and

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 114 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

invertebrate taphonomy has stressed marine set- in fluvial settings by controlling the balance of
tings in both actualistic and stratigraphic inves- such processes as lateral versus vertical aggra-
tigations. The characteristics and origin of ma- dation.
rine bone beds and freshwater shell beds are thus Climate is another controlling factor in the
less well understood despite their abundant rep- preservation of organic remains, and probably
resentation in the record. one of the least understood. Paleobotanists rec-
Taphonomic comparisons among major taxo- ognize a clear latitudinal gradient from low pres-
nomic groups are usually confounded by pa- ervation potential in the tropics, where produc-
leoenvironmental differences in the typical set- tivity is high but destruction rates are also high,
tings of fossil accumulation. Deposits of mixed to good macrofloral records in temperate settings
Composition such as shelly bone beds and strati- where seasonality keeps production and destruc-
graphic sequences that are alternately rich in mac- tion out of phase, to poor arctic records related
rofloral remains and vertebrates are ideal for to low overall production. Rates and patterns of
study because many potentially critical paleoen- net bone input may vary similarly. Probable lat-
vironmental factors can be controlled. Nonethe- itudinal variation in net supply of marine inver-
less, because taphonomists almost always con- tebrate remains is not as apparent—how do rates
centrate on a single major group, taxonomically of hardpart production, bioerosion, sedimenta-
mixed deposits are usually examined only from tion, microbial decomposition, and dissolution
one perspective. The advantages of a compara- correlate with climate?
tive approach have been exemplified, however, Another target for comparative taphonomy
by the results of Brett and others (Brett and (which we anticipate with considerble excite-
Baird 1984; Speyer and Brett 1984; Brett et al., ment) is the analysis of Phanerozoic trends in
in press) on bathymetric trends in the preserva- post mortem processes, including preservational
tion of different taxonomic groups in Devonian biases and dynamic interactions between biolog-
strata. Mode of death, history of hardpart con- ical and geological processes. Opportunities and
centration and interment, and porewater chem- pathways for taphonomic feedback, for example,
istries differ along the gradient as a function of have no doubt changed through the Phanerozoic
sediment dynamics, water energy, and oxygen- as a consequence of the evolution of hardpart
ation. Through such investigations, taphonomic producers, hardpart utilizers, and hardpart de-
fades models are beginning to emerge. stroyers (Kidwell and Jablonski 1983) The pres-
Comparative taphonomy of depositional ba- ervation potential of physically concentrated fos-
sins is another promising area for study. To the sil beds and the chemical diagenesis of fossils in
extent that rates of sedimentation govern the general have probably also undergone qualita-
concentration of fossil hardparts, the distribution tive changes through the Phanerozoic, as evi-
of fossil deposits and their degree of time aver- denced by the diminishing preservation potential
aging should differ among active margins, pas- of storm-generated beds (Sepkoski 1982; Larson
sive margins, and cratonic settings. On a smaller and Rhoads 1983), shifting relative abundance
scale, condensed hardpart concentrations associ- of hardpart mineralogies, and increasing depth
ated with unconformities and other breaks in the of penetration and bathymetric range of biotur-
record should differ according to the history of bators and abundance of duraphagous predators
deposition: starved condensed sequences have dif- (Vermeij 1977; Thayer 1983; Miller 1984). Ac-
ferent dynamics of accumulation (and thus taph- tualistic models can play an important role in
onomic attributes) than sequences condensed framing hypotheses for this comparative taphon-
through alternating deposition and erosion (Kid- omy of the Phanerozoic, complementing and ex-
well 1981, 1983; Kidwell and Jablonski 1983). plaining system-by-system patterns.
Patterns of sediment accumulation affect the dis-
tribution of shell gravels within cyclic transgres- Conclusion
sive-regressive sequences (Kidwell and Aigner, The expansion of taphonomy beyond its tra-
1985). Tectonic setting may also determine the ditional role as the study of information loss and
nature and distribution of bone accumulations paleoecologic bias into broader problems in pa-

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY AND PALEOBIOLOGY 115

leobiology has opened up a wealth of new ave- on broad-scale problems and more comparative,
nues for research and interdisciplinary exchange. synthetic inquiries into processes of preservation
The past decade of growth in paleontology has and their consequences in the fossil record.
generated new questions and demands on infor-
mation contained in the fossil record, thus help- Acknowledgments
ing to stimulate more careful consideration of We are grateful to all of the colleagues who
biases in this information. The increasingly rig- have contributed to the growth of our ideas con-
orous approach to bias is, however, only half the cerning taphonomy over the past several years.
story of taphonomy's new identity since tapho- In particular, we thank Catherine Badgley, Da-
nomic processes and effects are now seen as an vid Jablonski, and Carl Brett for their careful
important source of information about the in- readings of earlier versions of this paper and for
terrelationships and dynamics of geological and their perceptive suggestions and discussions. We
biological processes throughout life's history. We have also benefited from conversations with
are now looking more explicitly at the " infor- Marty Buzas and Scott Wing.
mation filters" as well as the remains and traces
that passed through them to become the fossil Literature Cited
record. ABEL, O. 1912. Grundzuge der Palaeobiologie der Wirbeltiere.
The new perspectives in taphonomy which we E. Schweizerbartsche Verlagsbuchhandlung Nagele; Stuttgart.
AHRENS, T. J. AND J. D. O'KEEFE. 1983. Impact of an asteroid
feel are likely to set the course for its immediate or comet in the ocean and extinction of terrestrial life. In: Boyton,
future are: W . V. and T. H. Abrams, eds. Proc. 13th Lunar and Planet. Sci.
1) that taphonomic processes and biases cover Conf., J. Geophys. Res. B88(Suppl.2):A799-A806.
ALLER, R. C. 1982. Carbonate dissolution in nearshore terrigenous
the full range of scales in paleontological prob- muds: the role of physical and biological reworking. J. Geol. 90:
lems, from studies of single species to Phanero- 79-96.
zoic diversity trends. There is a need for research ALVAREZ, L. W., W . ALVAREZ, F. ASARO, AND H. V. MICHEL. 1980.
Extraterrestrial cause for the Cretaceous-Tertiary extinction. Sci-
in middle- to large-scale taphonomic phenome- ence. 208:1095-1108.
na that draws upon knowledge of smaller-scale ANDREWS, P. AND E. NESBIT EVANS. 1983. Small mammal bone
patterns and looks for ways in which these are accumulations produced by mammalian carnivores. Paleobiology.
9:289-307.
magnified or canceled out by longer-term, larg- BADGLEY, C. E. 1982a. How much time is represented in the
er-scale processes. present? The development of time-averaged modern assemblages
2) that processes of preservation can be viewed as models for the fossil record. Proc. 3d N . Am. Paleontol. Conv.
7:23-28.
as active forces which not only have affected the BADGLEY, C. E. 1982b. Community reconstruction of a Siwalik
preserved record of biological evolution but which mammalian assemblage. Ph.D. Dissertation, Yale Univ. 364 pp.
have the potential to direaly influence biological BAMBACH, R. K. 1977. Species richness in marine benthic habitats
through the Phanerozoic. Paleobiology. 3:152-168.
evolution itself. The dynamic interactions be- BEHRENSMEYER, A. K. 1975. The taphonomy and paleoecology of
tween biological and geological processes in par- Plio-Pleistocene vertebrate assemblages of Lake Rudolf, Kenya.
ticular deserve more intensive research by pale- Bull. Mus. Comp. Zool. 246:473-578.
BEHRENSMEYER, A. K. 1976a. Fossil assemblages in relation to
ontologists, biologists, sedimentologists, and sedimentary environments in the East Rudolf succession. Pp. 3 8 3 -
geochemists. 401. In: Coppens, Y., F. C. Howell, G. L. Isaac, and R. E.
Although continuing research in modern an- Leakey, eds. Earliest Man and Environments in the Lake Rudolf
Basin. Univ. Chicago Press; Chicago. 615 pp.
alogues and theoretical modeling is critical to the BEHRENSMEYER, A. K. 1976b. Taphonomy and paleoecology in
further growth of taphonomy, a more careful the hominid fossil record. Yb. Phys. Anthropol. 79:36-50.
consideration of the limits of analogic reasoning BEHRENSMEYER, A. K. 1978. Taphonomic and ecologic informa-
tion from bone weathering. Paleobiology. 4:150-162.
is overdue, as is the development of methods for BEHRENSMEYER, A. K. 1982a. Time resolution in fluvial vertebrate
better understanding processes that are not rep- assemblages. Paleobiology. 8:211-228.
resented in recent biological and geological sys- BEHRENSMEYER, A. K. 1982b. Time sampling intervals in the ver-
tebrate fossil record. Proc. 3d N . Am. Paleontol. Conv. 7:41-45.
tems. Practically speaking, however, perhaps the BEHRENSMEYER, A. K. AND D. SCHINDEL. 1983. Resolving time in
greatest challenge faced by taphonomy is to im- paleobiology. Paleobiology. 9:1-8.
pose order on the vast body of multidisciplinary BEHRENSMEYER, A. K., D. WESTERN, AND D. E. DECHANT BOAZ.
1979. New perspectives in vertebrate paleoecology from a recent
data that can contribute to its goals. This order bone assemblage. Paleobiology. 5:12-21.
is beginning to appear with the growing focus BERGER, W . H. 1976. Biogenesis in deep sea sediments: produc-

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 116 ANNA K. BEHRENSMEYER & SUSAN M. KIDWELL

tion, preservation and interpretation. In: Riley, J. P. and R. Ches- M. B., ed. Advances in Archaeological Method and Theory, Vol.
ter, eds. Chem. Oceanogr. .5:265-388. 4. Academic Press; New York.
BIRKELUND, T. AND E. HANKASSON. 1982. The terminal Cretaceous GINGERICH, P. D. 1982. Time resolution in mammalian evolution:
extinction in Boreal shelf seas—a multicausal event. Geol. Soc. sampling, lineages and faunal turnover. Proc. 3d N . Am. Paleon-
Am. Spec. Pub. 190:373-384. tol. Conv. 1:205-210.
BONNEFILLE, R. 1979. Methode palynologique et reconstitutions GRAYSON, D. K. 1978. Reconstructing mammalian communities:
paleodimatiques au CenozoTque dans le Rift Est Africain. Bull. a discussion of Shotwell's method of paleoecological analysis. Pa-
Soc. Geol. France 27:331-342. leobiology. 4:77-81.
BRAIN, C. K. 1958. The Transvaal Ape-Man-Bearing Cave De- HAVINGA, A. J . 1967. Palynology and pollen preservation. Rev.
posits. Transvaal Mus. Mem. 13:1-125. Palaeobot. Palynol. 2:81-98.
BRAIN, C. K. 1967. Hottentot food remains and their meaning HAYNES, G. 1983. A guide for differentiating mammalian car-
in the interpretation of fossil bone assemblages. Sci. Pap. Namib. nivore taxa responsible for gnaw damage to herbivore limb bones.
Desert Res. Sta. 32:1-11. Paleobiology. 9:164-172.
BRAIN, C. K. 1969. The contribution of Namib Desert Hottentots HILL, A. P. 1975. Taphonomy of contemporary and late Cenozoic
to an understanding of Australopithecine bone accumulations. Sci. East African vertebrates. Ph.D. dissertation, Univ. London.
Pap. Namib. Desert Res. Sta. 39:13-22. HILL, A. P. 1978. Taphonomical background to fossil man-prob-
BRETT, C. E. AND G. C. BAIRD. 1984. Comparative taphonomy: lems in paleoecology. Pp. 87-102. In: Bishop, W . W., ed. Geo-
a key to interpretation of sedimentary sequences using fossil pres- logical Background to Fossil Man. Univ. Toronto Press; Toronto.
ervation. GSA Abstracts (Southeastern and North-Central Sec- 585 pp.
tions) 16:126. HOLTZMAN, R. C. 1979. Maximum likelihood estimation of fossil
BRETT, C. E., S. E. SPEYER, AND G. C. BAIRD. In press. Storm- assemblage composition. Paleobiology. 3:77-89.
generated sedimentary units: tempestite proximality and event JABLONSKI, D. 1980. Apparent versus real biotic effects of
stratification in the Middle Devonian Hamilton Group. In: Brett, transgressions and regressions. Paleobiology. 6:397-407.
C. E., ed. Stratigraphy and Depositional Environments of the JABLONSKI, D. 1983- Apparent versus real extinctions at the end
Hamilton Group (Middle Devonian) in New York. New York of the Cretaceous Period. Geol. Soc. Am. Abstr. 13:602.
State Mus. Sci. Serv. Bull. JABLONSKI, D. 1985. Causes and consequences of mass extinction:
BUZAS, M. A., C. F. KOCH, STEPHEN J . CULVER, AND N . F. SOHL. a comparative approach. In: Elliott, D. K., ed. Dynamics of Ex-
1982. On the distribution of species occurrence. Paleobiology. tinction. Wiley; New York.
8:143-150. JOHNSON, R. G. 1957. Experiments on the burial of shells. J.
CADEE, G. C. 1968. Molluscan biocoenoses and thanatocoenoses Geol. 65:527-535.
in the Ria de Arosa, Galicia, Spain. Zool. Verh. Rijksmus. Nat. JOHNSON, R. G. I960. Models and methods for analysis of the
Hist. Leiden. 93:1-121. mode of formation of fossil assemblages. Bull. Geol. Soc. Am.
CHEETHAM, A. H. AND E. THOMSEN. 1981. Functional morphology 71:1075-1086.
of arborescent animals: strength and design of cheilostome bryo- JOHNSON, R. G. 1962. Mode of formation of marine fossil assem-
zoan skeletons. Paleobiology. 7:355-383. blages of the Pleistocene Millerton Formation of California. Bull.
CLARK, J., J . R. BEERBOWER, AND K. K. KIETZKE. 1967. Oligocene Geol. Soc. Am. 73:113-130.
sedimentation, stratigraphy and paleodimatology in the Big Bad- JOHNSON, R. G. 1965. Pelecypod death assemblages in Tomales
lands of South Dakota. Fieldiana Geol. 3:1-158. Bay, California. J . Paleontol. 39:80-85.
DAMUTH, J . 1982. Analysis of the preservation of community JOHNSON, R. G. 1972. Conceptual models of benthic marine com-
structure in assemblages of fossil mammals. Paleobiology. 8:434- munities. Pp. 148-159. In: Schopf, T. J . M., ed. Models in
446. Paleobiology. W . H . Freeman; San Francisco.
DAVIS, M. B. 1969. Climatic changes in southern Connecticut JONES, D. S. 1980. Annual cycle of shell growth increment for-
recorded by pollen deposition at Rogers Lake. Ecology. 30:409- mation in two continental shelf bivalves and its paleoecological
422. significance. Paleobiology. 6:331-340.
DAVIS, M. B. 1975. Reconstructions of local and regional Holo-
KIDWELL, S. M. 1981. Stratigraphic patterns of fossil accumulation
cene environments from the pollen and peat stratigraphies of some
and their paleobiologic significance. Geol. Soc. Am. Abstr. 13:
Driftless Area peat deposits. Ph.D. thesis, Univ. Wisconsin. 187
486.
pp.
KIDWELL, S. M. 1982. Time scales of fossil accumulation: patterns
DAVIS, M. B. AND T. WEBB III. 1975. The contemporary distri-
from Miocene benthic assemblages. Proc. 3d N . Am. Paleontol.
bution of pollen in eastern North America: a comparison with
Conv. 1:295-300.
the vegetation. Quat. Res. 3:435-465.
KIDWELL, S. M. 1983. Stratigraphic taphonomy: predicting oc-
DINGUS, L. AND P. M. SADLER. 1982. The effects of stratigraphic
currence, time scales, and modes of formation of fossil deposits.
completeness on estimates of evolutionary rates. Syst. Zool. 31:
Geol. Soc. Am. Abstr. 13:612.
400-412.
EFREMOV, I. A. 1940. Taphonomy: a new branch of paleontology. KIDWELL, S. M. AND T. AIGNER. 1985. Sedimentary dynamics of
Pan-Am. Geol. 74:81-93. complex skeletal accumulations: paleoecological and evolutionary
EFREMOV, I. A. 1953. Taphonome et annales geologiques. Ann. implications. In: Bayer, U. and A. Seilacher, eds. Cycles in Sedi-
Centre d'Etud. et de Doc. Paleontol. No. 4:1-164. (Transl. by mentation and Evolution. Springer-Verlag; Berlin.
S. Ketchian and J . Roger) KIDWELL, S. M. AND D. JABLONSKI. 1983. Taphonomic feedback:
FLESSA, K. W . AND J . J . SEPKOSKI, J R . 1978. On the relationship ecological consequences of shell accumulation. Pp. 195-248. In:
between Phanerozoic diversity and changes in habitable area. Pa- Tevesz, J. S. and P. L. McCall, eds. Biotic Interactions in Recent
leobiology. 4:359-366. and Fossil Benthic Communities. Plenum; New York.
FURSICH, F. T. 1982. Rhythmic bedding and shell bed formation KOCH, C. F. AND N . F. SOHL. 1983. Preservational effects in
in the Upper Jurassic of East Greenland. Pp. 208-222. In: Ein- paleoecological studies: Cretaceous mollusc examples. Paleobiol-
sele, G. and A. Seilacher, eds. Cyclic and Event Stratification. ogy. 9:26-34.
Springer-Verlag; Berlin. LARSON, D. W . AND D. C. RHOADS. 1983. The evolution of in-
GIFFORD, D. P. 1981. Taphonomy and paleoecology: a critical faunal communities and sedimentary fabrics. Pp. 627-646. Biotic
review of archeology's sister discipline. Pp. 365-438. In: Schiffer, Interactions and Fossil Benthic Communities. Plenum; New York.

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 T A P H O N O M Y A N D PALEOBIOLOGY 117

LASKER, H . 1976. Effects of differential preservation on the mea- SCOTT, D. B. AND F. S. MEDIOLI. 1980. Living vs. total forami-
surement of taxonomic diversity. Paleobiology. 2:84-93. niferal populations: their relative usefulness in paleoecology. J .
LASKER, H. 1978. The measurement of taxonomic evolution: pres- Paleontol. .5*814-831.
ervational consequences. Paleobiology. 4:135-149. SCOTT, R. W . AND R. R. WEST. 1976. Structure and Classification
LAWRENCE, D. R. 1968. Taphonomy and information losses in of Paleocommunities. Dowden, Hutchinson & Ross; Stroudsburg,
fossil communities. Bull. Geol. Soc. Am. 79:1315-1330. Pa. 291 pp.
LAWRENCE, D. R. 1971. The nature and structure of paleoecology. SEILACHER, A. 1970. Begriffund Bedeutungder Fossillagerstatten.
J. Paleontol. 45:593-607. N. Jahrb. Geol. Palaontol. Abh. 7970:34-39.
LAWRENCE, D. R. 1979. Taphonomy. Pp. 793-799. In: Fair- SEILACHER, A. 1976. Allgemeiner Uberblick, Bericht 1970-1975
bridge, R. and D. Jablonski, eds. Encyclopedia of Paleontology. des Sonderforschungsbereichs 53 Palokologie. Zentralblatt fur
Vol. 7. Dowden, Hutchinson & Ross; Stroudsburg, Pa. Geologie und Palaontologie, Teil 77:206-210.
LEVINTON, J. S. AND R. K. BAMBACH. 1975. A comparative study SEPKOSKI, J . J., J R . 1975. Stratigraphic biases in the analysis of
of Silurian and Recent deposit-feeding bivalve communities. Pa- taxonomic survivorship. Paleobiology. 7:343-355.
leobiology. 1:97-124. SEPKOSKI, J . J., J R . 1976. Species diversity in the Phanerozoic:
MACCARTHY, B. 1977. Selective preservation of mollusc shells in species-area effects. Paleobiology. 2:298-303.
a Permian beach environment, Sydney Basin, Australia. N . Jahrb. SEPKOSKI, J . J., J R . 1978. Taphonomic factors influencing the
Geol. Palaontol., Mh. 8:466-474. lithologic occurrence of fossils in Dresbachian (Upper Cambrian)
MACDONALD, K. B. 1976. Paleocommunities: toward some con- shaley facies. Geol. Soc. Am. Abstr. 70:490.
fidence limits. Pp. 87-106. In: Scott, R. W . and R. R. West, SEPKOSKI, J . J., JR. 1982. Flat-pebble conglomerates, storm de-
eds. Structure and Classification of Paleocommunities. Dowden, posits, and the Cambrian bottom fauna. Pp. 371-385. In: G.
Hutchinson, & Ross; Stroudsburg, Pa. Einsele and A. Seilacher, eds. Cyclic and Event Stratification.
MILLER, M. F. 1984. Distribution of biogenic structures in Paleo- Springer-Verlag; Berlin.
zoic nonmarine and marine-margin sequences: an actualistic mod- SEPKOSKI, J. J., J R . , R. K. BAMBACH, D. M. RAUP, AND J . W.
el. J . Paleontol. .58:550-570. VALENTINE. 1981. Phanerozoic marine diversity and the fossil
OLSON, E. C. 1952. The evolution of a Permian vertebrate chron- record. Nature. 293:435-437.
ofauna. Evol. 6:181-196. SHEEHAN, P.M. AND D. M. RAUP. 1977. Species diversity in the
OLSON, E. C. 1958. Fauna of the Vale and Choza: 14. Summary, Phanerozoic: a reflection of labor by systematists? or systematists
review, and integration of the geology and the faunas. Fieldiana follow the fossils. Paleobiology. 3:325-329.
Geol. 70:397-448. SHOTWELL, J. A. 1955. An approach to the paleoecology of mam-
OLSON, E. C. 1980. Taphonomy: its history and role in com- mals. Ecology. 36:327-337.
munity evolution. Pp. 5-19. In: Behrensmeyer, A. K. and A. SHOTWELL, J . A. 1958. Inter-community relationships in Hemp-
Hill, eds. Fossils in the Making. Univ. Chicago Press; Chicago. hillian (mid-Pliocene) mammals. Ecology. 39:271-282.
PETERSON, C. H . 1976. Relative abundances of living and dead SIGNOR, P. W . 1978. Species richness in the Phanerozoic: an in-
molluscs in two California lagoons. Lethaia. 9:137-148. vestigation of sampling effects. Paleobiology. 4:394-406.
POLLACK, J . B., O. B. TOON, T. P. ACKERMAN, C. T. MCKAY, AND SIMPSON, G. G. I960. The history of life. Pp. 117-180. In: Tax,
R. P. TURCO. 1983. Elemental effects of an impact-generated S., ed. Evolution after Darwin. Vol. 7. Univ. Chicago Press;
dustcloud: implications for the Cretaceous-Tertiary extinctions. Chicago.
Science. 229:287-289. SLITER, W . V. 1975. Foraminiferal life and residue assemblages
RAUP, D. M. 1976. Species diversity in the Phanerozoic: a tabu- from Cretaceous slope deposits. GSA Bull. 86:897-906.
lation. Paleobiology. 2:279-288. SPEYER, F. E. AND C. E. BRETT. 1984. Comparative taphonomy of
RAUP, D. M. 1979. Biases in the fossil record of species and Middle Devonian trilobite beds. GSA Abstr. (Southeastern and
genera. Bull. Carnegie Mus. Nat. Hist. 13:85-91. North-Central Sections). 76:198.
RAUP, D. M. 1982. Large body impacts and terrestrial evolution THAYER, C. W . 1983. Sediment-mediated biological disturbance
meeting, October 19-22, 1981. Paleobiology. 8:1-3. and the evolution of the marine benthos. Pp. 480-625. In: Tev-
RETALLACK, G. 1984. Completeness of the rock and fossil record: esz, M. J. S. and P. L. McCall, eds. Biotic Interactions in Recent
some estimates using fossil soils. Paleobiology 70:59-78. and Fossil Benthic Communities. Plenum; New York.
RICHTER, R. 1928. Aktuopalaontologie und Palaobiologie, eine VAN VALEN, L. M. 1984. Review of Silver, L. T. and P. H .
Abgrenzung. Senckenbergiana. 70:285-292. Schultz, eds. Geological Implications of Impacts of Large Aster-
SADLER, P. M. 1981. Sediment accumulation rates and the com- oids and Comets on the Earth. Paleobiology. 70:121-137.
pleteness of stratigraphic sequences. J . Geol. 89:569-584. VELBEL, D. B. 1984. Sedimentology and taphonomy in a clastic
SADLER, P. M. AND L. DINGUS. 1982. Expected completeness of Ordovician sea. Geol. Soc. Am. Abstr. 76:204.
sedimentary sections: estimating a time-scale dependent, limiting VERMEIJ, G. J. 1977. The Mesozoic marine revolution: evidence
factor in the resolution of the fossil record. Proc. 3d N . Am. from snails, predators and grazers. Paleobiology. 3:245-258.
Paleontol. Conv. 2:461-464. VERMEIJ, G. J., E. ZIPSER, AND E. C. DUDLEY. 1980. Predation in
SANDERS, H . L. 1968. Marine benthic diversity: a comparative time and space: peeling and drilling in terebrid gastropods. Pa-
study. Amer. Nat. 702:243-282. leobiology. 6:352-364.
SCHAFER, W . 1972. Ecology and Palaeoecology of Marine Envi- VOORHIES, M. 1969. Taphonomy and population dynamics of an
ronments. Univ. Chicago Press; Chicago. 568 pp. early Pliocene vertebrate fauna, Knox County, Nebraska. Contrib.
SCHINDEL, D. E. 1980. Microstratigraphic sampling and the limits Geol., Spec. Paper No. 1. Univ. Wyoming; Laramie.
of paleontologic resolution. Paleobiology. 6:408-426. WALKER, K. R. AND W . C. PARKER. 1976. Population structure
SCHINDEL, D. E. 1982. Resolution analysis: a new approach to the of a pioneer and a later stage species in an Ordovician ecological
gaps in the fossil record. Paleobiology. 8:340-353. succession. Paleobiology. 2:191-201.
SCHOPF, T. J . M. 1978. Fossilization potential of an intertidal WARME, J . E. 1969. Live and dead molluscs in a coastal lagoon.
fauna: Friday Harbor, Washington. Paleobiology. 4:261-270. J. Paleontol. 43:141-150.
SCHOPF, T. J . M., K. O. COLLIER, AND B. O. BACH. 1980. Relation WARME, J . E. 1971. Paleoecological aspeas of a modem coastal
of the morphology of stick-like bryozoans at Friday Harbor, lagoon. Univ. Cal. Publ. Geol. 87:1-131.
Washington, to bottom currents, suspended matter and depth. WARME, J . E., A. A. EKDALE, S. F. EKDALE, AND C. H . PETERSON.
Paleobiology. 6:466-476. 1976. Raw material of the fossil record. Pp. 169-243. In: Scott,

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 118 A N N A K. B E H R E N S M E Y E R & S U S A N M . K I D W E L L

R. W . and R. R. West, eds. Structure and Classification of Pa- The primary orientation of the article as presented by the author(s) was used
leocommunities. Dowden, Hutchinson, & Ross; Stroudsburg, Pa. to determine its category in the above scheme. Studies of specific fossil occur-
rence, for example, were placed in 3, even though processes were also a topic of
WASMUND, E. 1926. Biocoenose und thanatocoenose. Arch. Hy- discussion. Studies in which processes themselves were the primiary focus were
drobiol. 77:1-116. placed in 2. The results of this analysis are shown in Fig. 4.
WATKINS, R. AND J . M. HURST. 1977. Community relations of The alphabetical listing below shows the categorization of articles with ta-
phonomic content. Primary concern with taphonomy is indicated as "Pri," in-
Silurian crinoids at Dudley, England. Paleobiology. 3:207-217. direct relevance as "ind," and incidental relevance as "Inc."
WEIGELT, J . 1927. Resente Wirbeltierleichen und ihre Palaobio-
logische Bedeutung. Verlag von Max Weg, Leipzig. 227 pp.
WESTERMANN, G. E. G. AND P. WARD. 1980. Septum morphology
Author Vol. Num. Category Status
and bathymetry in cephalopods. Paleobiology. 6:48-50.
W I N G , S. L. 1984. Relation of paleovegetation to geometry and Ackersten et al. 9 3 Bias-fossil Inc
cydicity of some fluvial carbonaceous deposits. J . Sedimentol. Andrews & Evans 9 3 Process-biological Pri
Andrews & Evans 5 1 Bias-combo Ind
Petrol. .54:52-66. Antia 3 4 Bias-recent Pri
ZANGERL, R. AND E. S. RICHARDSON, J R . 1963. The paleoecological Ashton & Rowell 1 2 Bias-fossil Ind
history of two Pennsylvanian black shales. Fieldiana—Geol. Mem. Bam bach 3 2 Bias-fossil Ind
Behrensmeyer & Schindel 9 1 Bias-fossil Ind
4:1-352. Behrensmeyer 8 Process-physical Pri
3
Behrensmeyer 4 2 Process-combo Pri
Behrensmeyer et al. 5 1 Bias-recent Pri
Appendix 1. Bell & Haglund 8 3 Bias-fossil Ind
Berg & Nishenko 1 3 Process-biological Ind
Results of the Georef Database search Bolt & Wassersug 1 3 Process-biological Inc
Computer access to the Georef Database was through the Dialog Information Bretsky & Bretsky 1 3 Bias-fossil Ind
Retrieval Service, provided in conjunction with Smithsonian Institution Libraries. Brett & Lindell 4 3 Bias-fossil Ind
Georef s database includes over 4500 international journals plus books, disser- Buzas et al. 8 2 Bias-recent Pri
tations, conference publications, and government publications. It is relatively Campbell & Valentine 3 1 Bias-combo Pri
comprehensive back to about 1970. Approximately 60% of the indexed publi- Chaloner 7 3 Bias-fossil Ind
cations originate from outside the United States. The keyword "taphonomy" Chamberlain 4 4 Structure Ind
was given for the search, with the specified period from 1975 to 1983. Chamberlain & Wester 2 4 Structure Inc
The search resulted in 546 tides. After careful screening for duplications, dates Chamberlain et al. 7 4 Process-biological Pri
from 1975 to 1983, and elimination of obvious misfits, about 440 of these were Cheetham & Thomsen 7 3 Structure Ind
used for our analysis. These do not include some (such as a number of articles Cloud 2 4 Bias-fossil Ind
in Paleobiology) that are known to us as important taphonomic publications. Crick 7 2 Bias-fossil Ind
Most omissions appear to be due to problems in coding particular subjects as Damuth 8 4 Bias-combo Pri
taphonomy. For instance, studies of preservation potential in modem environ- Dodson 6 1 General review Pri
ments often were not picked up by the search unless their relevance to paleo- Dodson & Wexlar 5 3 Process-biological Pri
ecology was noted in the title. The total cataloguing for 1983 also was not Dodson et al. 6 2 Bias-fossil Pri
available from Georef at the time of the search. Thus, the 440+ titles represent Dudley & Vermeij 4 4 Process-biological Ind
only part of the total output from taphonomic research over the last nine years. Eldredge 2 2 Bias-fossil Ind
Nevertheless, we feel this reflects a generally valid overview of activities in the Erben et al. 5 4 Structure Inc
field based on literature that is available to the scientific community at large. Farlow 9 3 Bias-fossil Inc
Country of publication is given for by Georef for most titles back to 1975. Finney 5 1 Structure Inc
Doctoral dissertations, which do not have this information, were mostly from Fisher 7 2 Process-biological Pri
U.S. universities (probably from Dissertation Abstracts). Dissertations do not Fleagle 4 1 Bias-fossil Inc
amount to a large proportion of the 440+ tides, but they do help to increase Flessa 1 2 Bias-combo Ind
the apparent dominance of the United States in taphonomic productivity. Coun- Flessa & Bray 3 4 Bias-fossil Ind
tries represented in the search included Venezuela, China, India, Romania, Flessa & Jablonski 9 4 Bias-fossil Ind
Czechoslovakia, Hungary, Poland, New Zealand, Norway, South Africa, and Flessa & Sepkoski 4 3 Bias-fossil Pri
Japan in addition to the Western European block, Canada, and the United Gingerich 7 4 Bias-fossil Inc
States. Gould & Calloway 6 4 Bias-fossil Ind
Gould et al. 3 1 Bias-fossil Ind
Grayson 4 1 Bias-fossil Pri
Hallam 8 4 Bias-fossil Inc
Appendix 2. Hallam 4 1 Bias-fossil Inc
Hallam 3 1 Bias-fossil Inc
Results of a review of Paleobiology Hayami 4 3 Bias-fossil Inc
In terms of activity in different fields, invertebrate paleontology accounted for Haynes 6 3 Process-biological Pri
41% of the 150 articles compared with vertebrate paleontology's 22%. Of those Haynes 9 2 Process-biological Pri
40 primarily concerned with taphonomy, however, the percentages were reversed, Hill 5 3 Process-combo Pri
IV with 23% and VP with 41%. Either way, this contrasts with the pattern Hoffman 5 4 Bias-fossil Ind
presented by the Georef database. Also of interest is the large number of papers Holtzman 5 2 Bias-combo Pri
involving combinations of fields, for example, those dealing with such topics as Hopson & Radinsky 6 3 Bias-fossil Inc
differential preservation and ecological diversity (Lasker 1976) and biases in Hulbert 8 2 Bias-fossil Ind
Phanerozoic species sampling (Signor 1978). Many of these are mainly concerned Jablonski 6 4 Bias-fossil Pri
with marine invertebrates, but they also include other major groups of organisms Jones 6 3 Structure Ind
and reflect the more interdisciplinary orientation of articles published in Paleo- Kanie et al. 6 1 Structure Inc
biology. A comparison of the first five and most recent four years of Paleobiology Kaufman 7 4 Process-biological Inc
showed a relative increase in articles concerned with vertebrate taphonomy. Kellogg 9 4 Bias-fossil Inc
Articles were categorized according to the type of problem or information Kier 3 2 Bias-fossil Pri
which was relevant to taphonomy. The categories were as follows: Kippel & Parmalee 8 4 Bias-fossil Ind
1) Structural attributes of organisms influencing preservation Kitchell et al. 4 2 Bias-recent Inc
2) Post mortem proceses (including agencies of death) Kitchell et al. 7 4 Process-biological Ind
—biological Klein 8 2 Process-biological Pri
—physical: primarily sedimentological Klein 8 2 General review Pri
—chemical Klein & Cruz-Uribe 9 1 Structure Ind
—combination of the above Kobluk et al. 4 2 Process-biological Ind
3) Patterns of preservation and bias (usually case studies) Koch 4 3 Bias-combo Pri
—based on knowledge of the fossil record Koch 6 2 Bias-fossil Ind
—based on studies of recent analogues Koch & Sohl 9 1 Bias-fossil Pri
—based on models of natural systems Kohlberger 9 1 Bias-fossil Ind
—pertaining to a combination of the above Kranz 3 4 Bias-modeled Pri
4) General: discussions of taphonomy as a field, etc.

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X
 TAPHONOMY AND PALEOBIOLOGY 119

Author Vol. Num. Category Status Author Vol. Num. Category Status

Krohn 5 2 Bias-combo Ind Seilacher 5 3 Structure Inc

Kurten 9 1 Bias-fossil Ind Sepkoski 4 3 Bias-fossil Ind
LaBarbera 7 4 Bias-combo Ind Sepkoski 1 4 Bias-fossil Pri
LaBarbera 3 3 Structure Inc Sepkoski 7 1 Bias-fossil Ind
Lasker 4 2 Bias-modeled Pri Sepkoski 5 3 Bias-fossil Ind
Lasker 2 1 Bias-combo Pri Sepkoski 2 4 Bias-fossil Ind
Levinton & Bambach 1 1 Bias-combo Ind Sheehan & Raup 3 3 Bias-combo Pri
Liddell & Brett 8 1 Process-biological Ind Shipman & Walker 6 4 Process-biological Pri
Lindberg & Kellogg 8 4 Process-biological Pri Signor 4 4 Bias-combo Pri
Lipps 7 2 General review Ind Stanley 7 3 Structure Ind
Malmgren et al. 9 4 Bias-fossil Inc Stearn 8 3 Bias-combo Ind
Maynard 2 2 Bias-recent Ind Strathmann 7 3 Structure Inc
Nichols & Pollack 9 2 Bias-modeled Ind Thayer 1 1 Structure Pri
Nicklas 7 1 Process-chemical Ind Thayer 1 4 Structure Inc
Noe-Nygaard 3 2 Process-biological Pri Thayer 3 1 Bias-combo Ind
Oliver 6 2 Structure Ind Thomas & Foin 8 1 Bias-modeled Ind
Paine 9 1 General review Ind Thomsen 3 4 Bias-fossil Ind
Peterson 9 4 Process-combo Ind Tipper 6 1 Bias-modeled Ind
Petry 8 1 Bias-recent Ind Valentine et al. 4 1 Bias-combo Ind
Pielou 5 4 Bias-combo Inc Vermeij 3 3 Bias-fossil Ind
Radinsky 8 3 Process-biological Ind Vermeij et al. 6 3 Process-biological Ind
Raup 1 4 Bias-combo Pri Walker & Alberstadt 1 3 Bias-fossil Ind
Raup 8 1 Bias-fossil Inc Walker & Parker 2 3 Bias-fossil Ind
Raup 9 2 Bias-fossil Inc Ward 7 1 Structure Inc
Raup 2 4 Bias-fossil Ind Ward 6 3 Structure Ind
Raup & Crick 8 2 Bias-fossil Ind Ward & Signor 9 2 Bias-fossil Inc
Raup & Marshall 6 1 Bias-fossil Ind Ward et al. 3 4 Structure Inc
Richardson & Watson 1 4 Structure Inc Watkins & Hurst 3 2 Bias-fossil Ind
Ronan 3 4 Process-biological Ind Weaver & Chamberlain 2 1 Structure Pri
Sambol & Finks 3 1 Bias-fossil Ind Webb 2 3 Bias-fossil Ind
Saunders & Wehman 3 1 Structure Ind Weber et al. 1 2 Process-chemical Inc
Schindel 8 4 Bias-fossil Ind Weiner 6 1 Process-chemical Ind
Schindel 6 4 Bias-fossil Ind Weiner et al. 5 2 Process-chemical Ind
Schindel & Gould 3 3 Bias-fossil Inc Westbroek et al. 5 2 Process-chemical Ind
Schopf 4 3 Bias-recent Pri Westerbroek 9 2 Process-combo Ind
Schopf 7 2 Bias-fossil Ind Westermann & Ward 6 1 Structure Inc
Schopf 6 4 Bias-fossil Ind Wise & Schopf 7 3 Bias-fossil Ind
Schopf 5 3 Bias-fossil Ind Wolff 1 2 Bias-fossil Pri
Schopf et al. 6 4 Structure Ind Wolff 7 2 General review Pri

Downloaded from https://www.cambridge.org/core. Stockholm University Library, on 06 Nov 2017 at 06:32:35, subject to the Cambridge Core terms of use,
available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S009483730001143X