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KIDWELL S.M. 1998. Time-averaging in the marine fossil record: overview of strategies and uncertainties.
[D~termination de la dur~e d'accumulation dans le registre fossile marin: revue des strategies et des incertitudes].
GEOBIOS, 30, 7: 977-995. Villeurbanne, le 31.03.1998.

Manuscrit d~pos~ le 25.08.1997.

ABSTRACT - The paleontologic reasoning that led to the recognition of time-averaged assemblages in the early
1970s, and to elaborations of the concept in the 1980s and 1990s, has now undergone considerable actualistic tes-
ting, primarily using marine mollusks. These studies have confirmed the fundamental elements of the concept,
including paleontologic estimates of the absolute durations and short-term dynamics of time-averaging: despite the
rapid rates of shell destruction that can be documented in modern environments, death assemblages in nearshore
and shelf settings are age-mixtures that have commonly formed over thousands to tens of thousands of years.
Although many more actualistic studies are needed to document environmental and taxonomic variation, the results
so far greatly increase our confidence in the indirect sedimentologic, stratigraphic, and paleontologic evidence used
to evaluate assemblages in the older fossil record. The strategy outlined for evaluating fossil assemblages stresses
breadth of evidence, most particularly geological context and origins of the host interval. Numerous questions
remain. One is whether a "taphonomic clock" operates: does damage, either on an individual shell or for an assem-
blage overall, accrue at a sufficiently steady rate that it can be used to estimate the relative age of a shell (i.e., if it
is from one of the earlier or from one of the later generations to be incorporated in the time-averaged assemblage),
or used to estimate the duration of time-averaging in the overall assemblage (maximum range of shell ages)?
Unfortunately, damage accrues very quickly apparently within molluscan assemblages (e.g. within the first few hun-
dred years), so that assemblages that accumulate over much longer time scales (hundreds of thousands of years) do
not differ significantly in taphonomic grade than those formed over a few thousand years. Comparative analysis of
fossil assemblages suggests that other features, such as diagenetic heterogeneity and microstratigraphic complexi-
ty within host intervals, may be more useful "clocks" for ranking the time-resolution of assemblages, but these ideas
need wider testing. Other uncertainties and directions for investigation are probable differences in the relative
degrees and absolute durations of time-averaging (a) among major taxonomic groups, (b) among environments
within single embayments, (c) through depositional sequences, owing to shifting rates of sedimentation as well as
migrating environments, and (d) over the course of Phanerozoic time, owing to changes in the nature of skeletons
produced as well as in the organisms that influence post-mortem preservation. We have testable hypotheses and
even preliminary evidence for these larger scale patterns, and they are exciting avenues for future research.


Rl~SUM]~ - Le raisonnement pal~ontologique qui a conduit ~ la reconnaissance de la dur~e d'accumulation des assem-
blages au d6but des ann6es 1970 puis aux 51aborations du concept dans les ann~es 1980 et 1990, a d6sormais subi un
test actualiste important en utilisant principalement les mollusques marins. Ces ~tudes ont confirm~ les ~l~ments fon-
damentaux du concept y compris les estimations pal6ontologiques des dur~es absolues et la dynamique ~ court terme
de l'~valuation de la dur~e d'accumulation malgr~ les taux rapides de destruction des coquilles qui peuvent ~tre
observes dans les environnements actuels, les assemblages des sites littoraux et de plate-forme sont, quant ~ leur ~ge,
des m~langes qui se sont commun~ment constitu~s sur des milliers ou des dizaines de milliers d'ann~es. Bien que des
analyses actualistes plus nombreuses soient n~cessaires pour documenter la variation environnementale et taxino-
mique, les r~sultats acquis accroissent fortement notre confiance dans les ~l~ments indirects, s6dimentologiques, stra-
tigraphiques et pal6ontologiques utilisSs pour appr~cier les assemblages dans le registre fossile plus ancien.Les
strategies esquiss6es pour l'interpr6tation des assemblages fossiles font ressortir l'ampleur des preuves plus particu-
li~rement pour le contexte g~ologique et les origines de l'intervalle concernS. De nombreuses questions demeurent.
L'une est de savoir si une "horloge taphonomique" intervient: les dommages, soit d'une coquille particuli~re, soit d'un
assemblage tout entier, parviennent-ils avec des taux snffisamment r6guliers ~ contribuer ~ estimer l'fige relatif d'une
coquille (c'est-~-dire si elle est de l'une des premieres ou de l'une des derni~res g~n~rations qui ont ~t~ incor-

por~es dans l'assemblage dont la dur~e d'accumulation est estim~e), ou peuvent-ils ~tre utilis~s pour ~valuer la
dur~e d'accumulation de l'assemblage tout entier (r~partition maximale des ages des coquilles)? Malheureusement,
les dommages se r~alisent tr~s rapidement dans les assemblages de mollusques (par exemple dans les tout premiers
cent ans) et ainsi les assemblages qui s'accumulent sur des p~riodes beaucoup longues (centaines de milliers
d'ann~es) ne different pas significativement de grade taphonomique de ceux form,s en quelques milliers d'ann~es.
Une analyse comparative des assemblages fossiles sugg~re que d'autres faits, comme l'h~t~rog~n~it~ diag~n6tique
et la complexit~ microstratigraphique dans les intervalles concern~s, peuvent ~tre des "horloges" plus utiles pour le
classement de la r~solution temporelle des assemblages, mais ces idles exigent d'etre plus largement test~es.
D'autres incertitudes et voles d'investigations concernent les differences probables dans les degr~s relatifs et les
temps absolus de la dur~e d'accumulation (a) parmi les groupes taxinomiques majeurs, (b) parmi les environne-
ments de lieux proteges, (c) ~ travers des s~quences de d~p6t en raison des taux variables de s~dimentation et des
environnements migrants et (d) au cours du Phan~rozo~que ~ cause des changements intervenus dans la nature des
squelettes ~labor~s et dans celle des organismes qui influent sur la preservation post-mortem. Nous avons des hypo-
theses testables et m~me une preuve pr~liminaire pour ces patrons ~ plus grande ~chelle et ce sont l~ des voies sti-
mulantes pour des recherches futures.

INTRODUCTION are there large-scale patterns in time-resolution -

do analytically important differences in resolution
How much time is represented by an assemblage exist among taxa, among environments, through
of fossils from a single bed or other narrowly deft- depositional sequences (transgressive-regressive
ned stratigraphic interval? Does the set of co- cycles), across paleolatitudes, or over the evolutio-
occurring fossils r e p r e s e n t an i n s t a n t a n e o u s nary sweep of Phanerozoic time? These questions
"snapshot" of formerly coexisting organisms, or of possible taphonomic megabias are as important
the remains of multiple generations, community for classical paleontologic investigations of single
states, or evolutionarily distinct faunas? The abi- beds and outcrops as they are for broad biogeogra-
lity to answer these questions for individual fossil phic and evolutionary studies, because all rely on
assemblages, both in relative and absolute time, is the fundamental reliability and comparability of
important to virtually all paleobiologic and histo- individual fossil assemblages.
rical geologic studies, because collections of fossils
from single beds or sets of associated beds are the
f u n d a m e n t a l sampling increment for paleoecolo- WHAT IS T I M E - A V E R A G I N G ?
gic, morphometric/taxonomic, and species-level
biostratigraphic and evolutionary studies. PALEONTOLOGIC REASONING
This paper briefly discusses how scales of time- "Time-averaging" is the process by which organic
averaging are evaluated in shallow marine macro- remains from different time intervals come to be
benthic records, using the well-studied mollusks preserved together. More precisely, following
as a primary source. Actualistic studies of modern Walker and Bambach's (1971) original definition,
death assemblages are corroborating the basic it refers to situations where remains "accumulate
paleontologic t e n e t s of time-averaging for from the local living community during the [rela-
example, t h a t mixed-age assemblages exist, and tively long] time required to deposit the contai-
t h a t the oldest generations represented in an ning sediment", and is the result of biological
assemblage leave a weaker signal t h a n the youn- generation times typically being short relative to
gest - and are also corroborating paleontologic net rates of sediment accumulation, or of mixing
estimates of absolute duration of time-averaging depths in the sedimentary column being large
within habitats, thus increasing our confidence in relative to sediment accumulation. During time-
paleontologic and geologic reasoning. This latter averaging, biological remains m a y undergo repea-
point is especially important for our confidence in ted burial/exhumation cycles due to physical
evaluating time resolution in the older fossil reworking of host sediment, and m a y be mixed
record, particularly for taxonomic groups and sedi- over tens of centimeters to a meter or more within
m e n t a r y environments without modern analogs. the sedimentary column by bioturbation. The
Many uncertainties remain, however, even for input of new dead material m a y be continuous or
relatively well-studied mollusks. For example, is discontinuous over this period, and each cohort of
there a taphonomic clock - t h a t is, can the condi- dead material is subject to taphonomic culling as
tion of surviving fossils be used to gauge the a function of tissue type, time in the system, and
degree of time-averaging in an assemblage? Also, post-mortem environmental conditions. Moreover,

the composition of dead input m a y itself change, stress, just as seen in many modern examples
owing to small-scale fluctuations in state of the (Brett & Seilacher 1991). The resulting assembla-
local source community or changes in the envi- ge is a temporally high-resolution "snapshot" that
ronment. The final biomass of a time-averaged may nonetheless be biologically or taphonomically
assemblage is thus expected to be completely dif- skewed, rather than a complete or random sub-
ferent from the instanteous pictures of communi- sample of the source community.
ty structure provided by biological grab samples In absolute terms, the time represented by a cen-
or mass kills. Time-averaged assemblages will sus assemblage could be truly instantaneous (days,
also be fundamentally different from assemblages weeks, single season of year), but in some instances
composed entirely or partially of exotic (allochtho- could range up to several years owing to the pro-
nous) elements. longed nature of some mass mortalities and the
Based on the geological context of fossil assem- long windows for carcass preservation afforded by
blages, paleontologists have recognized that the some low-temperature or low-oxygen seafloors.
general phenomenon of time-averaging can opera- Within-habitat t i m e - a v e r a g e d a s s e m b l a g e s
te over a broad range of time-scales, such that (strict s e n s e of Walker & B a m b a c h 1971).
organic remains from successive generations, suc- These are time-averaged over a period of relative
cessive communities, and even successive chrono- environmental stability, so that only individuals
zones can become amalgamated within a single from a single temporally persistent community
bed or narrow stratigraphic interval (F~irsich are mixed. Short-term processes that mix succes-
1978). For the practical purpose of grouping sive generations and community states include
assemblages with biologically and taphonomically bioturbation (both diffusive and advective move-
comparable degrees of time-resolution, and of dis- ment of remains within the sedimentary column;
tinguishing non-comparable assemblages, the this is inferred for homogeneously amalgamated
spectrum of time-averaging can be subdivided beds, usually with pervasive burrow-mottling)
into four qualitatively distinct categories (Kidwell and reworking (exhumation and redeposition of
& Bosence 1991), as follows. remains, associated with a fluctuating seafloor
C e n s u s a s s e m b l a g e s (original term of Hal- position; inferred for lithologically heterogeneous
lam 1972). These reflect zero or minimal time- intervals, commonly with traces of anastomosing
averaging of individuals, such as when mass discontinuity surfaces or r e m n a n t sedimentary
death of all or some part of a community is follo- structures). The lower the long-term (net or back-
wed by p e r m a n e n t burial, sealing the assemblage ground) rate of sediment accumulation on which
from later additions or modification. These are the these processes are superimposed, and/or the dee-
closest paleontologic analogs of ecological grab per the mixing/reworking depth, then the larger
samples or censuses of living communities in the number of generations that have potential to
m o d e r n environments, and p r e s u m a b l y also be mixed in the final assemblage.
record the shortest periods of time. " S n a p s h o t " is Hypothetically, we can visualize situations where
an excellent alternative term. remains are added to the assemblage continuously
Although these assemblages have the highest time over the interval of time-averaging; such records
resolution, they are not necessarily unbiased pic- are relatively complete with respect to elapsed time
tures of the source community. For example, esca- (that is, most sub-increments of elapsed time are
pe traces and missing life styles in a fossil assem- represented by fossils in the final assemblage; =
blage (e.g., absence of vagile benthos and nekton) " c o n t i n u o u s time-averaging" of F~irsich &
would suggest that some members of the commu- Aberhan 1990). Alternatively, remains may be
nity are missing in the final record; if mineralized added and/or preserved only sporadically, for
hardparts are not accompanied by soft-tissues, example associated with episodic events of mass
then organisms composed exclusively of soft tis- mortality or storm reworking, resulting in a tempo-
sues are also absent and taphonomic omission is rally less complete assemblage ("discontinuous
suspected. However, groups may be missing becau- time-averaging" of F~irsich & Aberhan 1990).
se of the ecology of death itself. Recent "mass mor- Several independent lines of evidence suggest
talities" are rarely fatal to all species and age- within-habitat time-averaged assemblages form
classes, much less to every living individual. over a range of absolute time-scales. These scales
Moreover, the peculiar taxonomic compositions of vary among environments but are on the order of
some fossil censuses suggest that, as in modern a minimum of years and decades to a m a x i m u m of
habitats, many ancient mass mortalities occurred thousands or perhaps ten thousand years depen-
during unusual rather than "everyday" conditions ding on the environment (Fig. 1). Indirect eviden-
in the life of a community, for example coinciding ce used to reach these estimates (Kidwell &
with spawning or culminating a long period of Bosence 1991) includes:

Biostratigraphically Condensed d, ecological information on the minimum num-

ber of years of replicate sampling necessary for
slopes& deepbasins cumulative live diversity to resemble the diversi-
Condensed shelves
ty of the local death assemblage. This is on the
order of years for temperate coastal settings and
Assemblages ~ coastalsettings m a n y decades to a century for the open shelf.
Within-Habitat ~ slopes& deepbasins Environmentally condensed assemblages
Time-Averaged shelves (original term of Ffirsich 1975). These are
Assemblages ~ coastalsettings time-averaged over a period of environmental
change, so that ecologically unrelated species
Census Assemblages become mixed. (The short-term processes and
I I I I I I I I I I l
issues of temporal completeness t h a t are of
1day 1month1year lOyr lOOyr lkylOkylOOkylMylOMylOOMy concern in within-habitat time-averaging also
years of time-averaging per assemblage apply here.) Deciding how large an environmental
change is necessary to categorize the assemblage
as "environmentally condensed" depends on the
FIGURE 1 - Estimated absolute durations for different relative
scales of time-averaging in marine assemblages, inferred from scale of the paleobiologic question.
dated biostratigraphic zones and estimates of average species'
durations, modern and ancient rates of facies migration, and As an example of a very fine-scale definition,
data from biological and hydrographic studies in modern envi- ~iirsich & Kauffman (1984) were able to distingui-
ronments (from Kidwell & Bosence 1991). The lines indicate Sh seasonal salinity-controlled differences in fau=
the range of possible time-scales over which a particular hal composition in some finely bedded Cretaceous
degree of time-averaging might develop. Durdes absolues
estimdes pour diffdrentes dchelles relatives de l~valuation de la lagoon deposits: beds that contained mixtures of
durge d'accumulation d'assembIages marins, infdrdes des zones both low-salinity and high-salinity faunas were
stratigraphiques datdes et des estimations de dur4e moyenne categorized as environmentally condensed. In
des esp~ces, des taux actuels et anciens de migration des facies many lagoonal and other records, however, com-
et des donnges fournies par les dtudes biologiques et hydrogra-
phiques des environnements actuels (d'apr~s Kidwell & munities do not alternate so starkly on a seasonal
Bosence 1991). Les traits indiquent l'extension des dchelles de basis: faunal variation may be a more complex and
temps possibles durant lesquelles pourrait se ddvelopper un intergradational spatial and temporal mosaic, and
degrg particulier de la durde d'accumulation estimg. thus the definition of a community or habitat is
broader. In these circumstances, environmental
a, hydrographic data on the minimum period over condensation is recognizable only at a coarser,
which storms and other physical perturbations facies scale (e.g. subtidal lagoon versus lagoon
become a steady-state process in Recent environ- margin, or lagoon versus open shelf, rather than
ments. This is the set of ordinary conditions pre- season to season within the subtidal lagoon). Also,
ferred or survived by the resident benthic com- the question at hand might only require a coarser
munity, and thus indicates the minimum period of subdivision of environmental variability, for
time-averaging for recurrent fossil assemblages; example if the aim were to compare the biodiversi-
this is on the order of a few decades in coastal set- ty of Cretaceous lagoons with those of the
tings b u t on the order of a century for the open Paleogene, or of tropical lagoons and temperate
shelf; the lifespan of the longest-lived organisms lagoons. Kidwell & Bosence (1991) suggested limi-
in the target community provides another means ting the term environmental condensation to these
of estimating these steady-state periods; coarser environmental changes that would normal-
b, geological information on rates of habitat ly yield distinct lithofacies (as opposed to distinct
migration and facies change during Cenozoic laminae within a facies). The question at hand,
t r a n s g r e s s i v e - r e g r e s s i v e cycles. This includes however, should determine the operational defini-
rates calibrated radiometrically and biostratigra- tion adopted in any given study.
phically; the longer the period that an environ- In terms of absolute time, environmentally conden-
m e n t remains stable in space, then the longer the sed assemblages by definition fall at the high end
m a x i m u m possible duration of within-habitat of within-habitat time-averaged assemblages,
time-averaging (on the order of a few thousands of since environments must have time to change for
years m a x i m u m for shelf facies); different communities to be telescoped at a single
c, ecological information on the minimum number site. For fine-scale condensation sensu Ffirsich &
of years of replicate sampling of a living commu- Kauffman (1984), two seasons could be sufficient to
nity t h a t is required for the cumulative diversity generate an environmentally condensed assembla-
curve to plateau. Several decades to a century are ge, whereas for facies-scale condensation sensu
needed to sample molluscan communities adequa- Kidwell & Bosence (1991), individual assemblages
tely in t e m p e r a t e coastal and shelf settings; and, probably signify at least multiple decades even in

coastal settings where facies patches are small and associated fossils). Reconstructing the short-term
can be quite mobile. Thousands of years are proba- dynamics of condensation commonly depends
bly more typical minimum times on open shelves upon such sedimentologic and taphonomic evi-
and deeper basin settings, where environmental dence, because biostratigraphic evidence alone is
bands are generally broader and driven by slower often not adequate to distinguish between inter-
processes of climate change and transgression/ vals condensed via episodic winnowing or erosio-
regression. Millenial-scale climatic "flickers" in the hal truncation, and those condensed via a slow-
Pleistocene, for example, drove latitudinal shifts in down in pelagic rain, even in highly resolved
species geographic ranges within a single sealevel Cenozoic successions (e.g., Aubrey 1995).
highstand, so that species from two different bio- In absolute terms, biostratigraphically condensed
geographic provinces can be time-averaged (envi- assemblages require evolutionarily long periods
ronmentally condensed) within a single transgres- of time for formation. This might comprise at
sive shellbed (Roy et al. 1996). minimum a few hundreds of thousands of years
Biostratigraphically c o n d e n s e d assem- for geological records characterized by rapidly
blages (sensu Heim 1934). These are time-ave- evolving or diverse groups with very short bio-
raged over geologically long periods of time; spe- zones, and several million y e a r s for records
cies from successive chronozones are mixed containing slowly evolving groups.
within a single horizon or occur in very close stra-
tigraphic succession. Such assemblages a r e com-
m o n l y e n v i r o n m e n t a l l y condensed as well. D E T E R M I N I N G THE D U R A T I O N A N D
Examples are known from both shallow and deep
water settings, and m a y reflect any combination D Y N A M I C S OF TIME-AVERAGING IN
of sediment winnowing, sediment bypass, and MODERN DEATH ASSEMBLAGES
sediment starvation (Jenkyns 1971; Wendt &
Aigner 1985; Kidwell 1991a,b; Fels & Seyfried In modern death assemblages, the duration of time-
1993; GSmez & Fern~ndez-LSpez 1994). averaging - that is the difference between the time
of death of the oldest individual in the assemblage
Condensed deposits are idealized as relatively and the time of death of the youngest - can be deter-
complete records of prolonged low net sedimenta- mined by direct dating of shells. Both radiocarbon
tion, but examples vary in completeness (e.g., some (for deposits younger than ~40,000 years) and
subzones from the normal series may be unrepre- amino-acid racemization (for deposits as old as 1.5
sented; this is a variant of discontinuous time-ave- million years in mid-latitudes) are appropriate (see
raging sensu Fiirsich & Aberhan 1990), in total Flessa 1993 for this specific application).
duration (number of zones condensed), and in the
degree of stratigraphic thinning (expressed either Few studies have addressed this problem directly,
as a percentage of normal section thickness, or as a but these yield consistent results for bivalves
rock accumlation rate, generally <1 m per my). In (Wehmiller et al. 1988, 1995; Nelson et al. 1988;
some instances, the input or preservation of fossils Flessa et al. 1993; Martin et al. 1996; Meldahl et al.
is so discontinuous that there is a great discrepan- 1997): shell ages range up to several thousand
cy in the ages of mixed fossils. years in actively forming death assemblages on
beaches, tidal flats, and nearshore subtidal habi-
The end-member situation where the assemblage tats (<10 m), and can be several thousands to a few
is composed entirely of fossils exhumed from tens of thousands of years old on the open shelf,
much older strata is best categorized as a remani~ where conditions of low net sedimentation among
or lag assemblage, b u t all gradations exist in the other factors promote thin shelly records of marine
quantities of remani~ that m a y be incorporated transgression. A compilation of 734 published
into a younger fossil assemblage, and in the age- radiocarbon dates from 276 localities produces
discrepancy between the remani~ and younger similar patterns: median maximum ages of 1,250
fossils. Assemblages condensed (telescoped) via years in nearshore localities (<10 m depth) and
r e p e a t e d winnowing and erosional truncation 9,190 years in shelf localities (Flessa &
m a y in fact be difficult to distinguish from those Kowalewski 1994) (Fig. 2). These constitute esti-
mixed with remani~, because some degree of dia- mates for within-habitat time-averaging in behch
genetic modification and stabilization is practical- and nearshore settings and for within-habitat
ly axiomatic for the oldest cohorts to survive time- time-averaged and environmental condensed
averaging via winnowing over such long time assemblages on the open shelf, and indicate that
scales (e.g., Wendt & Aigner 1985; Machalski & such long periods of time-averaging are typical
Walaszczyk 1987; for this reason, GSmez & rather than unusual for modern molluscan assem-
Fern~ndez-LSpez 1994 refer to all reworked speci- blages. These coincide with estimates using inde-
mens as remani~, even if they are the same age as pendent indirect methods (described above, Fig. 1),


30t m31

A ==
N = 63
o" U.
p- O
500 1000 1500 2000 2500 3Q00
9 U. 10 11

1 1 1 2

6,000 12,000 18,000 24,000 30,000 36,000 >36,000 6,000 12,000 18,000 24,000 30,000 36,000 >36,000
Maximum age (uncorrected radiocarbon age in years BP) Maximum age (uncorrected radiocarbon age in years BP)

FIGURE 2 - Frequency distribution of the m a x i m u m ages of shells in actively accumulating molluscan d e a t h assemblages in (A)
n e a r s h o r e localities (median m a x i m u m age = 1,250 years) and (B) shelf localities (median m a x i m u m age = 9,190 years), based on
a review of t h e radiocarbon literature (reprinted from Flessa & Kowalewski 1994). N u m b e r above b a r indicates absolute frequen-
cy in t h a t age class. These data document the durability and potential of shells to be time-averaged over t h o u s a n d s to a few tens
of t h o u s a n d s of years, and confirm the indirect reasoning used to e~timate absolute scales of time-averaging in the fossil record
(compare w i t h age estimates for w i t h i n - h a b i t a t time-averaging and ~nvironmental condensation in the same e n v i r o n m e n t s in Fig.
1). Distribution de frdquence des ~tges m a x i m a des coquilles dans des assemblages post-mortem d'accumulation active de mollusques
(A) dans des sites littoraux (etge mddian m a x i m u m = 1 250 ans) et (B) dans des sites de plate-forme ((tge mgdian m a x i m u m = 9 190
ans), basge sur une revue de la littdrature du radiocarbone (repris de Flessa & Kowalewski 1994). Le nombre au-dessus de chaque
barre indique la frdquence absolue dans chaque classe d~ge. Ces donndes montrent la durde possible et la capacitd des coquilles
d'avoir une durde d'accumulation de quelques milliers & quelques dizaines de milliers d'annges, elles confirment le raisonnement
indirect utilisd pour dvaluer des dchelles absolues d'estimation de la durde d'accumulation dans le registre fossile (& comparer avec
les dvaluations d'~ge pour l'estimation de la durde d'accumulation dans un habitat et & la condensation environnementale dans les
m~mes milieux de la Fig. 1).

and thus confirm the reliability of paleontolo- contradict observations and inferences of rapid
gic/geologic reasoning for the fossil record (Flessa shell destruction in modern environments (e.g.
& Kowalewski 1994; Meldahl et al. 1997). Peterson 1976; Cummins et al. 1986; Davies et al.
The strongly right-skewed frequency distributions 1989), only that those high rates do not remove all
for mollusk shells from modern death assemblages shell material from the record before it reaches a
("hollow curves" in Fig. 2) indicate that, although depth of permanent burial. Moreover, the very
assemblages contain some very old shells, most short "half-lives" calculated by Cummins et al.
dated shells are quite young (a few hundred years (1986) for mollusk shells in Texas lagoons (e.g. 40-
or less). This does not prove but is certainly consis- 307 days for half of all shell input to have disap-
tent with the idea that individual cohorts of dead peared) applied only to very small specimens (adult
shells "fade"in numbers duringthe period of time- and post-larval shells less than a few mm long);
averaging, so that the final assemblage sampled by half-lives were "immeasurably long" for all others,
the paleontologist is dominated by shells of relati- and these surviving elements are those dated in
vely young vintage. The quite good agreement of
other studies of death assemblages. The model pre-
species relative abundances in death assemblages
with that in the local live community is also consis- ferred by actualists to explain long periods of time-
tent with this model of how the compositions of averaging of mollusk shells -i.e., that intermittent
assemblages develop over the course of time-avera- burial protects shells from attack for some or much
ging (Kidwell & Flessa 1995). This would be very of the period of time-averaging (Driscoll 1970;
good news for paleobiologists if generally true, Seilacher 1985; Powell et al. 1989; Flessa et al.
because it suggests that time-averaging largely 1993)- is identical to that inferred by paleontolo-
adds a few old shells to mixed assemblages (ghosts gists to explain time-averaged and condensed
of old generations), rather than thoroughly and assemblages in the fossil record (Jenkyns 1971;
equally admixing shells of diverse ages. Wendt & Aigner 1985; Kidwell 1982, 1989; Kidwell
As discussed by Flessa & Kowalewski (1994), the & Aigner 1985), further corroborating the power of
long durations of time-averaging documented by the indirect methods that paleontologists must use
radiocarbon and amino-acid racemization do not to evaluate the pre-Quaternary record.

For many paleontologic studies, being able to eva- unmodified hardparts
luate the relative scale of time-averaging in fossil
assemblages is adequate for the paleobiologic ques-
tion at hand, but in other instances an absolute
I bv,us1
census ?

no - have had
yes: many articulated specimens
life positions not necessary
+ sediment evidence for obrution
estimate of duration is desirable, and both types of taphonomic modification
analysis are discussed here. Absolute estimates
should not be assumed to be superior to strictly

relative rankings. For example, a thousand years of ~
admixture of species that ordinarily
time-averaging in one setting may result in only condensed ? ) yes: have mutually exclusive ranges
within-habitat time-averaging, whereas in another
instance it may involve environmental condensa- no SUGGESTIVE FEATURES
tion. Those two assemblages would present very heterolithic matrix
different kinds of records for paleobiologic analysis. association with discontinuity or
Ienvironmentally flooding surface
R E C O G N I Z I N G RELATIVE SCALES OF TIME- condensed ? 1 yes: fossils with distinctly different
t aphonomic/diagenetic signatures
AVERAGING fauna functionally dissonant with
no matrix
Identifying the relative scale of time-averaging is
largely a process of elimination. It requires mu]- lithologically homogeneous matrix
oo,.t: one style of taphonomic modification
tiple lines of taphonomic, sedimentologic, strati- within-habitat I y e s : & diagenesis
graphic, and paleontologic evidence - no single time-averaged J functional agreement between
type of evidence is usually sufficient (Fig. 3). fauna & matrix

S t e p 1. Test the given assemblage for census attri-

F I G U R E 3 - S t e p s i n i n t e r p r e t i n g t h e r e l a t i v e s c a l e of t i m e - a v e -
butes. Sedimentologic features such as mud-drapes, r a g i n g w i t h i n a f o s s i l a s s e m b l a g e . Etapes dans l'interpr~tation
graded deposition, an unusual concentration or de l'gchelle relative de l'estimation de la durge d'accumulation
diversity of rapidly precipitated authigenic mine- dans un assemblage fossil.
rals, and the restriction of fossils to a single bed-
ding plane are common ancillary lines of evidence wear, diagenetic alteration) is not a consideration
(see Brett & Seilacher 1991; Brett & Baird 1993), in establishing biostratigraphic condensation,
but the primary evidence for mass death and rapid although it may illuminate the conditions under
permanent burial will be taphonomic. Classic lea- which condensation occurred.
tures include a low proportion of broken specimens, Step 3. Most fossil assemblages are neither cen-
a high proportion of articulated specimens, and suses nor biostratigraphically condensed, b u t fall
some or many specimens in life positions. However,
between these extremes. To discriminate within
the absence of specimens in life-position does not
this part of the spectrum, the best strategy in my
necessarily mean that the assemblage is not a cen-
sus-type snapshot. Epifaunal organisms are less experience has been first to test for evidence of
likely to be preserved in life-positions than infaunal environmental condensation. Reasons to suspect
individuals even under the most benign circum- environmental condensation include (adapted
stances, and some censuses are created by en masse from Fiirsich 1978; Kidwell 1993a):
transport of organisms into an exotic environment a, discontinuity surfaces t h a t extend through the
of death and burial, so that none are in life-posi- stratigraphic interval containing the assemblage.
tions (e.g., Burgess Shale). In general, however, These can be present even within very thin
there should be minimal evidence for post-mortem "beds", and can be quite subtle lithologically or
modification of elements other than possible rota- even partly obliterated by burrowing and biotur-
tion or transport, and in some instances there bation (a particular problem in the post-Paleozoic
might be superb preservation of soft-tissues as well record); the host sediment m a y be heterolithic;
as of biomineralized skeletal elements. b, fossils with disparate qualities of preservation,
S t e p 2. If a census origin can be eliminated, then so that the taphonomic character of the total
the next step is to test for the opposite extreme of assemblage would be described as bimodal or
biostratigraphic condensation. By definition, bios- polymodal, rather than simply a range in grades
tratigraphically condensed assemblages contain from specimen to specimen;
taxa t h a t ordinarily have m u t u a l l y exclusive c, up-section variation in the quality of preserva-
c h r o n o s t r a t i g r a p h i c ranges. The taphonomic tion consistent w i t h change (or a series of
condition of the fossils (degree of articulation, changes) in water energy, oxygenation, depositio-

nal tempo, etc. Excluded are simple up-section to be within-habitat time-averaged, i.e. consisting
shifts in fossil condition linked to single-event of multiple generations from a single indigenous
p h e n o m e n a such as graded bedding; community. This is a default category. Positive evi-
d, association of the assemblage with a major dence consistent with, but not proof of, within-
stratigraphic discontinuity. Subsets of fossils m a y habitat time-averaging includes:
be filled with sediment t h a t differs from the sur- a, lithologically homogeneous matrix;
rounding matrix, or m a y have a different style of b, functional agreement between taxa and matrix;
skeletal preservation, indicating reworking from and,
older strata (including strata not otherwise pre- c, single style of preservation among taxa of simi-
served at the discontinuity); lar construction, or continuous range in quality,
e, association with an interval or surface of sharp including diagenesis.
deepening or shallowing, as inferred from litholo-
gic features; and, G e n e r a l C a v e a t s . Several kinds of error will
occur using these guidelines. For example, the
f, ecologically disparate faunas admixed within
most unambiguous census assemblages come
bed. Ecological disparity might be recognized by
from highly stressed environments where both
reference to (i) the ecological tolerances of living
descendants, (ii) functional analysis (e.g., any colonization and death are highly pulsed; black
shales and other intermittently anoxic settings
mixture of soup-, firm-, shell-, and hard-ground
dwelling forms), or (iii) species co-occurrences are good examples. In other settings, however,
t h a t diverge from those t h a t are typical for ste- some true census assemblages will be erroneous-
ly categorized as within-habitat time-averaged,
reotypic biofacies of the same geologic age and
province. In general, this ecological approach is especially where bioturbation might have disrup-
ted and disarticulated specimens w i t h o u t
least desirable because of the potential to become
admixing other generations of dead material. It is
circular. It should be used to corroborate an eva-
difficult to see how this kind of error could be
luation suggested by taphonomic, sedimentologic,
or stratigraphic evidence, r a t h e r t h a n used alone. avoided, but fortunately it is a conservative one.
These four categories subdivide a continuum of
Also suspect are assemblages from stratigraphic
intervals that are demonstrably thinner than coeval time-averaging, and so m a n y intergradations are
strata, for example based on the stratigraphic possible. For example, if a catastrophically censu-
sed community occupied a seafloor t h a t contained
convergence of marker beds or biozone boundaries
any quantity of organic remains from previous
(i.e. stratigraphically condensed intervals). Thin-
generations (i.e., if the census was preceded by a
ning may occur over paleohighs (because of local
period of attritional mortality or a background of
starvation or enhanced winnowing) but also in
exotic input), then the taphonomic signature of
paleolows (siliciclastic starvation or suppressed
the census will be diluted accordingly. The assem-
rates of carbonate production), on bathymetric "pla-
blage is actually a kind of hybrid.
teaus" (bypassing in a toplap situation), and on
slopes (focussing winnowing or bypassing), and thus Some environmentally condensed assemblages will
is not limited to a particular water depth or phase of probably be erroneously categorized as within-
basin development. Environmentally condensed habitat time-averaged, but the confusion should
assemblages are not limited to stratigraphically primarily concern examples where condensation is
condensed intervals, but these are suspect records of a very fine-scale (for example, situations of
(see discussion in section 'Variation through deposi- taphonomic feedback where dead shell input
tional sequences"). influences the living community, or situations
where community states fluctuate rapidly due to
The most difficult situations are where fossils
highly variable environmental conditions, such as
from successive environments have become tho-
common in closed lagoons, small lakes, and silled
roughly admixed, either because of bioturbation basins). The roost practical approach is to take par-
or because of storm or other physical reworking ticular care in evaluating: (a) assemblages from
(burial-exhumation cycles). This homogenization densely packed shellbeds of all scales, since these
obscures or obliterates sedimentologic and strati- commonly record multiple r a t h e r t h a n single
graphic evidence of a complex history of fossil events of shell reworking; (b) assemblages from
accumulation, and in t h a t way also removes the facies of high-stress environments as listed above,
most conservative basis available to the paleonto-
where the environment is recognized from indepen-
logist for dissecting the assemblage into a series dent sedimentologic and stratigraphic evidence;
of higher resolution sub-assemblages. and (c) assemblages that mantle or immediately
S t e p 4. In the absence of evidence for census, underlie stratigraphically significant discontinuity
biostratigraphic condensation, or environmental surfaces (e.g., flooding surfaces, sequence bounda-
condensation, the assemblage would be assumed ries, mid-sequence condensed intervals, etc).

ESTIMATING ABSOLUTE DURATION (Kidwell & Flessa 1993; and see discussion below).
IN FOSSIL ASSEMBLAGES Estimate the time i n h e r e n t in the state of fos-
There are several approaches to estimating the sil preservation. This is a logical approach, since
number years recorded by particular fossil assem- a shell's condition should only worsen with increa-
blages, and all should be attempted for concordance. sing exposure to taphonomic processes after death.
However, attempts to calibrate such a "taphono-
U s e h o r i z o n s of k n o w n age (e.g., biostratigra- mic clock" (Kidwell 1993a) in absolute years are
phic datums, ash beds, sequence boundaries) or yielding mixed results, and thus the method can
the known duration of other phenomena (e.g., ave- not yet be recommended (discussed below).
rage durations of species) to estimate the time
value of the larger stratigraphic interval that EXAMPLES OF ANALYSIS
includes the assemblage of interest. This yields a
m a x i m u m possible duration for time-averaging. Applying these approaches (methods 1-3 above) to
benthic assemblages in the Miocene of Maryland,
E s t i m a t e the time r e q u i r e d to deposit the bed Kidwell (1982) estimated tens to hundreds or a few
that h o s t s the assemblage, reasoning from its thousands of years for assemblages associated with
sedimentology. This is the original reasoning of bedding planes or contained within "minor simple"
Walker & Bambach (1971), and is especially useful shell beds (concentrations smaller in physical
for analyzing the duration of time-averaging dimensions than sparsely fossiliferous facies in the
within dense shell concentrations (Kidwell 1993a; section, and having internally homogeneous or
Brett & Baird 1993). There are several caveats for monotonic change in matrix and biostratinomy).
coquinas - for example storm beds that accumula- These contrasted with "major complex" shell beds
te in days may nonetheless include time-averaged (laterally more extensive than other facies or divi-
shells exhumed from the seafloor - but these are sible into facies, and having complex internal strati-
common sensical. In a few instances, time-avera- graphies), which provided thin (stratigraphically
ging may have operated over a shorter period than condensed) records of transgressive marine facies.
required to deposit the host bed (e.g., infaunal The fossils in major complex deposits clearly recor-
assemblages that penetrate only the upper part of ded more time than a simple time-averaged assem-
a unit, down from a capping discontinuity). But in blage in the sense of Walker & Bambach (1971)
most instances, the time required to accumulate since each major shellbed was environmentally
the host bed is a minimum period for time-avera- condensed, and recorded more time than a single
ging of the fossil assemblage that it contains. "minor simple" shell bed since many of these were
Extrapolate from m o d e r n analogs. The as- amalgamated or stacked within a single major com-
sumption of uniformitarianism is probably accep- plex bed. Each major complex accumulation lay
within a single planktonic foraminiferal zone (-0.5-
table for the younger fossil record (e.g., Cretaceous
1.0 my each) and comprised only part of the total
and Cenozoic): if the sedimentary deposit that
range of most of its molluscan species (average
contains the fossils is known in modern settings to
duration of North American Neogene species is 1-10
be characterized by low time-averaging, then the
my). Each complex shell bed thus must have formed
fossil assemblage might be inferred to have compa-
over a period shorter than -0.5 my, but probably
rable resolution. Absolute durations of time-avera-
longer than 105 years; 10 ~ to 10' years were estima-
ging (see above) might even be extrapolated for
ted. This estimate was corroborated by analogy with
molluscan assemblages of these ages. Some ancient
radiocarbon-dated shell-rich sands that have for-
depositional systems, however, and many extinct
med on modern shelves during the past 18,000
groups, have few or no analogs in the modern
years of marine transgression.
world, and many extant groups of marine orga-
nisms with important fossil records have received Brandt (1985) and Parsons et al. (1988) used simi-
little actualistic examination (e.g., scales of time- lar reasoning for assemblages in Ordovician and
averaging in modern death assemblages of bryo- Devonian strata, producing estimates of tens to
zoans, brachiopods, crustaceans, corals, coralline thousands of years for assemblages ranging from
algae, larger forams). For these, the best strategy at simple to condensed shell beds (and see estimates
present might be to apply the broad estimates of in Brett & Baird 1993).
absolute time in Figure 1, once the relative scale of
time-averaging in the fossil assemblage has been
established (as discussed above). I suspect, howe- IS THERE A TAPHONOMIC CLOCK?
ver, that these absolute estimates will be maxi-
m u m possible durations of time-averaging for We intuitively know that taphonomic damage is a
other taxa: bivalve shells seem to be exceptionally function of time as well as environment after
durable hardparts among marine macrobenthos death, as the condition of a carcass can only dete-

riorate. Shell condition has been used since the A Bahia Concepcion eastern fan-deltas (n = 24 shells)
earliest days of marine taphonomy to distinguish 20.0"
census from non-census assemblages (e.g., >- Grade 1

Johnson 1960), but the extent to which differences O
Z [ ] Grade 2
in the degree of skeletal deterioration can be used LLI 12.0-
to differentiate among the various scales of time- O [ ] Grade 3
I.U 8.0-
averaging is not clear, nor is it clear that shell Lt. Grade 4
condition can be used to calibrate the duration of 4.0-

time-averaging in absolute years. After all, des- 0.0

tructive conditions do not always operate conti-
nuously in a given environment (e.g., seasonality B Bahia Concepcion eastern fan-deltas (n = 24 shells
and longer variation in water energy, bioturba- 20.0"
tion, and corrosive porewaters), nor are individual
>- 16.0- [ ] Grade 1
shells necessarily exposed continuously to des- O
tructive processes (e.g., intermittent burial or Z 12.0- [ ] Grade 2
emersion, d e v e l o p m e n t of protective over- O 8.0- ~ Grade 3
growths). D a m a g e t h u s will not necessarily tZ:
U. 4.0-
accrue in a steady, clock-like m a n n e r after death.
Is there in fact a taphonomic clock for shell age- 0.0-
since-death in modern assemblages, such as might C Bahia Concepcion pocket bays (n = 24 shells)
then be applied to the fossil record? The answer has
consequences for two practical issues: (1) whether 20.0-
[ ] Grade 1
taphonomic damage can be used to reassign indivi- ~ 16.0-
dual shells into their original generational cohorts, [ ] Grade 2
allowing the investigator to "unmix" time-averaged [ ] Grade 3
assemblages; and (2) whether the overall condition 8.0-
Grade 4
of the assemblage (as opposed to that of individual 4.0-
shells) can be used to estimate the absolute scale of
0.0 , , , , , , , , , , , , , , , , , ,
time-averaging for the total assemblage.
These issues have now been tested in a number of
shallow marine settings (Powell & Davies 1990;
Flessa et al. 1993; Wehmiller et al. 1995; Martin et
al. 1996; Meldahl et al. 1997). The types of damage SHELL AGE (calendar years before 1950)
used to rank shell deterioration include disarticula- D
tion, fragmentation, epibiont encrustation, abra- Bahia la Choya tidal channel (n = 13 shells)
sion, corrosion, bioerosion (original criteria of 8.0

Johnson 1960), and loss of color, surface glossiness, >"

[ ] Grade 1
and organic coats (Powell & Davies 1990). These O 6.0-
Z [ ] Grade 2
studies have found only a weak relationship bet- uJ
4.0- [ ] Grade 3
n" Grade 4
FIGURE 4 Frequency distribution of shell ages in actively for-
- LL 2 . 0 -
ming bivalve death assemblages from five intertidal to shallow
subtidal environments in the Gulf of California, documenting 0.0 i i i J i J m i = i i
time-averaging over m a n y hundreds to a few thousands of
years (A-E; from Meldahl et al. 1997). Although the oldest E Bahia la Choya inner tidal flats (n = 17 shells)
shells in each time-averaged assemblage are typically highly 8.0

altered (Grade 4), the youngest shells range in condition from >-
unaltered (Grade 1) to moderately (Grade 3) or high altered. O 6.0-
[ ] Grade 1
Post-mortem damage accrues rapidly but erratically, and thus ~ [ ] Grade 2
is a poor clock of an individual shell's age-since-death.
Distribution de frdquence des (tges des coquilles dans des assem- ~ l!ili.T~,,,~ [ ] Grade 3
blages post-mortem en formation active de bivalves dans cinq
environnements de l'intertidal au subtidal peu profond du Golfe 2.0. i

Grade 4
de Californie, montrant la durde d'accumulation de plusieurs
centaines ~ quelques milliers d'anndes (A-E : d'apr~s Meldahl et 0.0
al. 1997). Quoique les plus anciennes coquilles dans chaque
assemblage de durde d'accumulation gvalude soient typiquement
' "~ 'T "T ,'; "7 ~ ~ ~ ~ ~ ~, (?, ~ ?
tr~s altdrdes (Grade 4), les coquilles les plus rdcentes se rdpartis-
sent d'inaltdrdes (Grade 1) & moddrdment (Grade 3) et haute-
ment altdrdes. Les dommages post-mortem s'accroissent rapide-
ment mais de fa~on stochastique, cela constitue donc une horlo- SHELL AGE (calendar years before 1950)
ge imprdcise de l'dge individuel des coquilles depuis la mort.

clock of shell damage, and intervals of exhumation

Pleistocene ooAO and exposure on the seafloor for individual shells
(~ 125,000yrs) are too erratic over periods of only a few millenia for
successively aged subpopulations of shells to have
2ooc I~ Site 5 Middle Flat acquired distinctive levels of taphonomic damage
u') Site 8 MiddleFlat (op cit., and Seilacher 1985). This is consistent with

t field experiments: shells quickly become damaged
when exposed continuously to destructive processes
on the seafloor, but acquire damage more erratical-
ly and thus slowly when permitted to undergo natu-
ral burial/exhumation cycles (e.g., tethered-shell
~ ] 00(
experiments of Driscoll 1970; Best & Kidwell 1995).
+ Several workers have reported that Pleistocene
{ + ++ shells can be readily distinguished from Recent
shells in modern transgressive beach and near-
shore assemblages on the basis of shell damage,
I I indicating that shell condition can be sensitive to
2 3 the period of time-averaging when shell subpopu-
TaphonomicGrade lations differ in age by m a n y tens of thousands to
hundreds of thousands of years (Frey & Howard
FIGURE 5 - Relationship between shell age and taphonomic 1986; Henderson & Frey 1986; Wehmiller & York
condition for bivalves from three actively forming death
assemblages from intertidal Gulf of California (within a few 1992; Wehmiller et al. 1995; Flessa 1993). Here
h u n d r e d meters of area E of Fig. 4) (data replotted from again, however, the distinction is based on shell
Martin et al. 1996). The youngest shells are a few hundred color (blackening) and/or polishing, which are
years old and are in the best condition (Grade 2-3); older shells unlikely to be preserved in the older stratigraphic
are in comparable or worse condition (Grade 3 or 4) but there
is no consistent deterioration with age, even when Pleistocene record; fragmentation varies among sites depen-
specimens are considered (-125,000 years old). Each shell was ding upon energy levels, and is thus unreliable. In
dated by both amino acid racemization and radiocarbon one instance (intertidal northern Gulf of Califor-
methods. The wide range of ages of shells exhibiting Grade 3 nia), specimens reworked from Pleistocene bea-
damage demonstrate the difficulty in using shell condition to
estimate age-since-death. Relations entre l'dge de la coquille et chrock are characterized by lack of color, chalky
les conditions taphonomiques pour les bivalves d'apr~s trois surface texture, and bits of adhering cemented
assemblages post-mortem se formant activement en zone inter- matrix (Flessa 1993), but nonetheless are not
tidale dans le Golfe de Californie (sur quelques centaines de significantly different in condition from shells
m~tres de l'aire E de la Fig. 4) (donndes reprises de Martin et
al. 1996). Les coquilles les plus rdcentes ont quelques centaines that are only one thousand years old (Martin et
d'annges et sont en bon ~tat de conservation (Grades 2-3); les al. 1996) (Fig. 5). The evidence that a taphonomic
coquilles plus anciennes sont de conservation comparable ou clock operates over these longer time-periods is
plus mauvaise (Grades 3 ou 4) mais il n'y a pas de dgtdriora- thus still weak.
tion en relation avec l'(tge, m~me q u a n d on consid~re des spdci-
mens du Pldistoc~ne (- 125 000 ans). Chaque coquille a dtg Although the relationship between the scale of
dat#e p a r les deux mgthodes de la racdmisation des amino- time-averaging and the condition of any indivi-
acides et du radiocarbone. La large distribution des (tges des
coquilles prdsentant des dommages du Grade 3 souligne la dif- dual shell is weak, the overall condition of an
ficultd d'interprgtation de l'dtat des coquilles pour en estimer assemblage might still degrade consistently with
l'~tge depuis la mort. elapsed time. The overall condition of an assem-
blage can be quantified several ways (average
shell condition, modal shell condition, frequency
ween shell age (time-since-death) and taphonomic distribution of shell condition) and can be based
condition ("grade") within the first few hundreds or on any subset of possible taphonomic features
thousands of years after death. Color and surface (disarticulation, fragmentation, glossiness, etc)
glossiness are the only exceptions (and see Wilson (e.g., Brandt 1989; Davies et al. 1989; Flessa et al.
1988; Van Der Valk 1991), but neither of these is 1993; Kowalewski et al. 1995). Actualistic studies
likely to survive later diagenesis. The oldest shells indicate, however, that, here again, there is no
in an assemblage are generally moderately to high- strong relationship between assemblage grade
ly altered, bat many younger shells may be in equal- and years of time-averaging within the first few
ly poor condition (Figs. 4,5). Thus, using the condi- thousands of years of time-averaging (Flessa
tion of a shell to reconstruct its age relative to 1993; Martin et al. 1996; Meldahl et al. 1997), not-
others in the assemblage is not yet demonstrably withstanding the strong tendency for the oldest
reliable for assemblages time-averaged over periods shells to be in poor condition (Fig. 4). The weak-
as short as a few centuries or millennia (Flessa ness in the relationship might be in part an arte-
1993). The presumption is that burial stops the fact of environmental differences among the

600" during low tides stops the taphonomic clock as

effectively as shallow burial in subtidal settings.
500- E Insights from the stratigraphic record are gene-
rally consistent with this provisional conclusion
about modern molluscan death assemblages: the
400- overall condition of the assemblage seems to be
as strongly or even more strongly linked to the
environment of accumulation (as originally
300- implied by Johnson 1960), and particularly the
short-term burial/exhumation dynamics of shells,
200 - than it is to the duration of time-averaging. The
A most common signature in all types of time-ave-
raged assemblages is a mixture of shells exhibi-
100- ting a full range of physical and biogenic damage.
Kidwell & Aigner (1985), for example, found
shells in all stages ofpost-mortem disintegration,
0 i i I I i including articulated and disarticulated, whole
2.0 2.2 2.4 2.6 2.8 3.0 3.2 and fragmented, pristine and biocorroded, sharp
and abraded, from a series of environmentally
condensed hiatal deposits where complex accu-
Average Taphonomic Grade mulation histories could be inferred on indepen-
Ill dent sedimentologic evidence. This makes sense,
increasing taphonomic alteration given that by definition new material is conti-
nually added to an assemblage during time-ave-
FIGURE 6 - Relationship between scale of time-averaging and raging, so that while early cohorts may have
average taphonomic grade for actively forming bivalve death degraded to taxonomically unidentifiable debris,
assemblages, based on data from Fig. 4 (reprinted from Meldahl the assemblage should include shells in all condi-
et al. 1997). Scale of time-averaging is expressed as shell half- tions up to perfectly fresh (and see same argu-
life, calculated from the curves in Fig. 4. Assemblages A-D are
all from subtidal settings, and, although the regression line is ment in Fiirsich & Flessa 1991 for modern
not significant, a relationship between damage and time-avera- assemblages). The spectrum of damage does not
ging is suggested. Outlying assemblage E shows much less broaden appreciably in biostratigraphically
damage than expected for the duration of time-averaging, but it condensed assemblages, nor is it notably narro-
is from an upper intertidal setting where diurnal emergence of
shells slows taphonomic attack. Relations entre l~chelle d'esti- wer for within-habitat time-averaged assem-
mation de la durde d'accumulation et le grade taphonomique blages, suggesting that this mixed condition
moyen pour les assemblages post-mortem de bivalves se formant develops quickly and persists even through long
activement, d'apr~s les donnges d e l a Fig. 4 (repris de Meldahl et periods of time-averaging (and see Fiirsich &
al. 1997). L~chelle de l'estimation de la durde d'accumulation est
exprimde comme la moitid de la durde de vie de la coqaille, cal- Aberhan 1990; Fiirsich & Oschmann 1993;
culde d'apr~s les courbes de la Fig. 4. Les assemblages A-D pro- Kidwell 1993a). Taphonomic grade based on bios-
viennent tous de sites subtidaux et, quoique la droite de rggres- tratinomic (pre-burial) criteria thus seems
sion ne soit pas significative, une relation entre les dommages et reliable only for distinguishing census assem-
l~valuation de vie moyenne est suggdrde. L'assemblage isold E blages from all other time-averaged assemblages,
montre beaucoup moins de dommages qu'on ne l'attend d'apr~s
la durge d'accumulation, mais il provient d'un site de l'interti- and not for discriminating among time-averaged
dal supgrieur oh l~mersion quotidienne des coquilles rgduit les assemblages formed over different time-scales.
ddgdts taphonomiques.
The stratigraphic record suggests that other
assemblages tested so far. The large age-calibra- taphonomic features do begin to accrue with pro-
ted dataset that Meldahl et al. (1997) are develo- gressive time-averaging, and would thus be useful
ping for intertidal and shallow subtidal assem- for distinguishing among these longer scales of
time-averaging. These features are (a) diagenetic
blages from the subtropical Gulf of California is
heterogeneity in the assemblage itself and (b)
providing important new information on the pos- microstratigraphic complexity in the sedimentary
sible effect of environment on taphonomic clocks interval that hosts the assemblage (Kidwell
(Fig. 6). They have found, for example, that the 1993a; Brett & Baird 1993). For example, conden-
average quality of shell preservation declines sed assemblages commonly contain mixtures of
with increasing duration of time-averaging for a original shell material, variously replaced shells,
series of four lower intertidal to shallow-subtidal and steinkerns. Not all condensed fossil assem-
assemblages, but that an upper intertidal assem- blages have a mixed diagenetic signature: many
blage with far greater time-averaging has very consist entirely of well-preserved aragonitic and
low damage, apparently because emergence calcitic shells. However, I have never encountered

e x t r e m e diagenetic h e t e r o g e n e i t y in skeletal OTHER U N C E R T A I N T I E S A N D

assemblages that lack independent evidence of NEW D I R E C T I O N S
condensation or association with a surface of
significant submarine erosion: for example, such
heterogeneity is not associated with parasequen- SCALES OF TIME-AVERAGING FOR OTHER
ce- and smaller-scale discontinuity surfaces (see MARINE TAXA
also Holland 1993). Diagenetic heterogeneity Actualistic studies have provided a wealth of evi-
appears to be a condition associated with assem- dence that the skeletons of marine macrobenthos
blages in which fossils differ in age by many tens differ significantly in their post-mortem durabili-
of thousands to hundreds of thousands or more ties - owing to such factors as shell size, shape,
years, at least in siliciclastic records. mineralogy, and microarchitecture - and thus we
Fine-scale complexity in the bedding of skeletal can be almost certain that their time-averaged
concentrations ("microstratigraphic complexity") assemblages represent different periods of time.
also appears to be related to the time-scale of Skeletons composed of fragile materials, for
accumulation r a t h e r t h a n to the environment of example, should degrade so rapidly post-mortem
a c c u m u l a t i o n alone (Kidwell 1991a, 1993a). that they might all have disappeared after only a
few seasons or years of time-averaging, so that
Minor discontinuity surfaces that cut through
only the simplest forms and briefest duration of
stratigraphically condensed shell beds typically
within-habitat time-averaging are likely. This, for
are a mixture of types (including scours, burro- example, appears to be the situation for organic-
w e d firmgrounds, hardgrounds) and/or form a rich lingulide brachiopod shells, which on modern
complexly anastomosing network of surfaces in subtropical intertidal flats disintegrate within a
cross-sectional view (i.e., split and reconverge few weeks or months after death (Kowalewski
laterally). This contrasts with the minor surfaces 1996). Extremely rapid post-mortem disintegra-
that occur in non-condensed assemblages, which tion probably also explains the very low agree-
are usually of a single type, or trend up-section ment between living communities and local death
from one type to another. It appears t h a t the assemblages of lightly calcified modern crabs
more prolonged the period of condensation, then (e.g., Plotnick et al. 1990).
the more varied or random the mix of internal
d i s c o n t i n u i t y t y p e s (e.g., b i o s t r a t i g r a p h i c a l l y Few such expected differences, however, have
c o n d e n s e d i n t e r v a l s and r e m a n i 6 - c o n t a i n i n g been verified or calibrated by direct-dating of
lags, as contrasted with environmentally conden- naturally occurring death assemblages. This is an
important direction for future studies, as it has
sed intervals, as contrasted with within-habitat
been very successful for mollusks (references dis-
time-averaged assemblages). cussed above) and for benthic foraminifera
These alternative metrics for a taphonomic clock (Martin et al. 1995). Such information would esta-
could be tested in modern environments using blish just how disparate in time resolution the
amino-acid racemization, which can date indivi- records of various groups might be, thus allowing
duals shells back to - 1.5 Ma. An initial test in the more informed comparison of evolutionary and
stratigraphic record used the relative durations of ecological patterns (e.g., one group might appear
discontinuity surfaces to i n d e p e n d e n t l y r a n k to support greater morphometric variability per
scales of t i m e - a v e r a g i n g in associated fossil species than another, but if the more variable
assemblages, whose taphonomic grade and other group was known to be subject to greater time-
f e a t u r e s could t h e n be d e t e r m i n e d (Kidwell averaging per collecting horizon, then its morpho-
metric variability might be inflated taphonomi-
1993a, 1993b). Other siliciclastic records and
calty). This information would also be useful in
especially carbonate records need examination community-level paleoecology, by indicating whe-
before the true reliability of these potential clocks ther different taxa within an assemblage might
will be known. have been subject to significantly different scales
Thus, in evaluating the time-resolution of fossil of time-averaging (disharmonious time-averaging
assemblages, a broad and strongly geological sensu Kowalewski 1996).
approach is the most powerful one. The target We should not assume that the result of such
assemblage should be evaluated not only in terms investigations will be bad news, that is, that the
of shell condition (i.e., Johnson's criteria) but also multitaxic assemblage is wildly disharmonic. For
in terms of its diagenesis, sedimentology (inclu- example, Martin et al. (1996) and Anderson et al.
ding bedding style), and larger stratigraphic (1997) both found that Holocene benthic foramini-
context, as these latter features appear to be the feral assemblages contain well-preserved speci-
most informative. mens several thousands of years old on subtropical

intertidal flats and open shelves (respectively), allochthonous shells are also present. The most
comparable to the scale of time-averaging docu- commonly documented p a t t e r n on aggrading
mented in associated molluscan assemblages. open shelves and ramps has three broad depth
zones: (1) within-habitat time-averaged assem-
LARGE-SCALE PATTERNS blages in foreshore and shoreface facies domina-
Environmental variation in time-averaging ted by storm erosion and reworking; (2) a mix of
census assemblages interstratified with within-
The variety of processes and circumstances that habitat time-averaged background sediments in
influence time-averaging suggests that time-reso- the nearshore transition zone, where post-storm
lution for a given taxonomic group m a y vary deposition is greatest; event-beds of allochthonous
significantly among coeval environments. Water fossils are also common here; and (3) within-habi-
energy, frequency of storm reworking, rates of tat time-averaged assemblages dominate the off-
bioerosion, etc. are all known to affect rates of ske- shore zone of distal deposition below m a x i m u m
letal destruction and can vary dramatically bet- storm wavebase; these assemblages take the form
ween habitats within a single marine embay- of dispersed shells in a fine-grained matrix, mixed
ment. Moreover, depths of sediment mixing by by bioturbation and other biological processes;
bioturbators v a r y both along-shore and offshore they m a y also have shelly patches produced by
(from a few cm to several meters penetration), as within-habitat taphonomic feedback.
do rates of sediment supply and deposition.
Offshore areas are generally seen as the most
Direct-dating of modern molluscan assemblages common setting for sediment starvation and thus
indicates that nearshore assemblages are less for prolonged time-averaging. The actual record,
time-averaged t h a n open shelf assemblages, which however, is more complex (reviewed in Kidwell
are also environmentally condensed due to silici- 1991a, 1993a). (1) Unless starvation persists
clastic starvation during Quaternary transgres- through a significant change in water depth or
sion (see discussion above). But we have virtually oxygenation, deep water assemblages will not
no information on how molluscan time-resolution necessarily be more complex t h a n within-habitat
varies among other environments, including open time-averaged due to the g r e a t b a t h y m e t r i c
shelves that are aggrading, and so we can only ranges of most bathyal and abyssal bio-lithotopes;
speculate on the basis of modern sedimentation the absolute duration of time-averaging m a y be
patterns (e.g., Fiirsich & Aberhan 1990; Kidwell & quite long, of course, if sediment accumulation
Bosence 1991; Kidwell & Flessa 1995). rates are low. (2) In tectonic troughs, deep sea-
As a more tangible source of information, the stra- floors m a y not be starved owing to axial transport
tigraphic record yields a great variety of patterns of clastics; shallower water parts of the slope and
(reviewed in Kidwell 1993a) (Fig. 7). For example, shelf may be starved instead. (3) During rapid
the records of coastal lagoons and beach com- transgression (rapid increase in eustatic rise, or
plexes are typically heterogeneous, with a mosaic increase in tectonic subsidence), broad areas of
of census (catastrophic burial, oxygen- and salini- the shallow shelf m a y be sediment-starved, pro-
ty-stress, etc.), w i t h i n - h a b i t a t t i m e - a v e r a g e d ducing hiatal concentrations with high potential
("normal" bioturbated facies), and environmental- for environmental condensation; these assem-
ly condensed assemblages (in current-generated blages m a y record less absolute time t h a n deep
bioclastic banks, channel lags, bioherms, biogeni- water within-habitat assemblages (1 above). (4)
cally graded beds); assemblages dominated by Shallow water hiatal concentrations with envi-

FIGURE 7 - Variation in typical

scales of t i m e - a v e r a g i n g across
shallow m a r i n e facies tracts, and
t h r o u g h transgressive-regressive
depositional sequences (adapted
from figures in Kidwell 1991a,
1991b, 1993a). Variations & des
dchelles typiques d~valuation tem-
porelle dans des facies marins de
faible profondeur et des sdquences
de ddp6ts transgressives-rdgres-
sires (adaptd de figures de KidweU
1991a, 1991b, 1993a). ~-,=skletal concentrations

r o n m e n t a l l y condensed assemblages m a y also ter subsidence; environmental condensation could

form in nearshore areas of intense sediment win- be associated with each flooding surface, but this
nowing and bypassing, and on tectonic and other appears to be rare in actuality; and (2) the poten-
paleohighs subjected to winnowing and located tial for thin, stratigraphically condensed facies
any distance from the shoreline. wherever time-lines converge (parasequence lap-
out), most particularly along the leading edges of
Thus, the record indicates that bathymetric and transgressive surfaces, in distal portions of the
other environmental trends in time-averaging are basin during m a x i m u m transgression, and late in
highly variable but strongly linked to short- and the highstand tract (toplap), although this latter
long-term patterns of sediment accumulation. condition is r a r e l y developed or preserved.
Areas of highly episodic sediment deposition favor Assemblages containing remani6 depend upon
the formation and especially the preservation of erosional reworking of older strata, and thus are
censuses, whereas areas of low net rates of sea- largely limited to sequence boundaries (especially
floor aggradation, either because of repeated win- the margins of incised valleys) and transgressive
nowing of episodic deposits or because of sediment surfaces. Aside from these facies of condensation
starvation, favor faunally condensed assemblages and erosion, the marine record is dominated by
of various sorts. Mixing by bioturbation can be census/snapshot and withinrhabitat time-avera-
superimposed on any of these regimes; its scale
ged assemblages. These non-random distribution
and intensity relative to clastic sedimentation are
patterns clearly have implications for studies on
further factors in the preservation of censuses, but
mixing depths determined by bioturbators appear modes of species ev01~ti0n, on alpha-diversity
to be only a secondary control on time resolution. variation through sealevel fluctuations, and on
confidence limits for biostratigraphic zones, since
Variation through depositional sequences m a n y bed-level datapoints - especially the impor-
tant collecting horizons immediately below and
Sediment accumulation rates clearly are one fac-
above stratigraphic hiatuses - m a y be derived
tor influencing scales of time-averaging in fossil
assemblages, and so time-resolution can be expec- from assemblages t h a t are condensed to some
ted to v a r y s y s t e m a t i c a l l y (non-randomly) degree (Holland 1995; Marshall 1997).
t h r o u g h depositional sequences. Assemblages Although the first-order control on time-resolution
with the most complex histories of time-averaging through depositional sequences is determined by
are expected to be related to settings of stratigra- the mosaic of net sediment accumulation rates -
phic condensation (sensu Jenkyns 1971), t h a t is, the time-space mosaic of erosion, condensation,
where the stratigraphic record is thin relative to and "normal" deposition - the short-term dynamics
coeval sections owing to low net sedimentation. of condensation are also important. Stratigraphic
This slow net accumulation m a y be due to sedi- intervals that are thin (condensed) because of win-
merit starvation, sediment bypassing, or episodic nowing and episodic erosion during accumulation
seafloor winnowing, so that condensed intervals will tend to have discontinuous time-averaging
are known from a wide variety of settings, inclu- and will commonly include fossils reworked from
ding both shallow and deep water and both proxi- older layers, whereas intervals that are thin becau-
mal and distal locations within basins (Jenkyns se of little or no sediment supply (sediment starva-
1971; Wendt & Aigner 1985; Kidwell 1991a,b; tion) will tend to have continuous time-averaging
G6mez & Fern~ndez-Ldpez 1994; Fels & Seyfried and little contribution of reworked fossils, at least
1993) (Fig. 7). when these conditions occur in quiet or deep water
Environmentally condensed assemblages are not (Fiirsich & Aberhan 1990; Kidwell 1991a, 1991b;
limited to stratigraphically condensed intervals, Brett & Baird 1993; Gdmez & Fern~dez-LSpez
but these intervals are good potential sites of 1994). Other variants include sediment starvation
environmentally and biostratigraphically conden- in relatively high-energy shallow water, such as
sed assemblages, and thus we expect - and in fact pertain during initial marine transgression and on
do find - non-random variation in the time-resolu- some paleohighs; these are characterized by dis-
tion of assemblages t h r o u g h depositional continuous time-averaging.
sequences (Kidwell 1991a, 1991b, 1993a, 1993b; Variation through the Phanerozoic
Ffirsich et al. 1992; Brett & Baird 1993; Brett
1995; Krawinkel & Seyfried 1996; Anderson & Evolution in the n a t u r e of skeletons produced as
McBride 1996). Sequence stratigraphers continue well as in the organisms t h a t influence skeletal
to discover new variants on the anatomy of third- preservation (e.g., bioeroders, deep-irrigating bio-
order (- i m y duration) sequences, but some basic turbators, duraphagous predators) suggests t h a t
elements of the model have been stable, particu- scales, absolute durations, biological biases, and
larly: (1)divisibility into finer-scale, shallowing- taphonomic s i g n a t u r e s of time-averaging might
up parasequences in settings of moderate or grea- also have changed over geologic time. To approach

this, we compiled a large, field-checked database of living brachiopods, bryozoans, and crinoids,
on the diversity and physical scale of densely pac- comparable to the time-resolution studies of
ked skeletal concentrations (non-reef), reasoning living mollusks (see also discussion above), and
t h a t these were a function of several factors inclu- (2) stratigraphic searches for taphonomic diffe-
ding skeletal durability, which in t u r n is a limi- rences in fossil assemblages associated with dis-
ting factor on scales of time-averaging (Kidwell & continuity surfaces of similar hiatal duration but
Brenchley 1996). Data for Ordovician-Silurian, different geological age. For example, hiatuses
Jurassic, and Neogene s t r a t a (and d a t a on associated with parasequence flooding surfaces
Cambrian s t r a t a from Li & Droser 1997) indicate are generally shorter t h a n those associated with
significant expansion over the Phanerozoic in the sequence boundaries, providing a means of ran-
bioclasticity of the shallow marine record, and king the m a x i m u m period of time-averaging in
suggests t h a t this expansion was driven by evolu- the fossil assemblages t h a t m a n t l e such surfaces.
tionary and ecological changes among shell pro- Such a survey in post-Paleozoic records (Kidwell
ducers r a t h e r t h a n by geological and biological 1993b) revealed consistent differences in the
changes in post-mortem environments. With the kinds of skeletal concentrations associated with
exception of crinoids, thick bioclastic concentra- different ranks of discontinuity surfaces, implying
tions appear to be a f u n d a m e n t a l l y modern phe- a different envelop of scales of time-averaging for
nomenon, with shellbeds expanding in thickness each r a n k of surface, but there are as yet no com-
as evolutionarily Modern clades diversified taxo- parable d a t a for t h e Paleozoic. Phanerozoic
nomically in the Mesozoic and Cenozoic and shif- declines in other aspects of time resolution in the
ted the global biota toward more durable low- marine record have also been suggested, related
organic microstructures, toward occupation of primarily to increasing bioturbation [e.g., appa-
h i g h e r - e n e r g y h a b i t a t s , and p e r h a p s t o w a r d rent declines in the frequency of discrete storm
higher rates of carbonate production. beds and of soft-tissue and fragile skeleton pre-
An increase in the post-mortem durability of ske- servation (Brandt 1986; Sepkoski e t al. 1991;
letons over Phanerozoic time suggests t h a t skele- Allison & Briggs 1994; B u t t e r f i e l d 1995;
tal assemblages from the younger fossil record are Kowalewski & Flessa 1996)]. Such broad scale
more time-averaged t h a n those from the older los- changes in the quality of the sampled record are
sil record: the more durable the hardparts, the still little explored and exciting directions for
greater the probability t h a t hardparts from suc- future taphonomic research.
cessive generations and/or communities at a site A c k n o w l e d g m e n t s - I am grateful to the editors of Geobios
will survive short-term reworking events and for the invitation to contribute to this anniversary issue, and
become mixed into a single time-averaged fossil to David Jablonski for a helpful review. This paper is an out-
assemblage. There have always been census growth and updating of notes for a Paleontological Society
assemblages formed by sudden burial or other shortcourse on time resolution in Boston in 1993.
catastrophic events, but the scope for time-avera-
ging appears to have expanded significantly. The
likely discrepancies between early Paleozoic and
late Cenozoic scales of time-averaging are difficult
to quantify, but could easily be an order-of-magni- ALLISONP.A. & BRIGGSD.E.G. 1994 - Exceptional fossil
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order of tens to hundreds of years at maximum for through the Phanerozoic. Geology, 21: 527-530.
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years documented for modern bivalves. The most genesis and history on the northestern Gulf of
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