RESEARCH REPORTS
Sequence Stratigraphy, Paleoecology, and Evolution:
Biotic Clues and Responses to Sea-Level Fluctuations
CARLTON E. BRETT
Department of Earth and Environmental Sciences, University of Rochester, Rochester, N.Y. 14627
PALAIOS, 1998, V. 13, p. 241–262
Paleoecology has a dual relationship with sequence stratig-
raphy. On one hand, body and trace fossils, together with
their taphonomy, may provide sensitive indicators of envi-
ronmental parameters, including depth, substrate consis-
tency, sedimentation rate/turbidity, and benthic oxygena-
tion, which are critical in recognizing and interpreting par-
asequences and sequences. Fossils may provide some of the
best guides to identifying key surfaces and inferring sedi-
mentation dynamics within sequences. Conversely, the se-
quence stratigraphic paradigm and its corollaries provide a
predictive framework within which to examine biotic
changes and interpret their probable causes. Such changes
include ecological epiboles (short-term, widespread prolif-
eration of normally rare species), outages (absence of nor-
mally common species), ecophenotypic changes, and long-
term (tens to hundreds of Ka) community replacement.
Community replacement should be carefully distinguished
from short-term (10 to a few hundred years) ecological suc-
cession, rarely resolvable at the scale of single beds, al-
though replacement series through shallowing-to-deepen-
ing cycles may display some features that parallel true suc-
cession. Replacement in marine communities may be rela-
tively chaotic, but, more commonly in offshore settings, it
appears to involve lateral, facies-related shifting of broad
biofacies belts, or habitat tracking. Tracking patterns may
be nearly symmetrical in areas of low sediment input. How-
ever, replacement cycles are commonly asymmetrical. The
asymmetries involve both apparent and real effects; dele-
tion of portions of facies transitions at sequence boundaries
or condensed sections leads to artifactual asymmetries. Al-
ternatively, in areas proximal to siliciclastic sources, track-
ing asymmetries arise from the markedly higher sedimen-
tation rates during regressive (late highstand) than trans-
gressive phases. Replacements may also involve immigra-
tion of species into a sedimentary basin, either as
short-lived events (incursion epiboles) or as wholesale fau-
nal immigrations. The latter will typically follow intervals
of extinction/emigration of the indigenous faunas. Both
large and small immigration events appear most common-
ly during highstands (transgressive peaks), which may be
associated with altered water-mass properties, and may
open migration pathways for nekton and planktonic larvae.
At least in isolated basins, allopatric speciation may also
occur during fragmentation of habitats associated with re-
gressions.
Finally, there are predicted and empirical correlations
between sequence-producing sea-level fluctuations and mac-
roevolution. Major extinctions may be associated with hab-
itat reduction during major regressions (lowstands), or
Copyright Q 1998, SEPM (Society for Sedimentary Geology)
241
with anoxic events during major transgressions. Generally,
rising sea level may be correlated with evolutionary radia-
tions. Hence, some ecological-evolutionary unit boundaries
may correlate either with sequence boundaries or maximum
flooding surfaces. However, in other cases, no correlation
has been found between macroevolutionary patterns and se-
quence stratigraphy. The situation is obviously complex, but
sequence stratigraphy at least provides a heuristic frame-
work for developing and testing models of macroevolution-
ary process.
INTRODUCTION
One of the great challenges of the Earth sciences is the
development of ‘‘bio-geo-history.’’ This is an integrated
view of global change that encompasses the complex inter-
play of biological and physical processes over a hierarchy
of time scales ranging from the geologically instantaneous
to hundreds of millions of years. A potentially fruitful area
of integration lies at the interface between paleoecology
and evolutionary studies, on the one hand, and sequence
stratigraphy in its relationship to global sea-level and cli-
matic changes, on the other (Kauffman and Sageman,
1992; Sageman, 1992; Gómez and Fernandez-López,
1994).
The principles of sequence stratigraphy, originally
based on seismic studies of continental shelf strata and
formulated by Vail et al. (1977a, b) and Mitchum et al.
(1977), have subsequently been refined and applied to a
broad array of marine and non-marine depositional sys-
tems (Wilgus et al., 1988; Van Wagoner et al., 1988; Vail et
al., 1991; Emery and Meyers, 1996). Sequences are of vari-
able duration, spanning hundreds of thousands to millions
of years and have been subdivided by some authors into
different scales: second-order supersequences, spanning
tens of millions of years; third-order sequences with dura-
tions typically estimated at 1.5 to 5 million years; and
fourth-order sequences, or subsequences, that span about
100 thousand to a million years. Regardless of scale, how-
ever, sequences share common patterns in terms of re-
corded fluctuations in water depth and sedimentation. De-
positional sequences are internally conformable packages
of strata bounded by marine to subaerial erosion surfaces
(sequence boundaries) formed during sea-level lowstands.
For most shallow marine settings, lowstand deposits are
poorly developed. Typically, the sequence boundary also
represents a transgressive surface at which shallow ma-
rine deposits of a transgressive systems tract unconform-
ably overlie more offshore marine facies, the lowstand be-
ing represented only by the erosional unconformity itself
(Baum and Vail, 1988; Posamentier and Vail, 1988; Mey-
0883-1351/98/0013-0241/$3.00
242
ers and Milton, 1996). A second important surface, the
maximum flooding surface or surface of maximum star-
vation, separates retrogradational (overall deepening up-
ward) transgressive deposits from those of the overlying
early highstand. Deepest water conditions are typically re-
corded in highly condensed strata immediately above the
maximum flooding surface (Baum and Vail, 1988; Loutit
et al., 1988; Meyers and Milton, 1996). The highstand dis-
plays aggradational to progradational (shallowing up-
ward) stacking patterns of small-scale subsymmetrical to
shallowing upward cycles or parasequences. Thus, a typi-
cal depositional sequence displays a pattern of decreasing
terrigenous sedimentation rates in the transgressive sys-
tems tract, culminating in a condensed section and/or a
surface of maximum sediment starvation; depending upon
local conditions, intrabasinal carbonates may or may not
accumulate in shallow marine environments during this
time (Sarg, 1988; Emery, 1996). Siliciclastic sedimenta-
tion rates will increase in concert with shallowing in many
prograding highstand systems tracts, resulting in coars-
ening upward successions (Posamentier et al., 1988; Po-
samentier and Vail, 1988; Meyers and Milton, 1996). How-
ever, in some cases, actual drop in sea level may cause
forced regressions (Meyers and Milton, 1996); under these
conditions, a minor erosional surface or condensed bed
may separate early (aggradational) and later (prograda-
tional or regressive) portions of the highstand; Brett
(1995) referred to this type of condensed bed as a ‘‘precur-
sor bed.’’ Such beds are typically overlain by successions
displaying marked shallowing upward trends.
A number of patterns of paleoecological and even evolu-
tionary change appear to correlate with fluctuations in
water depth and sedimentation through the course of de-
positional sequences. Sequence stratigraphy, therefore,
may provide a key to the interpretation of these patterns,
as well.
In recent years, paleontologists have begun to develop
integrative models of depositional environments and pa-
leoecology (Valentine, 1973; Einsele and Seilacher, 1982;
Dodd and Stanton, 1990; Miller, 1990; Einsele et al., 1991)
and evolutionary ecology (Valentine, 1985; Walliser,
1986a; Bayer and Seilacher, 1985; Donovan, 1989; Kauff-
man and Walliser, 1990; Ross and Allmon, 1990; Eldred-
ge, 1990; Lees and Edwards, 1993; Erwin and Anstey,
1995). However, until recently, there has been little at-
tempt to meld these approaches with the paradigms of se-
quence stratigraphy. Because paleoecological changes are
closely correlated with fluctuations in sea level and sedi-
mentation, one may predict many genetic relationships
between fossil distributional patterns and depositional se-
quences.
The relationship between paleoecology and sequence
stratigraphy is actually twofold. First, data from the fossil
record may provide key information on depositional envi-
ronments critical to the interpretation of marine sedimen-
tary cycles and, thereby, to the recognition of ancient cy-
cles and sequences. Second, the sequence stratigraphic
framework provides a predictive model within which to ex-
plain various paleoecological phenomena: iterative cycles
of ecophenotypic change, faunal or community-level track-
ing, and various perturbations in marine paleoecology
that relate to changes in water depth, sedimentation, or
BRETT
other parameters that are tied to cycles of relative sea-lev-
el rise and fall.
Sequence stratigraphy provides a temporally con-
strained framework for the evaluation of ecological and
evolutionary events. For example, it may permit precise
evaluation of the timing of immigration, extinction, or
origination of new taxa in a region or, with less precise res-
olution, on global scales. Unfortunately, the processes of
sedimentation and erosion that produce sequence bound-
aries, condensed intervals, and flooding surfaces also im-
pose an inherent bias on the fossil record that may pro-
duce artifacts in the apparent distribution of fossils. It is
critical that paleontologists recognize and attempt to
avoid these potential pitfalls in literal interpretation of the
fossil record (Jablonski, 1985; Ross and Ross, 1988, 1994;
Holland, 1995). Nonetheless, the same processes of sea-
level, climate, and sediment-supply fluctuation that cre-
ate depositional sequences also may have strong biological
impacts. Immigration, origination, and extinction are
commonly related to these factors, so that there may be
important correlations between evolutionary patterns and
sequence stratigraphy.
In a companion paper (Brett, 1995), I considered the re-
lationships of sequence stratigraphy to biostratigraphic
and taphonomic patterns. Herein I focus upon the dual re-
lationship of sequence stratigraphy with paleoecology—
the use of paleoecological data in recognizing and inter-
preting sequences, and the real and apparent effects of se-
quence-generating phenomena on patterns of fossil organ-
ism distribution and evolutionary paleoecology.
FOSSILS AS PALEOENVIRONMENTAL TOOLS IN
SEQUENCE STRATIGRAPHY
Information derived from fossils in sedimentary se-
quences may be of critical importance in the interpreta-
tion of sequence patterns. This is particularly important
for the inference of ancient water depths or paleobathy-
metry; however, evidence for changes in substrate consis-
tency, sedimentation rate and turbidity changes, and bot-
tom-water oxygenation may also provide insights into the
environmental responses to sedimentary cycles and se-
quences. Taphonomic features related to sequence stratig-
raphy are discussed in Brett (1995); herein I focus on pa-
leoecological approaches.
Water Depth
Paleontologic data frequently provide a far more sensi-
tive gauge of relative water depth or even absolute depths
than do sedimentologic features. Paleontologic data pro-
vide a number of potential constraints on relative and ab-
solute water depths that may be extremely important in
establishing the magnitude of sedimentary cycles (Bene-
dict and Walker, 1978; Brett et al., 1993). Particularly
valuable indicators include marine algae, especially dasy-
cladaceans and certain types of endolithic green algae,
and cyanobacteria. For example, Beadle and Johnson
(1986) used cyclocrinitids to demonstrate that certain Si-
lurian facies accumulated in shallow photic shelf condi-
tions. Other lines of evidence for depth include evidence
from orientation and implosion of ammonites, eyes in var-
ious arthropods, and hermatypic corals (Brett et al., 1993).
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
Oxygen isotopic data derived from calcitic shells also
may be related to depth variations in some cases. For in-
stance, Adlis et al. (1988) observed cyclic variations in d18O
values in non-luminescent (unaltered) brachiopod shells
through Pennsylvanian cyclothems. Shells with highest
values (coolest temperatures) occurred in highstand facies
(deepest and coolest water). Conversely, Railsback et al.
(1990) used a combination of oxygen isotopes and punctal
density in Middle Ordovician brachiopods to infer the
presence of warm, saline bottom waters in the Taconic
foreland basin.
The relationships between ichnofacies and paleobathy-
metry, established by the seminal research of Seilacher
(1967), have been supported in a general sense by numer-
ous case studies (Fig. 1B–E; see revisions and updates of
Frey et al., 1990; Maples and West, 1992, and articles
therein; Droser and Bottjer, 1993; Bottjer and Droser,
1994; Goldring, 1995). However, factors other than water
depth (e.g., substrate consistency and oxygenation) exert a
very significant influence on the distribution of trace fos-
sils, leading Frey et al. (1990) to conclude that only weak
and indirect relationships exist between trace fossils and
depth. Moreover, certain trace makers may have shifted
their environmental positions, primarily in an onshore to
offshore direction through the Phanerozoic (see Droser
and Bottjer, 1993; Bottjer and Droser, 1994). Nonetheless,
for a given time interval and substrate type, ichnofacies
may still provide useful clues to relative water depth.
Thus, vertical dwelling burrows of the Skolithos ichnofa-
cies typify nearshore sands, especially in the early Paleo-
zoic. Conversely, grazing traces (Nereites ichnofacies)
characterize oligotrophic deep water muds, at least in the
Mesozoic-Cenozoic.
A further application of paleontologic data to interpre-
tation of paleobathymetry makes use of the predictable re-
currence of particular benthic biofacies with depth zones.
This approach, pioneered by Ziegler (1965), has proven ex-
ceptionally useful. Many biofacies do appear to be consis-
tently arrayed with respect to depth (Ziegler et al., 1968;
McKerrow, 1979; Boucot, 1975, 1982; Boucot and Lawson,
1998, and references therein). Furthermore, organisms
tend to be evolutionarily conservative with respect to their
preferred depth ranges. Hence, most genera and families
seem to have occupied similar bathymetric zones for mil-
lions of years (Boucot, 1990c; Aberhan, 1994). Similarly,
Holland (1997) observes that most Upper Ordovician bra-
chiopod and bivalve genera display environmental stasis,
although he also documents a few brachiopod species that
appear to shift their habitat preferences through time.
The generality of environmental stasis permits recogni-
tion of analogous depth positions in various systems even
though these are populated by suites of distinct species.
Fluctuations of fossil communities have been used exten-
sively in recognizing ancient sea-level fluctuation, partic-
ularly in the middle Paleozoic (e.g., Johnson, 1987; John-
son et al., 1985). Cisne and Rabe (1978) and Cisne et al.
(1984) outlined a detailed quantitative application of
depth-related fossil associations referred to as gradient
analysis. Application of these techniques may permit rec-
ognition of subtle cyclicity even in ‘‘basinal’’ mudrocks
where sedimentological cues are lacking.
243
Salinity
Many organisms are sensitive to slight fluctuations of
salinity. The presence of stenohaline organisms, such as
echinoderms, articulate brachiopods, or cephalopods, may
be a critical indicator of open marine conditions within
particular portions of depositional sequences; they should
typify later transgressive and early highstand deposits.
Conversely, environments of reduced and/or variable sa-
linity may be identified by the occurrence of low-diversity
euryhaline faunas (e.g., lingulids, oysters; see Fürsich,
1994; Aberhan, 1994). Because such faunas characterize
brackish-estuarine environments, they may be important
in recognizing paralic, lowstand deposits. Moreover, in
some instances, the appearance of marine ichnofossils at
certain horizons of otherwise non-marine sediments may
indicate minor marine incursions, associated with maxi-
mum flooding events (see Fig. 1C).
Substrate Consistency
The nature of sedimentary substrates typically varies in
a predictable manner through sequences and parasequ-
ences. In particular, during times of relative low sediment
input (e.g., in condensed sections), sediments may become
relatively firm to cemented. Submarine erosion, common-
ly associated with lowstand intervals, may also remove su-
perficial water-rich sediment, thereby exposing firm, ov-
ercompacted sediment to colonization by endofauna (Fig.
1D). Winnowing processes in shallow water occurring dur-
ing later parts of the highstand may produce shell or other
gravel deposits, which develop a firm armor or pavement
on the sea floor capable of supporting a number of species
that are restricted in habitat to colonization of hard sub-
strates (Figs. 1A, E). These organisms, in turn, may con-
tribute further skeletal debris, leading to positive taphon-
omic feedback (Fürsich, 1978; Kidwell and Jablonski,
1983; Miller et al., 1988; Kidwell, 1991). Finally, reduction
of sediment input following periods of winnowing and
shell-gravel accumulation may result in the formation of
submarine hardgrounds at flooding surfaces.
Consistencies of certain substrates (e.g., hardgrounds)
may be quite obvious; however, others cannot be so readily
recognized (Fig. 1A, E). For example, how may we distin-
guish between a firm, overcompacted mud versus sedi-
ment that was, at the time of final burial, soft and plastic?
Fossils, especially trace fossils, may provide very impor-
tant clues in this regard and may be very useful in identi-
fying key discontinuities within sequences (Figs. 1B–D).
For example, Sellwood (1970), Savrda (1991) and Jones
and Pemberton (1989), examined patterns of distribution
of ichnofabrics in sedimentary sequences. Sequence-
bounding unconformities and surfaces of maximum star-
vation display distinctive omission trace fossil suites (Figs.
1, 2). Where the latter are developed as firm- or hard-
grounds, sharply defined burrows or borings of hard sub-
strate-penetrating organisms are common (Fig. 1B,D,E).
Deep, well-defined burrows, including Thalassinoides and
Spongeliomorpha, are particularly well developed at ra-
vinement (transgressive) surfaces where overconsolidated
substrates have been exposed by submarine erosion (Kid-
well and Aigner, 1985; Bayer, Altheimer, and Deutchle,
1985; Landing and Brett, 1987; Savrda, 1991; Taylor and
244
FIGURE 1—Body and trace fossils as indicators of paleoenvironment
and of significant stratal surfaces. (A) Biogenic indicators of substrates
of varying consistency from soft (on left) to firm, and hard (right). (a)
Nuculid bivalve, shallow infaunal deposit feeder. (b) Soft-substrate
trace fossil Chondrites. (c) Pellet-lined burrow Ophiomorpha. (d) Bur-
rowing heart urchin. (e) Infaunal suspension-feeding bivalve. (f)
Sharply-defined dwelling burrow Thalassinoides. (g) Trace Sponge-
liomorpha. (h) mechanical mud-boring bivalve Petricola. (i) Chemical
rock-boring bivalve Lithophaga. (j, k) Crinoid holdfast and oyster ce-
mented to hardground surface. (l) Trypanites (borings). (m) Serpulid
worm tubes and bryozoans encrusting on cemented wall of relict bur-
row. (B–E) Ichnological signatures of stratigraphically important sur-
faces. (B) Marine flooding surface. (C) Minor marine incursion into
non-marine environment. (D) Omission surface associated with shal-
lowing and overlain by condensed bed (i.e., precursor bed; see text).
(E) Omision surface recording interval of sediment starvation and
hardground development in offshore facies (e.g., surface of maximum
starvation associated with early highstand). Adapted from Goldring
(1995), after Taylor and Gawthorpe (1993).
Gawthorpe, 1993). Sneden (1991) described spectacular
burrows, over a meter deep, at flooding surfaces of Upper
Cretaceous parasequences; he attributed these structures
to tilefish that excavated dwelling burrows in firm sand at
a ravinement surface.
Body fossils are less diagnostic of substrate consistency.
However, certain morphotypes, such as the deeply cup-
shaped valves of Gryphea in Jurassic mudstones, as op-
posed to broad, flattened and encrusting form, may be an
indicator of soft substrate consistency (Seilacher, Matyja,
and Wierzbowski, 1985; Figs. 1, 7). Likewise, Alexander
(1975) documents intraspecific differences in Ordovician
BRETT
FIGURE 2—Distribution of ichnofabrics and ichnogenera within two
depositional sequences from the Paleocene of Alabama. Note: intense
bioturbation by Thalassinoides suevicus (Th) in package 2 associated
with decrease in sedimentation in transgressive systems tract and pip-
ing of sediment at transgressive surface (top of package 1); diverse
ichnofossils in offshore facies of early highstand (especially in unit 24);
also diffuse burrowing in later highstands (packages 3 and 5) and
Teredolites woodground ichnofacies in transgressive lag deposits
overlying sequence boundary in unit 19 (from Savrda, 1991).
strophomenid brachiopods that relate to substrate consis-
tency. Obviously, the occurrence of encrusting (e.g., oys-
ters) and endolithic (e.g., lithophagid bivalves) organisms
directly on bedding planes indicates a very firm or hard
substrate consistency, and these fossils often provide key
evidence for the occurrence of hardgrounds in the marine
carbonate sedimentary record (Fig. 1; see Bromley, 1975;
Palmer and Palmer, 1977; Fürsich, 1979; Brett and Brook-
field, 1984; Brett, 1988; Wilson and Palmer, 1992; Gold-
ring, 1995). Conversely, the occurence of abundant nucu-
lid bivalves, as well as strong burrow deformation, may
imply very soft and thixotropic sediments (Fig. 1A).
Sedimentation Rate and Turbidity
Relative rates of sedimentation and associated levels of
water turbidity may be expected to vary predictably with-
in a sedimentary sequence. However, the assessment of
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
these rates is exceptionally difficult. It is closely tied to the
issue of time resolution in rocks, which has been ad-
dressed recently by Kidwell and Behrensmeyer (1993, and
papers therein). In addition to the taphonomic and trace
fossil indicators suggested above, paleoecology may pro-
vide an important tool in this endeavor.
Assemblages dominated by suspension feeding, endo-
and epibenthic organisms tend to be well-developed under
conditions of low sedimentation rates, low turbidity, and
firm substrates (Rhoads et al., 1972; Scott, 1978; Fürsich,
1978; Elder, 1987). Hence, a ratio of deposit to suspension
feeders may be a useful predictor of relative sedimentation
rates in sedimentary sequences.
If there is independent evidence for normal oxygena-
tion, variations in sedimentation rates may be inferred
from the relative concentration of trace fossils in vertical
successions (e.g., Savrda and Bottjer, 1994). Thus, Morti-
more and Pomerol (1991), Taylor and Gawthorpe (1993,
see Fig. 1B–E), and Savrda and Bottjer (1994) have docu-
mented the significance of concentrated, cross-cutting
trace fossils in recognizing minor omission and scour sur-
faces, including marine flooding surfaces (Figs. 1, 2). In
contrast, the occurrence of numerous discrete burrows
may indicate a higher sedimentation rate. Furthermore,
certain trace fossils, such as Zoophycos, appear to be sen-
sitive indicators of increased sedimentation rates (Brett,
1995). Storm-event beds may also be recognizable from
distinctive suites of ‘‘opportunistic’’ trace fossils (Vossler
and Pemberton, 1988; Pemberton and MacEachern,
1997). Such types of trace fossil occurrences should occur
predictably during middle-to-late highstands of deposi-
tional sequences that are characterized by shallowing and
increased sedimentation rates. In dysoxic facies it may be
difficult to distinguish the effects of sedimentation from
variable bottom-water oxygenation (see below), although
recognition of oxygen-related ichnofacies and trace fossil
tiering aid in this regard (see Savrda and Bottjer, 1994).
Truncation of trace fossils, such as borings with distinc-
tive profile shapes (e.g., Warme, 1975; Taylor and Gaw-
thorpe, 1993; Fig. 1D), may provide evidence for erosion of
the substrate. Such truncation may occur at hardgrounds
and other discontinuity surfaces. Goldring (1971) used al-
ternations of protrusive and retrusive spreiten in the trace
fossil Diplocraterion yoyo to infer fluctuating depositional/
erosional conditions (Fig. 1C). In a related line of reason-
ing, Wetzel and Aigner (1986) used tiered (i.e., depth par-
titioned) suites of trace fossils as a ‘‘yardstick’’ for seafloor
erosion. Because particular types of traces occupy approx-
imately consistent depths, it is possible to estimate the
amounts of sediment removed when only deeper tiers are
preserved below truncation surfaces. Such tools may be
useful in identifying submarine erosion surfaces associat-
ed with sequence or parasequence boundaries.
The abundance of light-sensitive or photosynthetic or-
ganisms may provide a good indicator of a low sedimenta-
tion rate and turbidity. Turbidity-sensitive taxa, such as
corals and suspension-feeding echinoderms, whose water
vascular systems may be clogged by excess fine-grained
sediments (Rosenkranz, 1971; Seilacher, Reif, and West-
phal, 1985), may also be useful indicators of low rates of
sedimentation and slight water turbidity.
The distribution of these organisms is expected to vary
consistently within a sedimentary sequence. For example,
245
higher-diversity assemblages of suspension-feeding or-
ganisms will typically occur during transgressive phases,
when turbidity and sedimentation were relatively low as a
result of nearshore sediment entrapment. Conversely, the
emptying of coastal sediment traps associated with sea-
level drop should greatly increase the influx of fine- to me-
dium-grained sediments into offshore areas (see Posamen-
tier et al., 1988; Posamentier and Vail, 1988; Meyers and
Milton, 1996). This will result in consequent negative ef-
fects on organisms that were intolerant of turbidity, and
positive effects on primarily deposit-feeding forms that re-
quired increased amounts of organic detritus and/or soft
substrate for their survival.
Bottom-Water Oxygenation
While not directly related to the processes generating
sedimentary sequences, the level of bottom-water oxygen-
ation is highly sensitive to relative rise and fall of sea level.
In particular, episodes of widespread anoxia or the devel-
opment of large masses of low-oxygen bottom water are
typically associated with transgressions (see Kauffman,
1981; Hallam, 1992; Wignall, 1994). Wignall has reviewed
biotic evidence for low oxygen or dysoxic conditions near
the substrate. While he finds no particular group of organ-
isms to be diagnostic of this zone throughout geologic time,
he does suggest a number of indicators, including overall
diversity and the presence or absence of burrowing organ-
isms, from which to interpret relative oxygenation. Batt
(1993) used proportions of inferred pelagic to nektobenthic
ammonoids as a proxy for paleo-oxygenation in Creta-
ceous cycles from the Western Interior basin.
A relatively sensitive tool for recognition of oxygenation
events and cycles within offshore marine mudrocks was
presented by Savrda and Bottjer (1991, 1994). Horizons of
extensive piping of burrowed, organic-free sediment into
black, laminated muds indicate periods of benthic oxygen-
ation. Likewise, a decrease in the diversity and depth of
trace fossils may point to decreasing oxygen levels or de-
oxygenation events. A tiered trace fossil model is generally
applicable for recognizing relative bottom-water oxygena-
tion on the basis of type and depth of burrows (Kauffman,
1981; Savrda and Bottjer, 1989, 1991, 1994; Sageman et
al., 1991). Many deepening events (e.g., marine flooding
surfaces in basinal sediments) are associated with the rise
of the anoxic boundary and, as such, may be recorded as
deoxygenation events.
BIOTIC RESPONSES TO SEDIMENTARY CYCLES
AND SEQUENCES
Sedimentary sequences are the record of fluctuations in
relative sea level, sedimentation, and other related envi-
ronmental parameters. Hence, one may predict that an ar-
ray of biotic changes will be reflected in their fossil assem-
blages. On the other hand, the preserved patterns of biotic
change may be strongly influenced by changing preserva-
tion potentials. For example, certain portions of a spec-
trum of community replacement may be absent due to a
lack of sedimentation or removal by erosion. Therefore,
distortion of the original pattern of biotic fluctuation will
be predictably severe at sequence boundaries and maxi-
mum flooding surfaces (condensed sections) of deposition-
246
al sequences (Holland, 1993, 1995; Ross and Ross, 1994).
This bias may have the effect of producing apparently
abrupt and simultaneous extinctions and biotic changes,
even if original patterns were gradual and staggered; or,
biases may sharpen truly abrupt turnover events (Hol-
land, 1995). Conversely, lateral shifting of facies within
depositional sequences may lead to a pattern of gradual
first appearances of species (e.g., during immigration
events) as a result of our inability to test for the presence/
absence of facies-controlled species in local sections in
which inappropriate facies are preserved. This is an ex-
pression of the smearing phenomenon originally docu-
mented by Signor and Lipps (1982).
We must carefully distinguish these artifacts of preser-
vation from real biotic responses to transgressive-regres-
sive cycles (see Jablonski, 1980, 1986). Probably the best
approach to this problem is the application of a ‘‘time-en-
vironment’’ approach; specifically, the regional study of fa-
cies spectra in carefully correlated stratigraphic intervals,
as advocated by Holland (1995, 1997). Clearly, such an ap-
proach is critical for a better assessment of the true timing
of biological events; studies of fossil-distribution patterns
in single sections or suites of locally restricted outcrops are
plagued with stratigraphic biases and cannot discern a
complete history of the original patterns. Sequence stra-
tigraphy provides a framework for constraining time rela-
tionships of strata across facies changes. Ideally, it is pos-
sible to trace laterally displaced biofacies with regional
study of the ‘‘time-slices’’ provided by single parasequ-
ences, systems tracts, or sequences (e.g., Elder et al., 1994;
Fig. 3). This will provide a more complete picture of the ac-
tual first and last appearances of fossil species, and
changes in the composition of biofacies. Also, it may be
possible to trace sequence-bounding unconformities and
condensation surfaces into areas of conformable and ex-
panded sections; this procedure will help to fill in the miss-
ing pieces of regional biofacies history.
Holland (1995) has also proposed a method in which the
apparent effects of hiatuses on species-range patterns
may be assessed. In this approach, the patterns of biotic
change, assuming gradual or constant species turnover,
may be simulated for unconformable gaps of different
magnitudes and those patterns compared with those of
the actual fossil record. Observed abrupt shifts that ex-
ceed those produced as the artifact of truncated ranges
may thereby be recognized as partially real biotic effects.
Assuming that spurious effects can be recognized and
accounted for, we may begin to interpret the ways in
which ancient organisms responded to the environmental
fluctuations that produced depositional sequences. The re-
sponses may be either ecological or evolutionary. Ecologi-
cal (facies-related) explanations must be carefully consid-
ered and ruled out before the observed biotic changes are
interpreted as evolutionary. As Jablonski (1980) succinct-
ly pointed out, patterns of changing diversity, endemism,
eurytopy, and other features typically recorded during
transgressive-regressive cycles may best be explained in
terms of migration of fixed onshore-offshore ecological gra-
dients. Hence, habitat tracking (see below), rather than
evolutionary responses to the ‘‘stresses’’ associated with
shifting sea level, may account for a majority of cases of
faunal change in local sedimentary sequences. This phe-
BRETT
nomenon of faunal migration was already discerned and
cogently analyzed long ago by Williams (1903, 1913).
Ecological (non-evolutionary) responses to sequence or
parasequence-producing environmental fluctuations in-
clude community replacement, ecophenotypic changes,
minor species-level substitutions and fluctuations (epibo-
les), and immigration/emigration of species into/from the
local ecosystem. These responses will be considered in the
next section. Evolutionary changes, including extinction,
species evolution and evolutionary radiation, in response
to sequence-producing environmental fluctuations, will be
dealt with in the last section of this paper.
COMMUNITY REPLACEMENT
Wholesale changes in species composition are recorded
in many sedimentary cycles. Such community replacement
commonly involves a predictable, facies-related recur-
rence of species. Where the latter can be related to lateral
shifting of biofacies it is termed habitat tracking (Figs. 3–
6). Before further discussion of replacement and tracking,
it is necessary to carefully distinguish these phenomena
from another pattern that has often been incorrectly con-
flated with replacement: succession.
Replacement vs. Community Succession
In the past, some paleontologists have used the term
succession to denote long-term changes in community
structure within local sections. Neontologists have gener-
ally defined succession as short-term (tens of years) se-
quential change in community structure, generally in re-
sponse to environmental changes induced by the organ-
isms themselves (Odum, 1971; Connell and Slaytor, 1977;
Levinton, 1982). As a result of time-averaging, it is unlike-
ly that such short-term succession can be resolved in most
strata (Schindel, 1980, 1982; Sadler, 1981), although
Walker and Alberstadt (1975) illustrate possible true
cases in level-bottom epibenthic communities and fossil
reefs. In any case, such examples lie at the lower end of
resolution for the fossil record, typically at the scale of in-
dividual rapidly buried bedding planes. Such short-term
‘‘autogenic’’ succession is clearly of a different temporal
magnitude than the faunal replacements, which typically
are observed through several meters of strata in sequenc-
es or parasequences, and must span thousands of years
(Bretsky and Bretsky, 1975; Walker and Alberstadt,
1975). Rollins et al. (1979) tried to distinguish between re-
placement and succession in a Pennsylvanian cyclothem.
However, their example of succession still represented a
significant gradational change in assemblage composition,
spanning a major portion of a cycle. Hence, it is best inter-
preted as a shifting faunal gradient; i.e. gradual replace-
ment, not succession.
Use of the term succession for the longer-term processes
has caused considerable confusion; this term should be re-
stricted to those rare cases in which changes in ecological
time can be documented in the fossil record. Nonetheless,
there may be an indirect relationship between short-term
succession and longer-term replacement, as suggested by
the work of Bretsky and Bretsky (1975; see Fig. 3 herein).
For example, early successional stages in marine commu-
nities are typified by low-diversity assemblages of gener-
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY 247

FIGURE 3—Pattern of asymmetrical faunal replacement (tracking) within probable sedimentary cycles of the Upper Ordovician Nicolet River
Formation of Quebec. Vertical columns represent an ordered series of distinctive species associations with diversity generally increasing from
1 to 4 but declining slightly in 5 and 6. These associations were termed ‘‘successional stages’’ by Bretsky and Bretsky (1975), but represent
relatively long-term changes herein interpreted as different depth-related community types (or biofacies). Original diagram has been modified
from Bretsky and Bretsky (1975) to reflect reinterpretation of ‘‘successional seres’’ as community replacements in about 17 small-scale de-
positional sequences, labelled S-1 to S-17. Alternate sequences are shaded to highlight sequence boundaries (SB). Inferred maximum flooding
surfaces (MFS) are denoted by dotted lines. Typical asymmetrical pattern of community replacement (left to right shift in types 1–6) during
inferred main, shallowing-upward highstand portions of cycles are best exemplied in S-2, S-5, and S-6: Abrupt shifts back from community
types 5 or 6 to type 1 are inferred to represent sediment-starved discontinuities at flooding surfaces: Minor intervals of right to left shifting of
biofacies (e.g., base of S-7) are inferred to represent transgressive systems tracts. Community types (biofacies) 3, 4, and 5A appear to be
absent in cycles above S-7. Coarse intervals occur at 400–5009, 625–10509, and 2100– 23009.

alized eurytopic species (Johnson, 1972; Walker and Al-
berstadt, 1975; Bretsky and Bretsky, 1975); as such, this
sere will be similar in many areas. If communities are
down-graded to pioneer seres by storm-generated distur-
bance of the seafloor (as in the model of Johnson, 1972),
then the average state of paleocommunities may be deter-
mined by the frequency of that disturbance. Thus, near-
shore assemblages may be dominated by ‘‘pioneer’’-type
communities, as these areas are frequently disturbed. Po-
ikiloaerobic settings, characterized by frequent fluctua-
tion of an oxic/anoxic boundary, may display similar pat-
terns (Wignall, 1994). In contrast, less disturbed, offshore,
and fully aerobic biofacies will more commonly have an op-
portunity to progress to more complex stages of coloniza-
tion (e.g., hard substrate encrusters settling on skeletons
of ‘‘pioneer colonizers;’’ Walker and Alberstadt, 1975; Kid-
well and Jablonski, 1983).
The time-averaged remains of these stable communities
will, on average, appear to be in more ‘‘mature’’ succes-
sional stages. Nonetheless, because they are effectively
random, major disturbances will still preserve earlier suc-
cessional stages occasionally (i.e., when two major storm
events happen to be closely spaced in time), and these will
resemble ‘‘nearshore biofacies.’’ True successional series,
worked out from such fortuitous samples in offshore areas,
will resemble the general pattern of time-averaged fossil
assemblages in an onshore-offshore gradient. Lateral fau-
nal tracking through transgressive sections might then
248
preserve a long-term pattern of facies replacement that
somewhat mirrors true succession, as in Bretsky and
Bretsky’s (1975) example.
Non-Tracking Species Replacements
A common perception, especially among neontologists
and paleobiologists working in the Pleistocene, is that en-
vironments are in a constant state of flux (see Bennett,
1990; DiMichele, 1994). Consequently, species associa-
tions occupying a given area are also very ephemeral. Al-
though broadly-defined biofacies may display some gross
similarities through time, local associations or communi-
ties are constantly reforming with new casts of characters
being assembled from pools of largely independent and
rather eurytopic species. Such chaotic changes in species
assemblages, or non-tracking replacements, should lead to
a rather unpredictable mosaic pattern of faunal distribu-
tion in the fossil record. A few authors have documented
patterns that might be interpreted in this way. For exam-
ple, Miller (1986) and Miller and Dubar (1988) infer that
some Neogene fossil molluscan communities represent
ephemeral groupings assembled from various species
pools following disturbances. Mosaic patterns of this type
may occur where habitats are highly patchy and/or subject
to frequent disturbances (Johnson, 1972), as in very near-
shore environments. However, even in such settings, time-
averaging may tend to smooth out the local patchiness,
leading to more predictable facies-related distributions.
A type of non-tracking replacement (‘‘long-term com-
munity succession’’) was postulated to have taken place in
‘‘unchanging’’ environments represented in the Lower De-
vonian of Oklahoma (Walker and Alberstadt, 1975). The
authors cite the occurrence of generally uniform carbonate
mudrock lithofacies as evidence for absence of environ-
mental change. However, this seems improbable given the
thickness of the unit (.30 m) and the extent of the faunal
change. Not all mudrock represents the same environ-
ment, and this situation is probably best interpreted as
faunal replacement in response to subtle environmental
changes.
Habitat Tracking
Biofacies may occupy very broad, monotonous belts
elongate parallel to shore in open shelf and epicontinental
environments. Migration of such belts in response to sea-
level fluctuations may yield a very predictable pattern of
replacement through sedimentary cycles that behaves in
accordance with Walther’s principle of facies (Anderson,
1971; Figure 4 herein). The repeated documentation of
predictable, recurrent associations of species that closely
parallel sedimentary cycles (Ziegler, 1965; Anderson,
1971; McKerrow, 1979; Cisne and Rabe, 1978) suggests
that a pattern of high-fidelity habitat tracking is typical of
many marine benthic communities. Provided that envi-
ronmental changes are not too pervasive or rapid, most
species associations appear to migrate or track their fa-
vored environment. Habitat tracking is fairly localized
(perhaps tens to a few hundreds of kilometers of migration
perpendicular to shoreline over tens to 400 Ka) for smaller
scale sea-level oscillations recorded by parasequences or
minor sequences (Elder et al., 1994). In major sea-level
BRETT
FIGURE 4—Schematic diagram illustrating hypothetical idealized pat-
terns of biofacies distribution and migration during development of a
third- or fourth-order depositional sequence. Diagram depicts a time-
space cross section across a depositional basin with a one-sided input
of siliciclastics on the east (right) side, which develops as a prograding
clastic-dominated ramp, and a carbonate ramp on the west (left) side.
Vertical axis is time. Horizontal axis is distance across the basin from
one ramp to the other (about 300 km) with basin axis at center of
diagram. Letters A–F represent distinct, depth-related biofacies for rel-
atively low-turbidity, low-sedimentation environments, with F repre-
senting deepest water biofacies, here shown in dark gray tones to
indicate anoxic to dysoxic conditions. A9 through E9 represent an al-
ternate set of biofacies developed at comparable depth ranges to A–
E but under conditions of higher sedimentation/turbidity. Vertical lines
represent hiatuses (minor disconformities) developed during different
phases of a sequence; stipple patterns indicate condensed (time-rich)
thin beds; darkest gray shading indicates deepest water facies. CB-
condensed bed associated with maximum flooding surface; PB-pre-
cursor bed (late highstand condensed bed that overlies surface of
forced regression); MFS-maximum flooding surface (a discontinuity
produced during maximum sediment starvation and/or marine erosion
associated with early highstand); SB-sequence bounding erosion sur-
face; SFR-surface of forced regression (a local non-sedimentation/
submarine erosion surface associated with abrupt lowering of sea lev-
el). Note that biofacies track preferred bathymetric/sedimentologic
conditions. During late highstand, distinctly different biofacies (E9 to
A9) are developed on the eastern ramp due to increased sedimenta-
tion rates. At lowstand, shallow-water biofacies are confined to areas
near local basin center, while erosion takes place on both basin mar-
gins. During transgression, shallow-water condensed facies typified by
the low-sediment end member biofacies (A–B) onlap sequence-
bounding unconformities on both margins due to general sediment-
starved conditions. In early highstand, relatively similar communities
are developed on both sides of the basin, while in basin center sedi-
ment starvation/submarine erosion may delete a portion of the tran-
sitional (deepening upward) record.
fluctuations, recorded by sequences, the geographic dis-
placement may amount to hundreds of kilometers, and
faunal recurrence within a particular region may be on the
order of hundreds of thousands or even a few million
years. Thus, in the middle Paleozoic of the Appalachian
Basin, certain (shallowest or deepest water) biofacies may
only appear once in each third-order sequence, and yet re-
turn with considerable fidelity. An example of this is pro-
vided by recurrence of coral beds in the Middle Devonian
Hamilton Group, where the shallowest water facies are
associated with transgressive systems tracts of each of
seven sequences. In all of these cases, however, the biofa-
cies appear to have continuously occupied particular mi-
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
grating environments; that is, the faunal gradients were
more or less static throughout the interval. However, an-
other form of very long-term recurrence apparently takes
place by reintroduction of suites of species after their elim-
ination from an entire region, and the development of en-
tirely different biofacies gradients. This form of tracking,
originally recognized by H.S. Williams (1903, 1913) in the
Devonian of the Appalachian Basin, is a distinct phenom-
enon reflecting emigration and immigration of entire fau-
nas (see below). Patterns of complete, species-by-species
tracking, in which communities recur with absolute simi-
larity, are rarely observed. Just as Walther’s ‘‘law’’ is a val-
id and highly useful generality with many exceptions in
detail, so too the notion of faunal tracking should be
viewed as a reasonable first approximation to the actual
process. In actuality, no two sedimentary cycles have de-
veloped under exactly the same conditions, so it is not sur-
prising that biofacies in distinct sedimentary cycles or
even different phases of the same cycle may differ in de-
tails of species composition, dominance, or rank position,
etc. (Bennington, 1994; Bambach and Bennington, 1996).
What is surprising is that biofacies may maintain very
similar compositions, species richness, rank ordering, etc.,
for intervals of time up to several million years.
Symmetrical Tracking
Within a single stratigraphic section, habitat tracking
may produce a predictable vertical stacking pattern of bio-
facies that appear in a nearly symmetrical cycle (Figs. 3–
5). Such ‘‘symmetrical tracking’’ patterns represent simple
lateral shifting of bathymetrically-zoned biofacies, per-
pendicular to facies strike (shoreline), in response to rela-
tive rise or fall of sea level. This simple Waltherian pattern
is assumed in models of ‘‘coenocorrelation’’ that make use
of fluctuations in fossil ‘‘communities’’ or species associa-
tion gradients to determine relative sea-level curves (e.g.,
Ziegler, 1965; Anderson, 1971; Cisne and Rabe, 1978; Cis-
ne et al., 1984; Johnson et al., 1985).
In offshore areas where the rate of siliciclastic sedimen-
tation remains fairly low throughout a sedimentary cycle,
a nearly symmetrical pattern of faunal tracking may be
observed (Figs. 4, 5). For example, many carbonate and
mixed siliciclastic-carbonate successions of the Ordovician
to Devonian in the Appalachian basin display a predict-
able spectrum of litho- and biofacies changes (Cisne and
Rabe, 1978; Brett et al., 1990). Figure 5 shows a typical cy-
cle in the Middle Devonian Hamilton Group of western
New York State (see Savarese et al., 1986), a depth-related
gradient of change from (a) black laminated shales with
sparse, low-diversity, dysoxic faunas, through (b) gray
mudstone facies with moderate-diversity assemblages of
small brachiopods, nuculid and other small mollusks, that
proceeds upward to (c) highly calcareous mudrocks and
limestones with high-diversity assemblages of corals, bra-
chiopods, bryozoans, and pelmatozoans. A ‘‘mirror-image’’
spectrum of biotas occurs in the upper or transgressive
half cycle, which is typically about as thick as the shallow-
ing portion of the cycle. Hence, species associations appear
to shift back and forth in a cyclic pattern that mimics the
sedimentary cycles.
Basinal areas may display similarly symmetrical, but
more subtle, patterns of faunal change. Such sections are
249
typically somewhat expanded as a result of entrapment of
sediments near depocenters. These sections display a
smaller range of facies change than do those representing
shallower water areas. In both transgressive and high-
stand (including regressive) phases, organisms sensitive
to higher rates of sedimentation or turbidity, such as cor-
als, may be rare or absent.
Although there are minor differences in relative fre-
quency of the most common taxa and perhaps in the ap-
pearance or absence of a few rare taxa, the assemblages on
either side of these symmetrical cycles appear remarkably
similar. Furthermore, detailed lateral correlation indi-
cates that these assemblages do in fact persist and mi-
grate in a Waltherian manner during rise and fall of sea
level. However, in areas in which erosion or siliciclastic in-
put are major factors, the sedimentary cycles and the fos-
sil communities within them may appear more markedly
asymmetrical (Figs. 3, 4, 6).
Asymmetrical Patterns: Artifacts of Preservation
A good deal of the biofacies asymmetry observed in some
sedimentary cycles may be attributed to incomplete pres-
ervation of intermediate facies. In the epicratonic seas,
where subsidence rates were low, transgressive to early
highstand systems tracts are best represented, while late
highstand (regressive) facies are largely missing (Einsele,
1985; see Figs. 3 and 4 herein). The shallowest water faci-
es, comprised of shell-rich or crinoidal transgressive lime-
stones, are abruptly juxtaposed onto early highstand faci-
es preserving a deeper-water faunal assemblage (Bayer,
Altheimer, and Deutchle, 1985; Brett and Baird, 1996; see
Fig. 4 herein). Erosion surfaces at the bases of the lime-
stones are marked by a distinctive omission suite of trace
fossils with well-preserved scratch marks indicating their
production within overcompacted muds exposed by re-
moval of considerable overburden (Kidwell and Aigner,
1985; Bayer, Alteimer, and Deutschle, 1985; Seilacher,
Reif, and Westphal, 1985; Landing and Brett, 1987). Tran-
sitional, shallowing biofacies of the late highstand in the
underlying sequence are absent because they have been
removed by erosion associated with a lowstand maximum
at the sequence boundary (Fig. 4). However, there are
hints that the actual pattern of faunal tracking was nearly
symmetrical. Where remnants of the truncated shallow-
ing-upward succession are preserved below a sequence-
bounding erosional surface, they appear very nearly iden-
tical to the analogous facies of the more completely pre-
served transgressive systems tract.
Elsewhere, particularly in siliciclastic-dominated shal-
low-water successions, portions of a tracking biotic spec-
trum may be lost at the surface of maximum starvation,
through extreme condensation or even submarine erosion
(Figs. 3, 4). This again imposes apparent asymmetries on
the pattern of vertical faunal replacement. A good illustra-
tion of this is found in Bretsky and Bretsky’s (1975) de-
tailed study of Late Ordovician faunas in the Nicolet River
Valley of Quebec. Figure 3 shows several ‘‘successional cy-
cles’’ (cycles of faunal replacement) that appear markedly
asymmetrical due to the absence of intermediate stages
(or biofacies). These patterns can be interpreted as pro-
gressions of community replacement through several
shallowing-upward cycles or parasequences. The absence
250 BRETT

FIGURE 5—Symmetrical community replacement (tracking) patterns in Middle Devonian biofacies. Representative cycle in distal, relatively
sediment starved position, based on Centerfield Limestone of western New York State. Note nearly mirror image ordering of biofacies through
a shallowing-deepening cycle ( late highstand portion of one depositional sequence and transgressive systems tract of overlying sequence)
Block diagrams illustrate biofacies or fossil associations plus typical bed stratinomy. (a) Dysoxic, low diversity leiorhynchid brachiopod and
cephalopod association in dark gray to black, laminated shale; (richness, R 5 5–10 species); (b,f) Moderate diversity (R 5 20–30 species)
Ambocoelia association, in medium gray mudstone, dominated by small brachiopods and molluscs; (c, e) Diverse brachiopod and small rugose
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
of mirror-image deepening series probably reflects sedi-
mentary condensation at the flooding surfaces of each se-
quence. Again, the asymmetrical pattern may be more ap-
parent than real (Holland, 1995).
Asymmetrical Patterns: Biotic Causes
True biotic asymmetries in tracking result from the ex-
pected variation of sedimentation rate throughout a depo-
sitional sequence. Complex, along-strike shifting of biofa-
cies may record the interplay of bathymetric and sedimen-
tologic/substrate factors that control organism distribu-
tion. Specifically, where sedimentation rates vary
markedly in shallowing (regressive) versus transgressive
phases of a cycle, producing asymmetrical parasequences
or sequences, faunal tracking patterns are complex and
asymmetrical (Rollins et al., 1979; Holterhoff, 1992).
For example, in depositional environments proximal to
sediment sources, cycles may be manifested as distinctly
upward-shallowing progradational packages that com-
mence with dark silty shales and pass upward through
relatively thick successions of bioturbated silty mudstone,
siltstone, and sandstones (Figs. 4, 6). In these areas, the
succession of fossil assemblages of the late highstand (re-
gressive) systems tract is quite distinct from that found in
the coeval thinner sections more distal to the siliciclastic
source area. Moreover, the pattern of faunal replacement
in the late highstand (regressive) facies is distinctly asym-
metrical with respect to that in the transgressive systems
tract. Epifaunal and infaunal deposit-feeding organisms
and their traces tend to dominate the regressive sedi-
ments. This presumably results from the greater avail-
ability of particulate food within the finer-grained por-
tions of the siliciclastic sediment, as well as a higher de-
gree of substrate instability (Fürsich, 1978; Scott, 1978;
Elder, 1987). Bivalves, and quasi-infaunal brachiopods, as
well as gastropods and certain trilobites, may occur abun-
dantly in these mudrocks (Fig. 6).
The shallowest portions of siliciclastic-dominated se-
quences commonly display hummocky cross stratification,
ripple marks, and other sedimentary structures that at-
test to physical processing of the sediments by storm
waves and currents (Fig. 6). Correspondingly, less intense
bioturbation is characteristic of these portions of these cy-
cles. Vertical traces, such as Skolithos, may be predomi-
nant in certain layers, reflecting minor cutoffs in sedimen-
tation. Coquinites of low-diversity, thick-shelled epiben-
thos may occur as discontinuous lenses within the upper
storm-dominated portion of many sedimentary cycles.
The biofacies of the transgressive portion of the cycle
typically resemble those in the symmetrical sediment-
starved cycles distal to terrigenous source areas (Fig. 4;
compare Figs. 5 and 6). They may include a wide array of
epifaunal species that are absent or exceedingly rare in
←
coral association in calcareous mudstone (R 5 40–50 species); and (d) Highest diversity (R 5 60–100 species) large rugose coral association;
note abundant bryozoans, brachiopods, molluscs and other taxa. Key to lithologies—(A) Limestone, fossiliferous wacke- to packstone, (B) gray
mudstone, (C) dark gray to black shale, (D) gray calcareous mudstone, (E) coarse siltstone and sandstone, (F) tabulate corals, (G) rugose
coral, and (H) brachiopod shell beds. Symbols for sequence stratigraphy (to left of stratigraphic column):EHST—(early) highstand systems
tract; PB—precursor bed (see text); LHST—regressive (late highstand) systems tract; SB—sequence boundary (here nearly conformable);
TST—transgressive systems tract; MFS—maximum starvation (flooding) surface.
251
the regressive half cycle. The matrix of these beds is typi-
cally a fine-grained, slightly calcareous mudstone, al-
though thin skeletal-hash beds or even semi-continuous
fossiliferous limestones may occur in the basal transgres-
sive systems tracts of these cycles.
These faunal asymmetries clearly reflect a distinct in-
equality of sediment supply in the regressive, as opposed
to transgressive, phase of the cycle (Fig. 4). Sedimentation
rates are higher and tend to increase during the late high-
stand as siliciclastic wedges prograde offshore, either in
response to, or as a cause of shallowing. Conversely, silici-
clastic sedimentation rates are lower and tend to decrease
during transgressions. Only fine-grained, suspended sili-
ciclastics are transported any distance offshore. Thus, the
transgressive half cycle appears sedimentologically and
faunally far more consistent over a broad area than does
the regressive half. Organisms that are sensitive to or ex-
cluded by high rates of sedimentation may appear even in
areas relatively proximal to the terrigenous source regions
during this time. The fact that two distinct types of biofa-
cies may occupy comparable bathymetric zones, and yet
each recur with considerable fidelity, suggests that these
biofacies are displaced laterally (along strike), tracking
variations in sedimentation as well as depth (Fig. 4).
ECOPHENOTYPIC CHANGE THROUGH
SEDIMENTARY CYCLES
Certain broadly adapted, eurytopic organisms may re-
main in a particular geographic area but display some
morphological modification in response to local environ-
mental change. Typically, this ‘‘modification’’ involves a
non-evolutionary, ecophenotypic response. In actuality,
this may involve a non-tracking response to environmen-
tal change, or ecomorphs of a species may themselves be
tracking environments.
A number of studies have sought to relate repetitive cy-
cles of morphological change to cycles of sedimentary en-
vironmental change. Such studies are not without difficul-
ties, however. Kidwell and Aigner (1985) discuss the prob-
lems of discriminating ecophenotypic variation in con-
densed sections, wherein mixing of remains of several
generations of organisms and, in some cases, those repre-
senting different environments may occur within the
same bed. Interpretation of ecophenotypic change, as op-
posed to microevolution, may also be ambiguous. Kelly
(1983) documented increasing mean values of some five of
eight variables in the bivalve Lucina anodonta from deep-
ening-upward successions in Miocene shell beds from
Maryland. Kelly suggested that the progressive change
represented a case of micro-evolution in an otherwise gen-
erally stable assemblage of fossils. However, as a similar
pattern repeats through three cycles, represented by ma-
jor shell beds, Kidwell and Aigner (1985) concluded that it
252 BRETT

FIGURE 6—Asymmetrical community replacement (tracking) in proximal correlative of same cycle illustrated in Figure 4 (based on Chenango
Siltstone Member in central New York State). Note that biofacies in middle regressive facies are dominated by low-diversity brachiopod and
mollusc faunas with abundant Zoophycos. Also, biofacies of analogous depths in transgressive phase are distinctly more diverse and closely
resemble those of the more sediment starved cycle shown in Figure 5. Fossil associations include: (a) Small brachiopod-dominated, moderate
diversity (R 5 20–30 species) Ambocoelia association with local thickets of auloporid corals found in precursor bed; (b) lower diversity (R 5
10–20 species) small brachiopod and mollusc-dominated association in silty mudstone; (c) patchy, moderate diversity (R 5 30–50 species)
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY 253

is more likely to represent a case of ecophenotypic change

that was reversible and repetitive through sedimentary
cycles.
A particularly fruitful line of investigation is the mor-
phological response of sedentary organisms to changes in
substrate that vary predictably in small-scale cycles or
parasequences. Seilacher, Matyja and Wierzbowski (1985)
and Bayer, Johnson and Brennan (1985) observed parallel
ecophenotypic gradients within Jurassic and Pleistocene
oysters that neatly parallel small-scale shallowing-up-
ward cycles (Fig. 7). Morphological trends, including flat-
tening and broadening of the shells, in each case reflect
the development of firm, sometimes shell-rich sediment
with decreasing mud content as a result of an increased
frequency of disturbance and winnowing events near the
tops of the cycles (Bayer, Alteimer and Deutchle, 1985).
Similarly, Kidwell and Aigner (1985) discussed a case of
variation and morphology of the Eocene clam Carolinia
placunoides, which apparently reflects alternation of colo-
nization strategies. Hard substrate encrusting forms were
broad and flat, while deeper, cup-shaped forms developed
stacks of mutually supported shells on soft substrate. Al-
exander (1975) observed similar ecophenotypic variation
in shells of Late Ordovician strophomenid brachiopods
from Ohio. Broadening and flattening of shells was char-
acteristic of offshore, muddy environments, while more
strongly concavo-convex morphotypes typified firmer sed-
iments of cycle caps.
In these cases, the skeletal morphology displays a con-
sistent trend of change through a small-scale sedimentary
cycle, which is readily related to the dynamics of sediment
input and winnowing and the development of variable FIGURE 7—Ecophenotypic changes in Jurassic (left) and Pleistocene
substrate consistencies. In other instances, repetitive eco- (right) oysters in response to increased substrate firmness in small-
morphological cycles may parallel sedimentary cycles scale shallowing upward cycle (from Seilacher, Matyja, and Wierz-
without being so obviously related to functional con- bowski, 1985).
straints. Bayer (1970) and Bayer and McGhee (1985) re-
port consistent ecomorphological gradients in genera of
Jurassic ammonites; smooth, evolute morphs typify off- versely, ‘‘outages’’ represent anomalous rarity or absence
shore claystone facies near bases of cycles, whereas evo- of a normally common species (Brett and Baird, 1997).
lute, highly sculptured forms occur in oolitic limestones, Ecological epiboles represent genuinely biological phe-
and spheroid forms with tricarinate keels characterize nomena in which conditions temporarily become condu-
condensed limestones. While the causes of such morpho- cive for colonization or proliferation of a species that is
logical response remain ambiguous, especially in nekto- normally rare within the basin. The species may flourish
benthic organisms, nonetheless, they may provide impor- over a wide area during these times. Particularly problem-
tant links between organism paleoecology and changing atical is the fact that these epiboles also may involve cross-
sedimentary environments. cutting relationships among facies. In outages, the oppo-
site situation pertains; some factor in the environment in-
ECOLOGICAL EPIBOLES AND OUTAGES hibits or locally eliminates a species that ordinarily is com-
mon or even dominant.
Ecological change in response to changing environmen- Certain taxa may display multiple epiboles, each occur-
tal parameters may involve simple changes in relative ring in analogous phases of sedimentary cycles, albeit not
dominance of species. For example, a normally rare spe- in each successive cycle. This indicates that conditions
cies becomes common and/or widespread in a thin strati- conducive to proliferation of the species are associated
graphic interval, forming an ‘‘ecological epibole.’’ Con- with certain combinations of environments that do not re-

←
Tropidoleptus brachiopod-dominated association (note also local erect bryozoans and pelmatozoans, interfingering with low diversity [5–15
species] Zoophycos dominated silty mudstone); (d) Low diversity (5–15 species) Allanella brachiopod and bivalve association in laminated
siltstone or fine-grained sandstone; (e) moderate to high diversity (R 550–100 species) large coral and pelmatozoan association in thin skeletal
hash limestone overlying sequence boundary; and (f) diverse brachiopod and small rugose coral association in calcareous mudstone (R 5
30–50 species); compare with associations c and e in Figure 5. Symbols for lithology and sequence stratigraphy as in Figure 5. Figure adapted
from Savarese et al. (1986) and Brett et al. (1990).
254
cur with every cycle. In mid-Paleozoic faunas, epiboles
tend to occur particularly during the transgressive sys-
tems tracts of third- and fourth-order sequences. The re-
gressive (late highstand) portions of cycles tend to be char-
acterized by eurytopic faunas that recur very predictably
in each cycle. In contrast, unusual environments associat-
ed with the development of variably sediment-starved
conditions may permit temporary proliferation of steno-
topic species that are generally rare. Ecological epiboles
and outages presumably reflect slight differences within a
generally persistent environment that favor or inhibit a
few species while leaving the majority of the fauna unaf-
fected. In most cases, the environmental perturbations re-
sponsible for epiboles or outages are not apparent from
sedimentary facies. However, clues as to causative agents
may be sought in isotopic or trace-element geochemistry.
REPLACEMENT BY LARGE-SCALE IMMIGRATION
Not all faunal replacements involve shifting or prolifer-
ation of indigenous species. If, and only if, replacement by
ecological tracking of local faunal elements can be ruled
out, it may be possible to discern patterns that are gener-
ated by immigrational and/or evolutionary processes in re-
lation to depositional sequences.
In some instances, faunal change involves influx of ‘‘ex-
otic’’ species (incursion epiboles of Brett and Baird, 1997)
or wholesale incursion of an entire fauna from another ba-
sin or province. The latter will commonly occur in associ-
ation with local extinction of many indigenous species.
However, this is not necessarily true. For example, in the
case of the Richmondian invasion, a number of inverte-
brate species were added to each of several biofacies in the
Upper Ordovician of the Cincinnati arch region without
loss of many indigenous taxa (Patzkowsky and Holland,
1996; Holland, 1997).
Episodic immigration events are particularly typical of
nektonic forms, such as ammonoids, and benthic forms
with planktonic larvae. Notable examples include wide-
spread ammonoid incursions that are closely associated
with marker beds in the Jurassic of Germany (see Bayer,
Johnson and Brennan 1985; Figure 8). Such incursions
may reflect unusual sedimentary conditions; thus, the de-
velopment of widespread shell gravels in the Jurassic ba-
sin seem to have permitted the influx of hard substrate en-
crusters as well as certain nektobenthic ammonoids (Bay-
er and McGhee, 1985; Bayer, Johnson and Brennan,
1985). Similarly, Hagdorn (1982, 1985) demonstrated that
widespread beds of brachiopods and crinoids typically oc-
cur at the caps of shallowing-up carbonate cycles within
the Triassic Muschelkalk Limestone of Germany. Cemen-
tation of the substrate associated with marine flooding
surfaces developed hardgrounds that permitted the exten-
sive influx and colonization of species that required hard
substrates for attachment. These species were normally
restricted to shoal areas on the margins of the Tethys sea.
Altered circulation and climatic patterns during sea-
level highstands may promote major immigration epi-
sodes. For example, the abrupt appearance of a diverse
fauna within a condensed concretionary limestone in the
Upper Cretaceous Graneros Shale may be related to the
influx of a warm subtropical water mass into the Western
Interior Basin during a major eustatic sea-level rise
BRETT
(Kauffman, 1986). Normally isolated biogeographic prov-
inces may also be connected during maximum flooding
events, allowing the transfer of larvae between provinces
(Eicher and Diner, 1985; Kauffman, 1986).
The existence of ‘‘recurrent’’ faunas separated by hun-
dreds of meters of strata bearing completely dissimilar
faunas indicates a long-term form of faunal migration, as
noted around the turn of the century by Williams (1903,
1913). In his study of recurrent Tropidoleptus or Hamilton
faunas, Williams observed that an entire suite of species
may recur long after their apparent extinction in a region.
This indicates a process similar to, but distinct from habi-
tat tracking, in that the recurrent fauna or biofacies seems
to have been replaced by another suite of tracking species
in its appropriate sedimentary environments, as inferred
from position in cycles and lithofacies. Such faunal recur-
rence may involve displacement and reintroduction of en-
tire or partial biofacies from a region within the basin not
generally represented by exposed strata, or another depo-
sitional basin or province. This faunal outage and recur-
rence is probably analogous to the phenomenon identified
as species outages, except that it involves wholesale mi-
gration of entire faunas or biofacies. Recently, Patzkowsky
and Holland (1996) and Holland (1997) have documented
a similar pattern within the Middle and Upper Ordovician
of eastern North America. The ‘‘Richmondian invasion’’
(Holland, 1997) involved the reappearance of a number of
lineages that were displaced from the region of the then-
tropical area of present day Arctic Canada. Evidently, lon-
ger-term secular variations in climate and/or water chem-
istry forced these organisms out of eastern North America
coincident with a second tectophase of the Taconic Oroge-
ny. However, later, when oceanographic conditions were
restored to more nearly their pre-Taconic condition, some
of the lineages were able to reinvade.
Long-term faunal recurrence appears to be a response
to factors other than simple sea-level variation, such as cli-
matic changes. As with incursion epiboles, preliminary ob-
servations indicate that major faunal migrations are as-
sociated with highstands of major second-order sequences,
typically occurring just above major black shales (Brett
and Baird, 1995, 1997). Thus, the periods of faunal outage
in a region may be initiated by particularly extreme sea-
level rises that coincide with climatic change. During
these times, shallower-water biofacies are displaced great
distances by faunal tracking and are introduced into a
new region. At the same time, climatic or other non-depth
related factors become sufficiently altered in the basin of
faunal origination such that, when appropriate bathymet-
ric conditions return to the area, it is no longer hospitable
to many of the original suite of species, which become ex-
tinct, at least locally. A new fauna appears in most biofa-
cies, derived from tolerant holdovers, newly evolved deriv-
atives of these endemics and, in some instances, a large
proportion of immigrant taxa, themselves displaced by the
same event from their original provinces. In some cases,
later environmental changes may partially reverse the
pattern, enabling some of the displaced faunas to reinvade
their original territory.
The phenomenon of large-scale faunal migrations clear-
ly requires more in-depth study. It may be at the root of
some of the abrupt faunal overturns that bound blocks of
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY 255

relative stasis or ecological-evolutionary subunits docu-

mented by Brett and Baird (1995; see below).

ITERATIVE EVOLUTION AND SEDIMENTARY

CYCLES
Stasis within and between sedimentary cycles (e.g., Hal-
lam, 1978) suggests that anagenetic change is an excep-
tional response to fluctuating sea level. Sheldon (1993)
has argued that anagenesis may be characteristic of sta-
ble, monotonous environments, whereas those with fre-
quent fluctuations favor stasis. Presumably, organisms
must remain relatively unspecialized to cope with envi-
ronmental fluctuations. However, some case studies have
documented long-term anagenetic changes that appear to
relate to sedimentary sequences.
The occurrence of iterative evolutionary patterns within
several sedimentary cycles offers the potential for testing
for relationships between evolutionary pattern and envi-
ronmental change (Fig. 8). For example, Bayer and Mc-
Ghee (1985) observed iterative cycles of heterochronic
morphological change (peramorphosis) in Middle Jurassic
ammonites from Klupfel-type sedimentary cycles in Ger-
many. Small, ribbed, evolute, and highly variable ammo-
nites typical of the heterolithic, storm-dominated facies of
transgressive portions of the cycles were replaced by large,
involute, and less variable forms in ‘‘monotonous’’ mud-
rocks in the highstand portion of each cycle. The authors
attribute the initial small, paedomorphic forms to adapta-
tion in stressed environments near the regressive maxi-
mum or lowstand; local heterogeneities favored a high de-
gree of morphological variability. Conversely, more uni-
form conditions of the highstand favored the less variable
FIGURE 8—Recurring lineages of phylloceratid and lytoceratid am-
peramorphic forms. This pattern agrees with the predic- monites in the Jurassic of the South German Basin in relation to global
tion of Hallam (1978) that evolution of ammonoid species sea-level (curve after Hallam, 1978). Note that several species display
during regressions involved r-selection, favoring paedo- abrupt appearances in the basin (incursions) during second-order sea-
morphs, while k-selection operated during transgressions. level highstands (from Bayer and McGhee, 1985).
With one exception, Bayer and McGhee (1985) observed
that the founders of these lineages did not arise within the
basin, but immigrated from Tethyan sources episodically
during sea-level lowstands. Many authors have documented correlations between
Based on this case study, Bayer and McGhee (1985) also large-scale extinctions/evolutionary radiations and fluc-
proposed a general model for evolution related to sea-level tuations in relative sea level (Moore, 1954; Newell, 1967;
fluctuation in local basins. They postulated that restric- Johnson, 1974; Ramsbottom, 1981; Jablonski, 1980, 1985,
tion and fragmentation of habitat area occurred during re- 1986; Fischer, 1986; Hallam, 1987, 1989, 1992). There are
gressions (lowstands), leading to preferential extinction a number of theoretical arguments for such a relationship,
and replacement of offshore species. Bayer and McGhee mainly related to species diversity-habitat area relation-
postulate that, if the basin was isolated, in situ evolution ships derived from study of island biogeography (MacAr-
might be accelerated at this time by allopatric speciation. thur and Wilson, 1967; Simberloff, 1974; Schopf, 1974).
Similarly, Kauffman (1977) postulated high rates of spe- The most obvious connection is undoubtedly that of sea-
cies evolution during shallowing intervals. However, Bay- level fall and major extinction. As a result of the area effect
er and McGhee also suggest that, if a basin had connec- (Simberloff, 1974; Schopf, 1974), substantial drops in sea
tions to other areas, replacement by immigration of ‘‘prea- level may produce major habitat-area reduction and stress
dapted,’’ exotic species might preclude local evolution be- in marine communities, resulting in mass extinctions. Ja-
cause immigration rates would far exceed evolutionary blonski (1986) and Hallam (1989, 1992) have stressed that
rates. This model has very important implications for pat- sea-level drop is the most pervasive environmental corre-
terns of extinction and speciation in stratigraphic se- late of major mass extinctions. Many of the larger mass ex-
quences. It needs to be very carefully tested by detailed tinctions show some element of association with sea-level
studies of faunal patterns in local basins. drop. Sea-level lowstands and regression should be ex-
pected to produce major erosional surfaces that bound
MACROEVOLUTION AND SEQUENCE large-scale depositional sequences. Several of the major
STRATIGRAPHY crises in life history, notably the late Ordovician (Ashgill),
Conditions responsible for producing sedimentary se- Permo-Triassic, and Cretaceous-Tertiary extinctions are
quences may also be tied to major evolutionary bioevents. associated with very major drops in global sea level. Such
256
extinctions are, thus, associated with major sequence
boundaries of second- or third-order cycles.
However, few of the largest sea-level drops, such as
those that form Sloss’s (1963) megasequence boundaries,
are associated with mass extinctions. Many other third-or-
der unconformities recognized by seismic stratigraphers,
such as the very major Oligocene sea-level lowstand, also
do not coincide with mass extinction. Jablonski (1985) ar-
gued that even complete draining of modern continental
shelves would not produce major extinction because off-
shore oceanic islands would still harbor plentiful species
from which continental shelf seas could be repopulated.
Hence, even rather major regressions may not directly im-
pact organisms as predicted by the area-effect model.
However, Johnson (1974) has stressed that, following
times of extensive cratonic flooding by shallow epiconti-
nental seas, organisms may suffer far greater extinction
during sea-level drop. Not only do relatively minor sea-lev-
el lowstands expose vast areas, but under these conditions
organisms may have become specifically adapted to local
conditions in the epeiric seas; such ‘‘perched faunas’’ may
be especially vulnerable to habitat loss. Thus, conditions
prior to the regression may be most critical in determining
its effect. Moreover, some sea-level drops may rather be
associated with major climatic change, especially cooling,
which actually produced the observed extinctions (Stan-
ley, 1984, 1988). Evolution of widespread, stenotopic cool-
water faunas may also occur during ‘‘icehouse’’ times or
associated with major regressions. This phenomenon is
well exemplified by the widespread Late Ordovician Hir-
nantia fauna that proliferated after the Ashgill extinc-
tions (Brenchley, 1989).
Conversely, a number of studies have pointed to the as-
sociation of transgressions and evolutionary radiations of
shallow-marine organisms (Moore, 1954; Kauffman, 1977;
Hallam, 1978; House, 1985). Flooding of continental areas
by shallow seas should increase habitable ecospace and fa-
vor evolution of stenotopic organisms (cf., the ‘‘perched
fauna’’ hypothesis of Johnson, 1974). Warming global tem-
peratures and weakening climatic gradients during trans-
gressions also may favor evolution of stenotopic faunas
(Kauffman, 1977; Fischer, 1986).
Several authors have documented a relationship be-
tween punctuational faunal changes and sea-level fluctu-
ations. For example, Hallam (1987, 1992) has documented
a close correspondence of extinctions with sea-level low-
stands and radiations with transgressions in Early and
Middle Jurassic ammonite families. Likewise, Dockery
(1986) relates major overturns in Paleogene molluscan
faunas with regressions that produced sequence bound-
aries in the Gulf Coastal area. Although Hansen (1987)
claimed no correlation between molluscan extinction and
sea-level for the same strata, a pattern of foraminiferal ex-
tinctions with sequence boundaries comparable to Dock-
ery’s molluscs was observed by Gaskel (1991). However,
the sharpness and apparently synchronous nature of
these extinctions may be partly an artifact of the unconfor-
mities (Holland, 1995). Bayer and McGhee (1985) docu-
mented repeated patterns of local extinction, followed by
in situ evolution and immigration, during lowstands in
Mesozoic third-order cycles.
On the other hand, other studies have revealed no rela-
tionship between faunal change and sea-level fluctuation.
BRETT
For example, Leckie et al. (1992) found that important
turnover events in Cretaceous plankton and reefs tended
to occur within sequences and not at their boundaries.
McGhee (1991, 1992) notes that most previous studies re-
lating evolutionary patterns to sea level have been quali-
tative. In detailed statistical studies, McGhee compares
rates of extinction and diversification for Devonian bra-
chiopods in New York to the sea-level curves of Johnson et
al. (1985). He concludes that there is no correlation be-
tween relative sea-level change (McGhee, 1991) or rate of
sea-level change (McGhee, 1992) and patterns of extinc-
tion or diversification (although the data on which this
study was based [Dutro, 1981] are incomplete, and, as not-
ed below, some faunal overturn events in the Appalachian
basin do appear to be linked with transgressions). Clearly,
there is a need for many more such critical studies. Thus,
despite probable correlation in some cases, no simple, pre-
dictable relationship exists between sea-level lowstands,
sequence boundaries, and mass extinctions.
A further relationship between sea-level effects and
macroevolution involves major transgressions and extinc-
tions. Episodes of widespread bottom-water anoxia in
shallow seas and oceanic anoxic events (OAE’s; Arthur
and Schlanger, 1979) very commonly correlate with maxi-
mum sea-level highstands (Hallam and Bradshaw, 1979;
Wignall and Hallam, 1991; Wignall, 1994). These anoxic
events may cause waves of extinction that propagate from
offshore basinal areas to shallow shelves as the extent of
the anoxic water column expands (Hallam, 1987, 1992).
Area-reduction effects again pertain, as major tracts of
shallow seafloor become anoxic and uninhabitable. How-
ever, in this case, offshore low-oxygen adapted organisms
may show differential survivorship. These extinctions
would be predictably associated with maximum flooding
surfaces or condensed sections.
Excellent examples of such highstand-related extinc-
tions are documented by Kauffman (1984) and Elder
(1987) for the Cretaceous Cenomanian-Turonian and Tu-
ronian-Coniacian extinctions. In both cases, major extinc-
tion of even eurytopic forms was associated with an anoxic
event during or just before peak transgression. The extinc-
tions followed periods of equable conditions in the early- to
mid-transgression, which favored establishment of high-
diversity faunas. These extinctions were then followed by
major faunal overturns and further extinctions coinciding
with stresses during regression. Kauffman (1984) noted
the importance of ameliorated conditions prior to the an-
oxic events that ‘‘set the faunas up for the kill.’’ Although
differing in details, both the Albian-Cenomanian and ter-
minal Cretaceous extinctions followed somewhat similar
patterns (Kauffman, 1984).
A comparable pattern has been found for local extinc-
tions and faunal turnover events in the Devonian of the
Appalachian Basin (Brett and Baird, 1995). For example,
the demise of the Eifelian Onondaga fauna and appear-
ance of the well-known Givetian Hamilton fauna is asso-
ciated with major eustatic sea-level rise and an interval of
widespread anoxia (Brett and Baird, 1996); this coincides
with the global Kacak-otomari bioevent (Walliser,
1986a,b, 1990; Boucot, 1986; 1990a,b). Similarly, the
abrupt loss of much of the Hamilton fauna occurs with an-
other eustatic highstand and widespread low oxygen con-
ditions, and coincides with the global Pharciceras bioevent
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
(House, 1985; Boucot, 1990b). In each case, a major eus-
tatic rise and anoxic event is associated with local exter-
mination of more than 70% of species, accelerated in situ
evolution, and immigration. The Kacak event in the Ap-
palachian Basin follows a pattern similar in detail to the
Cenomanian-Turonian event documented by Kauffman
(1984), in showing the introduction of a temporary, prob-
ably warm-water fauna at or near peak transgression that
was subsequently extinguished and replaced by stable
fauna. Likewise, the Frasnian-Fammenian extinction has
been associated with two intervals of widespread anoxia—
the lower and upper Kellwasser events (Schindler,
1990)—perhaps but not unquestionably associated with
maximum flooding surfaces (see McGhee, 1992).
The situation is clearly complex and not entirely pre-
dictable, but a number of faunal overturns do coincide
with lowstands or transgressive maxima. However, each
case must be carefully evaluated on its own merits. Far
more detailed studies, such as those of Kauffman (1984;
1986) and Elder (1987), must be done before any more de-
finitive generalizations can be made. However, at least the
sequence model provides an important framework within
which to test these ideas.
CONCLUSIONS
(1) Paleoecology has a dual relationship with sequence
stratigraphy. Fossils provide very important tools for
reconstructing patterns of environmental change that
relate to sequence-forming processes; and sequence
stratigraphy provides a predictive framework that fa-
cilitates recognition and interpretation of patterns of
biotic change in the geologic record.
(2) Paleontological cues to sequence interpretation in-
clude several parameters. Fossils provide indicators of
absolute and relative depth; fluctuations in body and
trace fossil associations or ‘‘communities’’ may permit
recognition of cycles even in seemingly monotonous
mudrock facies. Certain types of organisms are sensi-
tive indicators of low turbidity and sedimentation
rates; trace fossil concentration and crosscutting rela-
tionships may aid in recognition of condensed inter-
vals and submarine erosion surfaces. Variations in
substrate consistency, that may, in turn, relate to
changes in sedimentation rate, can be judged from the
occurrence of fossil taxa known to be substrate selec-
tive. Traces and body fossils also may elucidate fluc-
tuations in benthic oxygenation that relate to sea-level
variation.
(3) Temporal variations in fossil assemblages (‘‘commu-
nities’’) can be interpreted within the framework of se-
quence stratigraphy. Long-term (10’s to 100’s of Ka)
community replacement should be distinguished from
short-term ecological succession, although these two
processes, operating at different scales, may display
significant parallels that relate to position within on-
shore-offshore gradients of disturbance frequency.
(4) The recurrence of highly similar fossil assemblages in
analogous portions of sedimentary cycles indicates
that associations of species that require particular
combinations of depositional environments persist
with relatively little change through long intervals of
time. These associations appear to shift laterally over
257
considerable distances during intervals ranging from
a few thousands to perhaps a few millions of years.
Rather than adapting to the stress of changing envi-
ronments, marine organisms most commonly appear
to track their favored environments. Provided that the
lateral migration of these environments is not too rap-
id, or that these environments do not disappear alto-
gether from the local basin, most organisms appear to
be able to keep up with the shift of environments pro-
duced by sea-level fluctuation.
(5) Similar rates of sediment accumulation during trans-
gressions and highstands (including regressions) may
lead to nearly symmetrical cycles of faunal habitat
tracking. Many asymmetries in tracking can be readi-
ly interpreted as either artifacts of incomplete preser-
vation of facies transitions (e.g., at sequence bound-
aries or surfaces of maximum starvation), or as real bi-
otic responses to fluctuating sedimentation rates, tur-
bidity, and substrates within sedimentary cycles.
Sedimentation rates are much lower in proximal areas
during transgressive phases, due to coastal sediment
entrapment, than during late highstands or regres-
sions. Fossil assemblages vary accordingly. Low sedi-
mentation conditions or special substrates formed in
condensed intervals may permit proliferation of nor-
mally rare species, leading to ecological epiboles.
Moreover, some eurytopic species may not track but
remain in a particular area and show ecophenotypic
responses, for example, to changing substrate condi-
tions.
(6) Some faunal replacements involve immigration of spe-
cies into a local basin from other basins or provinces.
These incursions may be short-lived (but geographi-
cally widespread) events, involving a few species (in-
cursion epiboles), or wholesale influx of ‘‘exotic’’ asso-
ciations. In both cases, there appears to be an associa-
tion with transgressive highstands. This relationship
may involve climatic changes (typically warming) in-
directly related to the highstand, or the opening of dis-
persal routes because of high sea-level stands. How-
ever, in some cases, incursions may also be associated
with unusual conditions during lowstands.
(7) In situ evolution is a less common response to se-
quence-generating environmental changes. Lateral
shifting of tracking biotas, or incursion of immigrant
species, appears to occur at a much higher rate than
evolutionary adaptation. However, Bayer and Mc-
Ghee (1985) postulate that allopatric speciation may
result from habitat fragmentation during regressions
in some isolated basins.
(8) Patterns of large-scale evolutionary radiation and
mass extinction may be related in complex ways to
large-scale fluctuations of sea level and related effects
that also generate depositional sequences. Many em-
pirical and theoretical studies link extinctions to re-
duction in habitat area or climatic stress (typically
cooling) during major lowstands. Transgressions gen-
erate increased habitat space and ameliorated cli-
mates that may favor evolutionary radiations. How-
ever, loss of habitable seafloor may result from the
spread of bottom-water anoxia during transgressions,
and produce waves of extinction from deep-to-shallow
water environments. Hence, bioevents commonly may
258
be associated with either sequence boundaries or sur-
faces of maximum flooding. Indeed, major stable eco-
logical-evolutionary units may be bounded at strati-
graphically important discontinuities. However, some
studies have found little or no relationship between
macroevolution and sea-level fluctuations, and many
of the largest sequence-bounding unconformities and
flooding surfaces bear no evidence of dramatic bioev-
ents. The relationship between sequence stratigraphy
and macroevolution is obviously complex and multi-
factorial. Nonetheless, it warrants considerably more
detailed quantitative study.
ACKNOWLEDGMENTS
Dave Bottjer invited me to submit an article on se-
quence stratigraphy and paleontology to PALAIOS for the
tenth anniversary issue. The subject proved to be vast,
and, ultimately, that article had to be shortened; I very
much appreciate Dave’s willingness to consider splitting
the article into two natural divisions. The first, dealing
with the relationships of biostratigraphy and taphonomy
to sequence stratigraphy, appeared in the anniversary
volume. The present paper represents the paleoecological-
evolutionary ‘‘half’’ of my original manuscript, substan-
tially modified and improved by several perceptive re-
views. I thank Pete Allison, Art Boucot, Bob Gastaldo,
Steve Holland, Brad Sageman, Charles Savrda, and
Chuck Ver Straeten, among others, for their many excel-
lent and constructive suggestions regarding this manu-
script. Lars Olsen helped in preparation of two figures. My
work on sequence stratigraphy and paleontology has been
supported by NSF grants EAR 8313103 and 9219807, and
grants from donors to the Petroleum Research Fund,
American Chemical Society.
REFERENCES
ABERHAN, M., 1994, Guild-structure and evolution of Mesozoic ben-
thic shelf communities: PALAIOS, v. 9, p. 516–545.
ADLIS, D.S., GROSSMAN, E.L., YANCEY, T.E., and MCLERRAN, R.D.,
1988, Isotope stratigraphy and paleodepth changes of Pennsylva-
nian cyclical sedimentary deposits: PALAIOS, v. 3, p. 487–506.
ALEXANDER, R.R., 1975, Phenotypic lability of the brachiopod Rafines-
quina alternata (Ordovician) and its correlation with the sedimen-
tologic regime: Journal of Paleontology, v. 49, p. 607–618.
ANDERSON, E.J., 1971, Environmental models for Paleozoic commu-
nities: Lethaia, p. 287– 302.
ARTHUR, M.A., and SCHLANGER, S.O., 1979, Cretaceous ‘‘oceanic an-
oxic events’’ as causal factors in the development of reef-reservoir
giant oil fields: American Association of Petroleum Geologists Bul-
letin, v. 63, p. 870–885.
BAMBACH, R.K., and BENNINGTON, J.B., 1996, Do communities
evolve? A major question in evolutionary paleoecology: in ERWIN,
D.J., JABLONSKI, D.J., and LIPPS, J., eds., Evolutionary Paleobiol-
ogy: Essays in Honor of James W. Valentine: University of Chica-
go Press, Chicago, p. 123–160.
BATT, R.J., 1993, Ammonite morphotypes as indicators of oxygena-
tion in a Cretaceous epicontinental sea: Lethaia, v. 26, p. 49–62.
BAUM, G.R., and VAIL, P.R., 1988, Sequence stratigraphic concepts
applied to Paleogene outcrop, Gulf and Atlantic basins: in WILGUS,
C.K., HASTINGS, B.S., ST. C. KENDALL, C.G., POSAMENTIER, H.W.,
ROSS, C.A., and VAN WAGONER, J.C., eds., Sea level changes: An
integrated approach: Society of Economic Paleontologists and
Mineralogists, Special Publication, no. 42, p. 302–327.
BAYER, U., 1970, Anomalien bei Ammoniten des Aaleniums und Ba-
BRETT
jociums und ihre Beziehung Zur Lebensweise: Neues Jahrbuch
für Geologie und Paläontologie, Abhandlungen, v. 140, p. 306–327.
BAYER, U., and MCGHEE, G.R., 1985, Evolution in marginal epiconti-
nental basins: The role of phylogenetic and ecological factors (am-
monite replacements in the German Lower to Middle Jurassic): in
BAYER, U., and SEILACHER, A., eds., Sedimentary and Evolution-
ary Cycles, Lecture Notes in Earth Sciences, v. 1: Springer-Verlag,
Berlin, p. 164–220.
BAYER, U., and SEILACHER, A., EDS., 1985, Sedimentary and Evolu-
tionary Cycles, Lecture Notes in Earth Sciences, v. 1: Springer-
Verlag, Berlin, 465 p.
BAYER, U., ALTHEIMER, E. and DEUTCHLE, W., 1985, Environmental
evolution in shallow epicontinental seas: Sedimentary cycle and
bed formation: in BAYER, U., and SEILACHER, A., eds., Sedimenta-
ry and Evolutionary Cycles, Lecture Notes in Earth Sciences v. 1:
Springer-Verlag, Berlin, p. 347–381.
BAYER, U., JOHNSON, A.L.A., and BRENNAN, J., 1985, Ecological pat-
terns in Middle Jurassic Gryphaea: The relationship between
form and environment: in BAYER, U., and SEILACHER, A., eds.,
Sedimentary and Evolutionary Cycles, Lecture Notes in Earth
Sciences v. 1: Springer-Verlag, Berlin, p. 436–463.
BEADLE, S., and JOHNSON, M.E., 1986, Palaeoecology of Silurian cy-
clocrinitid algae: Palaeontology, v. 29, p. 585–601.
BENEDICT, G.L., and WALKER, K.G., 1978, Paleobathymetric analysis
in Paleozoic sequences and its geodynamic significance: American
Journal of Science, v. 278, p. 579–607.
BENNETT, K.D., 1990, Milankovitch cycles and their effects on species
in ecological and evolutionary time: Paleobiology, v. 16, p. 11–21.
BENNINGTON, J.B., 1994, Are similar fossil assemblages ever the
same? The statistical analysis of paleocommunity recurrence:
Geological Society of America, Abstracts with Programs, v. 26 (7),
p. A456.
BOTTJER, D.J., and DROSER, M.L., 1994, The history of Phanerozoic
bioturbation: in DONOVAN , S.K., ed., The Paleobiology of Trace
Fossils: John Wiley and Sons, Chichester, p. 155–178.
BOUCOT, A.J., 1975, Evolution and Extinction Rate Controls: Elsevier
Press, Amsterdam, 427 p.
BOUCOT, A.J., 1982, Ecostratigraphic framework for the Lower De-
vonian of the North American Appohimchi Subprovince: Neues
Jahrbuch für Geologie und Paläontologie, Abhandlungen, v. 163,
p. 81–121.
BOUCOT, A.J., 1986, Ecostratigraphic criteria for evaluating the mag-
nitude, character and duration of bioevents: in WALLISER, O.H.,
ed., Global Bio-events: A Critical Approach, Lecture Notes in
Earth Sciences, v. 8: Springer-Verlag, Berlin, p. 25–45.
BOUCOT, A.J., 1990a, Phanerozoic extinctions: how similar are they to
each other?: in KAUFFMAN, E.G., and WALLISER, O.H., eds., Ex-
tinction Events in Earth History, Lecture Notes in Earth Sciences,
v. 30: Springer-Verlag, Berlin, p. 5–30.
BOUCOT, A.J., 1990b, Silurian and pre-Upper Devonian bio-events: in
KAUFFMAN, E.G., and WALLISER, O.H., eds., Extinction Events in
Earth History, Lecture Notes in Earth Sciences, v. 30: Springer-
Verlag, Berlin, p. 125–132.
BOUCOT, A.J., 1990c, Evolutionary Paleobiology of Behavior and Co-
evolution: Elsevier Press, Amsterdam, 325 p.
BOUCOT, A.J., and LAWSON, J.D., EDS., 1998, Paleoecommunities—A
Case Study from the Silurian and Lower Devonian: World and Re-
gional Geology Series No.10, Cambridge University Press, New
York, 912 p.
BRENCHLEY, P.J., 1989, The Late Ordovician Extinction: in DONO-
VAN, S.K., ed., Mass Extinctions: Columbia University Press, New
York, p. 104–132.
BRETSKY, P.W., and BRETSKY, S.S., 1975, Succession and repetition of
Late Ordovician fossil assemblages from the Nicolet River Valley,
Quebec: Paleobiology, v. 1, p. 225–237.
BRETT, C.E., 1988, Paleoecology of hard substrate communities: An
overview: PALAIOS, v. 3, p. 374–378.
BRETT, C.E., 1995, Sequence stratigraphy, biostratigraphy, and ta-
phonomy in shallow marine environments: PALAIOS, v. 10, p.
597–616.
BRETT, C.E., and BAIRD, G.C., 1995, Coordinated stasis and evolution-
ary ecology of Silurian to Middle Devonian faunas in the Appala-
chian basin: in ERWIN, D., and ANSTEY, R., eds., New Approaches
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
to Speciation in the Fossil Record: Columbia University Press,
New York, p. 285–315.
BRETT, C.E., and BAIRD, G.C., 1996, Middle Devonian Sedimentary
cycles and Sequences in the northern Appalachian basin: in WITZ-
KE, B., and DAY, J., eds., Paleozoic Sequence Stratigraphy: Views
from the North American Craton, Geological Society of America,
Special Paper 306, p. 213–241.
BRETT, C.E., and BAIRD, G.C., 1997, Epiboles, outages and ecological
evolutionary bioevents’’: Taphonomic, ecological and biogeograph-
ical factors: in BRETT, C.E., and BAIRD, G.C., eds., Paleontological
Events: Stratigraphic, Ecological and Evolutionary Implications:
Columbia University Press, New York, p. 249–284.
BRETT, C.E., and BROOKFIELD, M.E., 1984, Morphology, faunas, and
genesis of Ordovician hardgrounds from southern Ontario, Cana-
da: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 46, p.
233–290.
BRETT, C.E., MILLER, K.B., and BAIRD, G.C., 1990, A temporal hier-
archy of paleoecological process in a Middle Devonian epeiric sea:
in MILLER, W., III, ed., Paleocommunity Temporal Dynamics: The
Long-Term Development of Multispecies Assemblages: Paleonto-
logical Society Special Publication, v. 5, p. 178–209.
BRETT, C.E., BOUCOT, A.J., and JONES, B., 1993, Absolute depths of
Silurian benthic assemblages: Lethaia, v. 26, p. 25–40.
BROMLEY, R.J.,1970, Borings as trace fossils and Entobia cretacea
Portlock, as an example: in CRIMES, T.P., and HARPER, J.C., eds.,
Trace Fossils: Seel House Press, Liverpool, p. 49–90.
CISNE, J.L., and RABE, B.D., 1978, Coenocorrelation: Gradient analy-
sis of fossil communities and its applications to stratigraphy: Le-
thaia, v. 11, p. 341–364.
CISNE, J.L., GILDNER, R., and RABE, B.D., 1984, Epeiric sedimenta-
tion and sea level: Synthetic ecostratigraphy: Lethaia, v. 17, p.
267–288.
CONNELL, J.H., and SLAYTOR, R.O., 1977, Mechanisms of succession
in natural communities and their role in community stability and
organization: American Naturalist, v. 111, p. 1119–1144.
DIMICHELE, W.A., 1994, Ecological patterns in time and space: Paleo-
biology, v. 20, p. 89–92.
DOCKERY, D.T., II, 1986, Punctuated succession of Paleogene mol-
luscs in the northern Gulf Coastal Plain: PALAIOS, v. 1, p. 582–
589.
DODD, J.R., and STANTON, R.J., JR., 1990, Paleoecology—Concepts
and Applications: John Wiley, New York, 502 p.
DONOVAN, S.K., ED., 1989, Mass Extinctions: Columbia University
Press, New York, 266 p.
DROSER, M.L., and BOTTJER, D.J., 1993, Trends and patterns of Phan-
erozoic ichnofabrics: Annual Review of Earth and Planetary Sci-
ences, v. 21, p. 204–225.
DUTRO, J.T., 1981, Devonian brachiopod biostratigraphy: in OLIVER,
W.A., JR. and KLAPPER, G., eds., Devonian Biostratigraphy of New
York: International Union of Geological Sciences, Washington,
D.C., p. 67–82.
EICHER, D.L., and DINER, R., 1985, Foraminifera as indicators of wa-
ter mass in the Cretaceous Greenhorn sea, Western Interior: in
PRATT, L.M., KAUFFMANN, E.G., and ZELT, F.G., eds., Fine-
grained deposits and biofacies of the Cretaceous Western Interior
Seaway: Evidence of cyclic sedimentary processes: Society of Eco-
nomic Paleontologists and Mineralogists, Field trip Guidebook,
no. 4, p. 60–71.
EINSELE, G., 1985, Response of sediments to sea-level change in dif-
fering subsidence storm-dominated marginal and epeiric basins:
in BAYER, U. and SEILACHER, A., eds., Sedimentary and Evolution-
ary Cycles, Lecture Notes in Earth Sciences, v. 1: Springer-Verlag,
Berlin, p. 68–97.
EINSELE, G., and SEILACHER, A., eds., 1982, Cyclic and Event Strati-
fication: Springer-Verlag, Berlin, 536 p.
EINSELE, G., RICKEN, W., and SEILACHER, A., EDS., 1991, Cycles and
Events in Stratigraphy: Springer-Verlag, Berlin, 955 p.
ELDER, W.P., 1987, Paleoecology of the Cenomanian-Turonian (Cre-
taceous) stage boundary extinctions at Black Mesa, Arizona: PA-
LAIOS, v. 2, p. 24–40.
ELDER, W.P., GUSTASON, E.R., and SAGEMAN, B.B., 1994, Correlation
of basinal carbonate cycles to nearshore parasequences in the Late
259
Cretaceous Greenhorn Seaway, Western Interior, U.S.A.: Geolog-
ical Society of America Bulletin, v. 106, p. 892–902.
ELDREDGE, N., 1990, Hierarchy and Macroevolution: in BRIGGS,
D.E.G., and CROWTHER, P.R., eds., Palaeobiology—A Synthesis:
Blackwell, Oxford, p. 124–129.
EMERY, D., 1996, Carbonate systems: in EMERY, D., and MEYERS,
K.J., eds., Sequence Stratigraphy: Blackwell, Oxford, p. 211–237.
EMERY, D. and MEYERS, K.J., EDS., 1996, Sequence Stratigraphy:
Blackwell, Oxford, 297 p.
ERWIN, D.J., and ANTSEY, R., 1995, New Approaches to Speciation in
the Fossil Record: Columbia University Press, New York, 342 p.
FISCHER, A.G., 1986, Climatic rhythms recorded in strata: Annual
Reviews of Earth and Planetary Sciences, v. 14, p. 351–376.
FREY, R.W., PEMBERTON, S.G., and SAUNDERS, T.D.A., 1990, Ichnofa-
cies and bathymetry: A passive relationship: Journal of Paleontol-
ogy, v. 64, p. 155–158.
FÜRSICH, F.T., 1978, The influence of faunal condensation and mixing
on the preservation of fossil benthic communities: Lethaia, v. 11,
p. 243–250.
FÜRSICH, F.T., 1979, Genesis, environments and ecology of Jurassic
hardgrounds: Neues Jahrbuch für Geologie und Paläontologie,
Abhandlungen, v. 158, p. 1–63.
FÜRSICH, F.T., 1994, Palaeoecology and evolution of Mesozoic salini-
ty-controlled benthic assemblages: Lethaia, v. 26, p. 327–346.
GASKEL, B.A., 1991, Extinction patterns in Paleogene benthic fora-
miniferal faunas relationship to climate and sea level: PALAIOS,
v. 6, p. 2–16.
GOLDRING, R., 1971, Shallow water sedimentation: Geological Society
of London, Memoir, v. 5, p. 1–88.
GOLDRING, R., 1995, Organisms and substrate response and effect: in
BOSENCE, D.W.J., and ALLISON, P.A., eds., Marine Palaeoenviron-
mental Analysis from Fossils: Geological Society of London, Spe-
cial Publication 83, p. 151–180.
GÓMEZ, J.J., and FERNANDEZ-LÓPEZ. S., 1994, Condensation process-
es in shallow platforms: Sedimentary Geology, v. 92, p. 147–159.
HAGDORN, H., 1982, The ‘‘Bank der Kleinen Terebrateln’’ (upper Mus-
chelkalk, Triassic) near Schwabisch Hall-Germany: Tempestite
condensation horizon: in EINSELE, G., and SEILACHER, A., eds., Cy-
clic and Event Stratification: Springer-Verlag, Berlin, p. 263–287.
HAGDORN, H., 1985, Immigrations of crinoids into the German Mus-
chelkalk basin: in BAYER, U., and SEILACHER, A., eds., Sedimen-
tary and Evolutionary Cycles: Lecture Notes in Earth Sciences, v.
1: Springer-Verlag, Berlin, p. 237–254.
HALLAM, A., 1978, How rare is phyletic gradualism and what is its
evolutionary significance? Evidence from Jurassic bivalves: Paleo-
biology, v. 4, p. 16–25.
HALLAM, A., 1981, A revised sea-level curve for the Early Jurassic:
Journal of the Geological Society of London, v. 138, p. 735–743.
HALLAM, A., 1987, Radiations and extinctions in relation to environ-
mental change in the marine Lower Jurassic of northwest Europe:
Paleobiology, v. 13, p. 152–168.
HALLAM, A., 1989, The case for sea-level change as a dominant factor
in mass extinction of marine invertebrates: Philosophical Trans-
actions of the Royal Society of London, v. 13, p. 152–168.
HALLAM, A., 1992, Phanerozoic Sea-Level Changes: Columbia Uni-
versity Press, New York, 266 p.
HALLAM, A., and BRADSHAW, M.J., 1979, Bituminous shales and oolit-
ic ironstones as indicators of transgression and regression: Jour-
nal of the Geological Society of London, v. 136, p. 157–164.
HANSEN, T., 1987, Extinction of late Eocene to Oligocene molluscs:
Relationship to shelf area, temperature changes and impact
events: PALAIOS, v. 2, p. 69–75.
HOLLAND, S.M., 1993, Sequence stratigraphy of a carbonate-clastic
ramp: The Cincinnatian Series (Upper Ordovician in its type
area): Geological Society of America Bulletin, v. 105, p. 306–322.
HOLLAND, S.M., 1995, The stratigraphic distribution of fossils: Paleo-
biology, v. 21, p. 92–109.
HOLLAND, S.M., 1997, Using time-environment analysis to recognize
faunal events in the Upper Ordovician of the Cincinnati Arch: in
BRETT, C.E., and BAIRD, G.C., eds., Paleontological Events: Strati-
graphic, Ecological and Evolutionary Implications: Columbia Uni-
versity Press, New York, p. 309–334.
HOLTERHOFF, P.F., 1992, Ecophenotypic variation and phylogeny
260
within the Erisocrinacea (Crinoidea): Linkage of morphology, ecol-
ogy, and sea-level in the Late Paleozoic: in LIDGARD S., and
CRANE, P.R., eds., Fifth North American Paleontological Conven-
tion, Abstracts and Program: The Paleontological Society, Knox-
ville, p. 131.
HOUSE, M.R., 1985, Correlation of mid-Paleozoic ammonoid evolu-
tionary events with global sedimentary perturbations: Nature, v.
313, p. 17–22.
JABLONSKI, D., 1980, Apparent versus real biotic effects of transgres-
sions and regressions: Paleobiology, v. 6, p. 397–407.
JABLONSKI, D., 1985, Marine regressions and mass extinctions: A test
using modern biota: in VALENTINE, J.W., ed., Phanerozoic Diver-
sity Patterns—Profiles in Macroevolution: Princeton University
Press, Princeton, p. 335–354.
JABLONSKI, D., 1986, Causes and consequences of mass extinctions:
Comparative approach: in ELLIOT, D.K., ed., Dynamics of Extinc-
tion: Wiley Press, New York, p. 183–229.
JOHNSON, R.G., 1972, Conceptual models of benthic marine commu-
nities: in SCHOPF, T.J.M., ed., Models in Paleobiology: Freeman
Cooper and Co., San Francisco, p. 148–159.
JOHNSON, J.G., 1974, Extinction of perched faunas: Geology, v. 2, p.
471–482.
JOHNSON, M.E., 1987, Extent and bathymetry of North America plat-
form seas in the Early Silurian: Paleoceanography, v. 2, p. 185–
211.
JOHNSON, R.G., KLAPPER, G., and SANDBERG, C.A., 1985, Devonian
eustatic fluctuations in Euramerica: Geological Society of America
Bulletin, v. 69, p. 567–587.
JONES, B., and PEMBERTON, S.G., 1989, Sedimentology and ichnology
of a Pleistocene unconformity bounded, shallowing-upward car-
bonate sequence: The Ironshore Formation, Salt Creek, Grand
Cayman: PALAIOS, v. 4, p. 343–355.
KAUFFMAN, E.G., 1977, Upper Cretaceous cyclothems, biotas, and en-
vironments, Rock Canyon anticline, Pueblo Colorado: The Moun-
tain Geologist, v. 14, p. 129–152.
KAUFFMAN, E.G., 1981, Ecological reappraisal of the German Posi-
donienschiefer (Toarcian) and the stagnant basin model: in GRAY,
J., BOUCOT, A.J., and BERRY, W.B.N., eds., Communities of the
Past: Hutchinson Ross, Stroudsburg, p. 311– 381.
KAUFFMAN, E.G., 1984, The fabric of Cretaceous marine extinctions:
in BERGGEN, W.A., and VAN COUVERING, J.A., eds., Catastrophes
and Earth History: The New Uniformitarianism, Princeton Uni-
versity Press, Princeton, p. 151–246
KAUFFMAN, E.G., 1986, High-resolution event stratigraphy: Regional
and global Cretaceous bio-events: in WALLISER, O., ed., Global
Bioevents, Lecture Notes in Earth Sciences, v. 8: Springer-Verlag,
Berlin, p. 279–335.
KAUFFMAN, E.G., and SAGEMAN, B.B., 1992, Biological patterns in se-
quence stratigraphy; Cretaceous of the Western Interior Basin,
North America: in LIDGARD, S., and CRANE, P.R., eds., Fifth North
America Paleontological Convention, Abstracts and Program: The
Paleontological Society, Knoxville, p. 158.
KAUFFMAN, E.G., and WALLISER, O.H., EDS., 1990, Extinction Events
in Earth History, Lecture Notes in Earth Sciences, v. 30: Springer-
Verlag, Berlin, 432 p.
KELLY, P.H., 1983, The role of within-species differentiation in mac-
roevolution of Chesapeake Group bivalves: Paleobiology, v. 9, p.
261–268.
KIDWELL, S.M., 1991, Taphonomic feedback (live/dead interactions)
in the genesis of bioclastic beds: Keys to reconstructing sedimen-
tary dynamics: in EINSELE, G., RICKEN, W. and SEILACHER, A.,
eds., Cycles and Events in Stratigraphy: Springer-Verlag, Berlin,
p. 268–282.
KIDWELL, S.M., and AIGNER, T., 1985, Sedimentary dynamics of com-
plex shell beds: Implications for ecologic and evolutionary pat-
terns: in BAYER, U., and SEILACHER, A., eds., Sedimentary and
Evolutionary Cycles, Lecture Notes in Earth Sciences, v. 1:
Springer-Verlag, Berlin, p. 382–395.
KIDWELL, S.M., and BEHRENSMEYER, A.K., EDS., 1993, Taphonomic
Approaches to Time Resolution in Fossil Assemblages: Paleonto-
logical Society Short Course No. 6, Knoxville, 302 p.
KIDWELL, S.M., and JABLONSKI, D., 1983, Taphonomic feedback: Eco-
logical consequences of shell accumulation: in TEVESZ, M.J.S., and
BRETT
MCCALL, P.L., eds., Biotic Interactions in Recent and Fossil Ben-
thic Communities: Plenum Press, New York, p. 195–248.
LANDING, E., and BRETT, C.E., 1987, Trace fossils and regional signif-
icance of a Middle Devonian (Givetian) disconformity in south-
western Ontario: Journal of Paleontology, v. 61, p. 205–230.
LECKIE, R.M., SCOTT, R.W., BRALOWER, T.J., and SLITER, W.V., 1992,
Relationship between sequence boundaries and the evolutionary
history of planktonic foraminifera, calcareous nannofossils, and
reef communities in the Mid-Cretaceous (Barremian-Cenoma-
nian): in LIDGARD, S., and CRANE, P.R., eds., Fifth North Ameri-
can Paleontological Convention, Abstracts and Program: The Pa-
leontological Society, Knoxville, p. 179.
LEES, D.R., and EDWARDS, D., 1993, Evolutionary Patterns and Pro-
cesses: Linnean Society Symposium, v. 14, Academic Press, Lon-
don, p. 19–31.
LEVINTON, J.S., 1982, Marine Ecology: Prentice-Hall, Inc., Engle-
wood, 526 p.
LOUTIT, T.S., HARDENBOL, J., VAIL, P.R., and BAUM, P., 1988, Con-
densed sections: The key to age dating and correlation of continen-
tal margin sequences: in WILGUS, C.K., HASTINGS, B.S., ST. C.
KENDALL, C.G., POSAMENTIER, H.W., ROSS, C.A., and VAN WAG-
ONER, J.C., eds., Sea-level Changes: An Integrated Approach: So-
ciety of Economic Paleontologists and Mineralogists, Special Pub-
lication, no. 42, p. 183–213.
MACARTHUR, R.H., and WILSON, E.O., 1967, The Theory of Island
Biogeography: Princeton University Press, Princeton, 203 p.
MAPLES, C.G., and WEST, R.R., EDS., 1992, Trace fossils: Paleontolog-
ical Society, Short Course No. 5, Knoxville, 238 p.
MCGHEE, G.R., JR., 1991, Extinction and diversification in the Devo-
nian brachiopoda of New York: No correlation with sea level?: His-
torical Biology, v. 5, p. 215–227.
MCGHEE, G.R., JR., 1992, Evolutionary biology of the Devonian
brachiopoda of New York State: No correlation with rate of change
of sea-level?: Lethaia, v. 25, p. 165–172.
MCKERROW, S.J., 1979, The Ecology of Fossils: The MIT Press, Cam-
bridge, 384 p.
MEYERS, K.J., and MILTON, N.J., 1996, Principles of sequence stratig-
raphy: in EMERY, D., and MEYERS, K.J., eds., Sequence Stratigra-
phy: Blackwell Science Ltd., Oxford, p. 11–44.
MILLER, W., III, 1986, Paleoecology of benthic community replace-
ment: Lethaia, v. 19, p. 225–231.
MILLER, W., III, ED., 1990, Paleocommunity Temporal Dynamics: Pa-
leontological Society, Special Publication No. 5, Knoxville, 421 p.
MILLER, W., III, and DUBAR, J.R., 1988, Community replacement of a
Pleistocene Crepidula biostrome: Lethaia, v. 21, p. 67–78.
MILLER, K.B., BRETT, C.E., and PARSONS, K.M., 1988, The paleoeco-
logic significance of storm-generated disturbance within a Middle
Devonian muddy epeiric sea: PALAIOS, v. 3, p. 35–52.
MITCHUM, R.M., JR., VAIL, P.R., and THOMPSON, S., III, 1977, The de-
positional sequence as a basic unit for stratigraphic analysis: in
PAYTON, C.E., ed., Seismic stratigraphy—Applications to hydro-
carbon exploration: American Association of Petroleum Geologists
Memoir No. 26, p. 53–62.
MOORE, R.C., 1954, Evolution of the late Paleozoic invertebrates in
the response to major oscillations of shallow seas: Bulletin of the
Museum of Comparative Zoology, Harvard, v. 122, p. 259–286.
MORTIMORE, R.N., and POMEROL, R., 1991, Stratigraphy and eustatic
implications of trace fossil events in the Upper Cretaceous chalk of
northwestern Europe: PALAIOS, v. 6, p. 216–232.
NEWELL, N.D., 1967, Revolutions in the history of life: Geological So-
ciety of America, Special Paper No. 89, p. 63–91.
ODUM, E.P., 1971, Fundamentals of Ecology: W.B. Saunders Co.,
Philadelphia, 574 p.
PALMER, A.R., 1984, The biomere problem: Evolution of an idea: Jour-
nal of Paleontology, v. 58, p. 599–611.
PALMER, T.W., and PALMER, C., 1977, Faunal distribution and colo-
nization strategy in a Middle Ordovician hardground community:
Lethaia, v. 10, p. 179–266.
PATZKOWSKI, M.E., and HOLLAND, S.M., 1996. Faunal extinctions, mi-
grations, and stratigraphic sequences: Middle and Upper Ordovi-
cian of the eastern United States: in WITZKE, B.D., BUNKER, W.,
and DAY, J., eds., Paleozoic sequences: View from the craton: Geo-
logical Society of America, Special Paper No. 306, p. 131–142.
SEQUENCE STRATIGRAPHY AND PALEOECOLOGY
PEMBERTON, S.G., and MACEACHERN, J.A., 1997, The ichnological sig-
nature of storm deposits: The use of trace fossils in event stratig-
raphy: in BRETT, C.E., and BAIRD, G.C., eds., Paleontologic
Events: Stratigraphic, Ecologic, and Evolutionary Implications:
Columbia University Press, New York, p. 73–109.
POMPECKJ, J.F., 1914, Die Bedeutung des Schwabischen Jura für die
Erdgeschichte: Schweizerbart, Stuttgart, 64 p.
POSAMENTIER, H.W., and VAIL, P.R., 1988, Eustatic controls on clastic
deposition II—Sequence and systems tract models: in WILGUS,
C.K., HASTINGS, B.S., ST. C. KENDALL, C.G., POSAMENTIER, H.W.,
ROSS, C.A., and VAN WAGONER, J.C., eds., Sea-level changes: An
integrated approach, Society of Economic Paleontologists and
Mineralogists, Special Publication No. 42, p. 125–154.
POSAMENTIER, H.W., JERVEY, M.T., and VAIL, P.R., 1988, Eustatic
controls on clastic deposition I—Conceptual framework: in WIL-
GUS, C.K., HASTINGS, B.S., ST. C. KENDALL, C.G., POSAMENTIER,
H.W., ROSS, C.A., and VAN WAGONER, J.C., eds., Sea-level
changes: An integrated approach, Society of Economic Paleontol-
ogists and Mineralogists, Special Publication No. 42, p. 109–124.
RAILSBACK, L.B., ACKERLY, S.A., ANDERSON, T.F., and CISNE, J.L.,
1990, Paleontological and isotope evidence for warm saline deep
waters in Ordovician oceans: Nature, v. 343, p. 156–159.
RAMSBOTTOM, W.H.C., 1981, Eustatic control on Carboniferous am-
monoid biostratigraphy: in HOUSE, M.R., and SENIOR, J.R., eds.,
The Ammonoidea: Systematics Association Special Publication, v.
18, Academic Press, London, p. 369–388.
RHOADS, D.C., SPEDEN, J.G., and WAAGE, K.M., 1972, Trophic group
analysis of Upper Cretaceous (Maastrichtian) bivalve assem-
blages from South Dakota: American Association of Petroleum
Geologists Bulletin, v. 56 , p. 1100–1113.
ROLLINS, H.B., CAROTHERS, M., and DONAHUE, J., 1979, Transgres-
sion, regression and fossil community succession: Lethaia, v. 12, p.
89–104.
ROSENKRANZ, D., 1971, Zur Sedimentologie and Okölogie von Echi-
nodermen- Lagerstätten: Neues Jahrbach für Geologie and Pa-
läontologie, Abhandlungen, v. 138, p. 221–258.
ROSS, C.R., and ROSS, J.P., 1988, Late Paleozoic transgressive-regres-
sive deposition: in WILGUS, C.K., HASTINGS, B.S., ST. C. KENDALL,
C.G., POSAMENTIER, H.W., ROSS, C.A., and VAN WAGONER, J.C.,
eds., Sea-level changes: An integrated approach, Society of Eco-
nomic Paleontologists and Mineralogists, Special Publication No.
42, p. 227–247.
ROSS, C.R., and ROSS, J.P., 1994, Permian sequence stratigraphy and
fossil zonation: in BEAUCHAMP, B., EMBRY, A.E., and GLASS, D.,
eds., Pangea Global Environments and Resources: Canadian So-
ciety of Petroleum Geologists Memoir 17, p. 227–247.
ROSS, R.M., and ALLMON, W.D., EDS., 1990, Causes of Evolution: A
Paleontological Perspective: University of Chicago Press, Chicago,
479 p.
SADLER, P.J., 1981, Sediment accumulation rates and the complete-
ness of stratigraphic sections: Journal of Geology, v. 89, p. 569–
584.
SAGEMAN, B.B., 1992, Paleoecology—A cure for sequence syndrome?
PALAIOS, v. 7, p. 485–486.
SAGEMAN, B.B., WIGNALL, P.B., and KAUFFMAN, E.G., 1991, Biofacies
model for oxygen-deficient facies in epicontinental seas: Tool for
paleoenvironmental analysis: in EINSELE, G., RICKEN, W., and
SEILACHER, A., eds., Cycles and Events in Stratigraphy: Springer-
Verlag, Berlin, p. 542–564.
SARG, J.F., 1988, Carbonate sequence stratigraphy: in WILGUS, C.K.,
HASTINGS, B.S., ST. C. KENDALL, C.G., POSAMENTIER, H.W., ROSS,
C.A., and VAN WAGONER, J.C., eds., Sea-level changes: An inte-
grated approach, Society of Economic Paleontologists and Miner-
alogists Special Publication No. 42, p. 155–181.
SAVARESE, M.L., BRETT, C.E., and GRAY, L.M., 1986, Faunal and lith-
ological cyclicity in the Centerfield Member (Middle Devonian)
Hamilton Group of western New York: A reinterpretation of de-
positional history: in BRETT, C.E., ed., Dynamic Stratigraphy and
Depositional Environments of the Hamilton Group (Middle Devo-
nian) in New York State, Part I: New York State Museum Bulletin
457, p. 78–101.
SAVRDA, C.E., 1991, Ichnology in sequence stratigraphic studies: An
261
example from the Lower Paleocene of Alabama: PALAIOS, v. 6, p.
39–54.
SAVRDA, C.E., and BOTTJER, D.J., 1989, Anatomy and implications of
bioturbated beds in ‘‘black shale’’ sequences: Examples from the
Jurassic Posidonienschiefer (southern Germany): PALAIOS, v. 4,
p. 330–342.
SAVRDA, C.E., and BOTTJER, D.J., 1991, Oxygen-related biofacies in
marine strata: an overview: in TYSON, R.V., and PEARSON, T.H.
eds., Modern and Ancient Continental Shelf Anoxia: Geological
Society of London Special Publication No. 58, p. 291–310.
SAVRDA, C.E., and BOTTJER, D.J., 1994, Ichnofossils and ichnofabrics
in rhythmically bedded pelagic-hemipelagic carbonates: Recogni-
tion and evaluation of benthic redox and scour cycles: in DE BOER,
P.L., and SMITH, D.G., eds., Orbital Forcing and Cyclic Sequences:
International Association of Sedimentologists, Special Publication
No. 19, p. 195–210.
SCHINDEL, D.E., 1980, Microstratigraphic sampling and the limits of
paleontologic resolution: Paleobiology, v. 6, p. 408–426.
SCHINDEL, D.E., 1982, The gaps in the fossil record: Nature, v. 297, p.
282–284.
SCHINDLER, E., 1990, The Late Frasnian (Upper Devonian) Kellwas-
ser crisis: in KAUFFMAN, E.G., and WALLISER, O.H., eds., Extinc-
tion Events in Earth History, Lecture Notes in Earth Sciences, v.
30: Springer-Verlag, Berlin, p. 151–159.
SCHOPF, T.J.M., 1974, Permo-Triassic extinctions: Relation to sea
floor spreading: Journal of Geology, v. 82, p. 129–143.
SCOTT, R.W., 1978, Approaches to trophic analysis in paleocommuni-
ties: Lethaia, v. 11, p. 1–14.
SEILACHER, A., 1967, Bathymetry of trace fossils: Marine Geology, v.
5, p. 413–428.
SEILACHER, A., MATYJA, B.A., and WIERZBOWSKI, A., 1985, Oyster
beds: Morphological response to changing substrate conditions: in
BAYER, U., and SEILACHER, A., eds., Sedimentary and Evolution-
ary Cycles, Lecture Notes in Earth Sciences, v. 1: Springer-Verlag,
Berlin, p. 421–435.
SEILACHER, A., REIF, W.E., and WESTPHAL, F., 1985, Sedimentologic-
al, ecological and temporal patterns of fossil Lagerstätten: in
WHITTINGTON, H.B., and CONWAY MORRIS, S., eds., Extraordinary
Biotas: Their Ecological and Evolutionary Significance: Philo-
sophical Transactions, Royal Society of London, v. 311, p. 5–23.
SELLWOOD, B.W., 1970, The relation of trace fossils to small scale sed-
imentary cycles in the British Lias: in CRIMES, T.P., and HARPER,
J.C., eds., Trace Fossils: Seel House, Liverpool, p. 489–504.
SHELDON, P., 1993, Making sense of microevolutionary patterns: in
LEES, R.B., and EDWARDS, D., eds., Evolutionary Patterns and
Processes: Linnear Society Symposium, v. 14, p. 19–31.
SIGNOR, P.W., III, and LIPPS, J. H., 1982, Sampling bias, gradual ex-
tinction patterns, and catastrophes in the fossil record: in SILVER,
L.T., and SCHULTZ, P.H., eds., Geological Implications of Impacts
of Large Asteroids and Comets on the Earth: Geological Society of
America Special Paper 190, p. 291–296.
SIMBERLOFF, D., 1974, Permo-Triassic extinctions: Effects of area on
biotic distribution: Journal of Geology, v. 82, p. 267–274.
SLOSS, L.L., 1963, Sequences in the cratonic interior of North Ameri-
ca: Geological Society of America Bulletin, v. 74, p. 93–119.
SNEDEN, J.W., 1991, Origin and sequence stratigraphic significance
of large dwelling traces in the Escondido Formation (Cretaceous
Texas, USA): PALAIOS, v. 6, p. 541–552.
STANLEY, S.M., 1984, Marine mass extinction: A dominant role for
temperature: in NITECKI, M., ed., Extinctions: Chicago University
Press, Chicago, p. 69–117.
STANLEY, S.M., 1988, Paleozoic mass extinctions: Shared patterns
suggest global cooling as a common cause: American Journal of
Science, v. 288, p. 334–352.
TAYLOR, A.M., and GAWTHORPE, R.L., 1993, Application of sequence
stratigraphy and trace fossil analysis to reservoir description: Ex-
amples from the Jurassic of the North Sea: in PARKER, J.R., ed.,
Petroleum Geology of Northwest Europe: Proceedings of the 4th
Conference: Geological Society, London, p. 317–335.
VAIL, P.R., MITCHUM, R.M., JR., and THOMPSON, S., III., 1977a, Part
three: Relative changes of sea-level from coastal onlap: in PAYTON,
C.E., ed., Seismic stratigraphy-Applications to hydrocarbon explo-
262
ration: American Association of Petroleum Geologists Memoir No.
26, p. 63–82.
VAIL, P.R., MITCHUM, R.M., JR., and THOMPSON, S., III, 1977b, Part
four: Global cycles of relative changes sea-level: in PAYTON, C.E.,
ed., Seismic stratigraphy—Applications to hydrocarbon explora-
tion: American Association of Petroleum Geologists Memoir No.
26, p. 83–97.
VAIL, P.R., AUDEMARD, F., BOWMAN, S.A., EISNER, P.N., and PERÊZ-
CRUZ, C., 1991, The stratigraphic signatures of tectonics, eustacy
and sedimentology-an overview: in EINSELE, G., RICKEN, W., and
SEILACHER, A., eds., Cycles and Events in Stratigraphy: Springer-
Verlag, Berlin, p. 617–659.
VAN WAGONER, J.C., POSAMENTIER, H.W., MITCHUM, R.M., JR., VAIL,
P.R., SARG, J.F., LOUTIT, T.S., and HARDENBOL, J., 1988, An over-
view of the fundamentals of sequence stratigraphy and key defi-
nitions: in WILGUS, C.K., HASTINGS, B.S., ST. C. KENDALL, C.G.,
POSAMENTIER, H.W., ROSS, C.A., and VAN WAGONER, J.C., eds.,
Sea-level changes: An integrated approach: Society of Economic
Paleontologists and Mineralogists Special Publication No. 42, p.
39–45.
VALENTINE, J.W., 1973, Evolutionary Paleoecology of the Marine Bio-
sphere: Prentice Hall, Inc., Englewood Cliffs, 511 p.
VALENTINE, J.W., 1985, Phanerozoic Diversity Patterns; Profiles in
Macroevolution: Princeton University Press, Princeton, 441 p.
VOSSLER, S., and PEMBERTON, S.G., 1988, Superabundant Chon-
drites: A response to storm buried organic matter?: Lethaia, v. 21,
p. 94.
WALKER, K.R., and ALBERSTADT, L.P., 1975, Ecological succession as
an aspect of structure in fossil communities: Paleobiology, v. 1, p.
238–257.
WALLISER, O.H., ED., 1986a, Global bio-events: Lecture Notes in
Earth Sciences, v. 8: Springer-Verlag, Berlin, 442 p.
WALLISER, O.H., 1986b, Towards a more critical approach to bio-
events: in WALLISER, O.H., ed., Global Bio-Events, A Critical Ap-
proach, Lecture Notes in Earth Sciences, v. 8, Springer-Verlag,
Berlin, Heidelberg, p. 1–4.
BRETT
WALLISER, O.H., 1990, How to define ‘‘global bioevents’’: in KAUFF-
MAN, E.G., and WALLISER, O.H., eds., Extinction Events in Earth
History, Lecture Notes in Earth Sciences, v. 30: Springer-Verlag,
Berlin, p. 1–4.
WARME, J.E., 1975, Borings as trace fossils, and the processes of ma-
rine bioerosion: in FREY, R.W., ed., The Study of Trace Fossils:
Springer-Verlag, New York, p. 181–227.
WETZEL, A., and AIGNER, T., 1986, Stratigraphic completeness:
Tiered trace fossils provide a measuring stick: Geology, v. 14, p.
234–237.
WIGNALL, P.B., 1994, Black Shales: Oxford Monographs on Geology
and Geophysics, v. 30, 127 p.
WIGNALL, P.B., and HALLAM, A., 1991, Biofacies, stratigraphic distri-
bution and depositional models of British onshore Jurassic black
shales: in TYSON, R.V., and PEARSON, T.H., eds., Modern and An-
cient Continental Shelf Anoxia: Geological Society of London Spe-
cial Publication No. 58, p. 291–310.
WILGUS, C.K., HASTINGS, B.S., ST. C. KENDALL, C.G., POSAMENTIER,
H.W., ROSS, C.A., and VAN WAGONER, J.C., EDS., 1988, Sea-level
changes: An integrated approach: Society of Economic Paleontol-
ogists and Mineralogists, Special Publication No. 42, 407 p.
WILLIAMS, H.S., 1903, Correlation of geological faunas: U.S. Geologi-
cal Survey Bulletin 210, 147 p.
WILLIAMS, H.S., 1913, Recurrent Tropidoleptus fauna of the Upper
Devonian in New York: U.S. Geological Survey Professional Paper
79, 103 p.
WILSON, M.A., and PALMER, T.J., 1992, Hardgrounds and hardground
faunas: Institute of Earth Studies Publications, University of
Wales, Aberystwyth, v. 9, 131 p.
ZIEGLER, A.M., 1965, Silurian marine communities and their environ-
mental significance: Nature, v. 207, p. 270–272.
ZIEGLER, A.M., COCKS, L.R.M., and BAMBACH, R.K., 1968, The com-
position and structure of lower Silurian marine communities: Le-
thaia, v. 1, p. 1–27.
ACCEPTED JANUARY 20,1998