444
Measuring Recurrence of Marine Biotic Gradients: A Case
Study from the Pennsylvanian-Permian Midcontinent
THOMAS D. OLSZEWSKI
Department of Geological Sciences, Indiana University, Bloomington, IN 47405
MARK E. PATZKOWSKY
Department of Geosciences, The Pennsylvania State University, University Park, PA 16802
PALAIOS, 2001, V. 16, p. 444–460
The aim of this study was to determine whether biotic as-
sociations of Pennsylvanian-Permian brachiopods and bi-
valves from the northern Midcontinent differ in their degree
of recurrence through time. The study interval includes 2.5
Myr that can be divided into 5 full and 2 partial composite
depositional sequences separated by subaerial unconformi-
ties. These stratigraphic packages represent replicate natu-
ral experiments in establishing the benthic marine ecosys-
tem of the basin. Based on cluster and ordination analyses,
two discrete biofacies can be recognized—one dominated by
brachiopods and the other by bivalves. Within each of these,
environmental gradients can be recognized. The brachio-
pod gradient is interpreted to reflect the degree of water-col-
umn oxygenation, whereas the bivalve gradient is interpret-
ed to reflect the transition from restricted to open-marine
conditions. Comparison of measured recurrence with ran-
domized data indicates that the ecological segregation of
the two biofacies is maintained to a significant degree
through the succession of depositional sequences in the
study interval. In contrast, the gradients within each bio-
facies, although recognizable, are not maintained rigidly
from sequence to sequence. There is also no significant dif-
ference in gradient recurrence between the two biofacies.
These results imply that there is no need to call upon strong
interspecific interactions to maintain the structure of these
paleocommunities through time.
INTRODUCTION
A question of long-standing interest to ecologists and
paleoecologists concerns the degree of integration in bio-
logical communities (Jackson, 1994). This issue frequently
is cast as a debate between advocates of strong interde-
pendence of species (Clements, 1916) and strong indepen-
dence of species (Gleason, 1926). Ecologists have tested
these alternative hypotheses by examining how species
compositions change along environmental gradients
(Whittaker, 1967). In a Clementsian world, species com-
position and abundance are expected to be very consistent
over a range of environmental conditions because the spe-
cies that make up a community form an integrated entity
(Ricklefs, 1990). In a Gleasonian world, species composi-
tion and abundance should vary from site to site as condi-
tions change because every species reacts to environmen-
tal change independently (Ricklefs, 1990). Ecologists gen-
erally study species associations in the single time slice
Copyright Q 2001, SEPM (Society for Sedimentary Geology)
RESEARCH REPORTS
represented by the modern world. In contrast, although
there are limitations imposed by taphonomy and strati-
graphic resolution (Kidwell and Flessa, 1995; Martin,
1999 and references therein), paleoecologists can examine
the fossil record of biological communities through time.
The aim of this paper is to identify and characterize the
consistency of species associations through time using
Pennsylvanian-Permian bivalves and brachiopods of the
North American Midcontinent as a case study.
Recent paleoecological studies of marine invertebrates
(e.g., Brett and Baird, 1995; Westrop, 1996; Tang and Bo-
ttjer, 1996; Patzkowsky and Holland, 1997; Olszewski and
Patzkowsky, in press) have focused on compositional
change at the scale of entire basins, combining taxonomic
lists from a variety of environments. In this study, the fo-
cus is on changes within biofacies rather than the entire
basin. Cluster analysis has been used to define paleocom-
munity types, gradient analysis to examine inter-specific
associations within paleocommunity types, and recur-
rence analysis to test for changes in associations through
time. In order to put the results in a broader context, they
are compared with those of other recent studies covering
different groups, environments, and ages (Bennington
and Bambach, 1996; Holterhoff, 1996; Pandolfi, 1996).
STRATIGRAPHIC FRAMEWORK
This study focuses on a 2.5-Myr interval of late Pennsyl-
vanian to early Permian rocks (Fig. 1) of the northern Mid-
continent (Fig. 2). At this time of global ‘‘icehouse’’ climatic
conditions (Fischer, 1984), an ice sheet was present at the
south pole (Ziegler et al., 1997), leading to cyclical eustatic
and climatic changes analogous in frequency, amplitude,
and effect to those of the Quaternary (Veevers and Powell,
1987). These are reflected in the cyclothemic stratigraphy
of Pennsylvanian-Permian rocks in the Midcontinent
(Heckel, 1984, 1986, 1995).
Cyclicity occurs at two scales in the study interval. At
the finest resolution, meter-scale cycles occur in both
open-marine, platform settings and nearshore, coastal set-
tings (Miller and West, 1993; Miller et al., 1996; Olszew-
ski, 2000). These cycles are very thin (1 to 5 m), unconfor-
mity-bounded depositional sequences and, therefore, rep-
resent temporally significant packets of rock (Posamentier
et al., 1988; Van Wagoner et al., 1988) averaging 50,000
years or less in duration.
At a larger scale (that of the classical cyclothems), com-
posite depositional sequences, composed of 8 to 13 meter-
scale cycles, can be recognized (Miller and West, 1993,
0883-1351/01/0016-0444/$3.00
PALEOCOMMUNITY RECURRENCE
FIGURE 1—Stratigraphic study interval. Stratigraphic column modified
from Zeller (1968) and Baars et al. (1994). Roman numerals denote
composite depositional sequences. I through V are complete sequenc-
es. 0 and VI are partial sequences.
1998; Olszewski, 2000). Open-marine carbonates, marine
mudrocks, and condensed sections dominate their trans-
gressive and early highstand phases. Late highstand de-
posits include peritidal carbonates, deltaic sandstones,
and pedogenic mudrocks. Correlation of meter-scale cycles
reveals angular unconformities at the boundaries of com-
posite sequences (Olszewski, 2000), indicating that these
too are temporally significant packages of rock.
The 51 meter-scale cycles identified in the study inter-
val (Olszewski, 2000) provide a means of examining
trends in paleoecological communities at relatively high
temporal resolution (;5·104 year). The five complete and
two partial composite sequences (Fig. 1) are separated by
extensive subaerial unconformities. This means that their
deposition required re-establishment of marine environ-
445
FIGURE 2—Study area. Shaded area indicates exposure of upper
Pennsylvanian through lower Permian rocks used in this study. North-
south geographic zones are also indicated (see Table 2).
ments on the platform. Each composite sequence, there-
fore, can be treated as a replicate natural experiment in
setting up a benthic marine ecosystem that provides a
means of measuring the recurrence of ecological associa-
tions.
DATA
Data consist of fossil bivalve and articulate brachiopod
collections (plus the calcareous inarticulate genus Petro-
crania) from two sources: 341 collections from Mudge and
Yochelson’s (1962) monograph on the stratigraphy and pa-
leontology of the Pennsylvanian-Permian Midcontinent
and 132 additional collections made in 1997 through 1999
(‘‘new collections’’). Each collection represents the assem-
blage of fossils found in a bed at a site; they were not com-
bined to create composite lists by geologic unit or location.
The entire data set is included in Olszewski (2000) and
can be accessed at the Pennsylvania State University’s
Electronic Theses and Dissertations website (http://
etda.libraries.psu.edu/).
All fossils were identified to as fine a taxonomic level as
possible. Although many specimens were identifiable to
species, a large proportion could only be determined to ge-
nus. Therefore, to make use of as much data as possible,
all statistical analyses were conducted at the genus level.
This choice did not have a strong influence on patterns of
ecological association for two reasons. First, most genera
in the study interval are monospecific. Second, most poly-
specific genera in the data set are dominated by one spe-
cies. For example, in Mudge and Yochelson’s (1962) data,
Neospirifer is reported 116 times: 38 as N. sp. indet., 7 as
N. cf. N. kansasensis, and 71 as N. dunbari. If the propor-
tion of 7 to 71 is representative of the relative frequency of
these two species, then only 11 of the 116 Neospirifer oc-
currences are expected to be N. cf. N. kansasensis. These
figures, which are typical of other genera as well, indicate
that the number of associations gained by using genera
rather than species provides more information (in a statis-
tical sense) than including many rare species and reject-
ing specimens not identified to the species level.
Because the interest of this study was in associations of
446 OLSZEWSKI & PATZKOWSKY

TABLE 1—Descriptive statistics of data subsets. S is the richness of collections included in each data subset. ‘‘New collections’’ are those
made for this study and added to those of Mudge and Yochelson (1962). ‘‘All collections’’ denotes Mudge and Yochelson (1962) collections
plus ‘‘new collections.’’ ‘‘All genera’’ indicates inclusion of both brachiopod and bivalve genera (as opposed to just one group or the other). An
occurrence denotes the presence of a taxon in a collection regardless of its abundance. In a presence–absence data matrix, where 1 indicates
presence and 0 indicates absence, the total number of occurrences is simply the sum of the matrix (i.e., the number of 1’s).

Data subset Taxa Collections Occurrences Specimens

All collections All genera, S $ 2 40 474 2,593 18,737

All genera, S $ 4 35 317 2,191 16,122
Brachiopods, S $ 2 23 382 2,080 16,317
Bivalves, S $ 2 16 128 367 1,745
Bivalves, S $ 3 12 60 225 927
New collections All genera, S $ 2 34 132 865 7,687
All genera, by sequence, I 31 157 844 4,902
S$2 II 27 61 314 2,002
III 31 147 925 6,209
IV 23 29 167 1,727
V 26 82 319 1,764
Brachiopods, by sequence, I 20 119 634 3,866
S$2 II 16 41 199 1,452
III 20 140 851 6,043
IV 17 25 143 1,628
V 16 49 187 977
Bivalves, by sequence, I 11 49 149 702
S$2 II 11 27 91 497
III 9 18 42 116
IV 4 5 12 53
V 10 39 109 646

two or more taxa, collections including only one taxon
were not included in statistical analyses. The final data
set of 474 collections includes a total of 18,737 specimens
and 2,593 occurrences (Table 1).
The geographic and stratigraphic distribution of collec-
tions is summarized in Table 2 for seven north-south geo-
graphic zones (Fig. 2) of about 48 km each (five townships
or about 30 miles) and the five complete composite se-
quences. Four hundred and twenty-four collections are in-
cluded in the table, leaving 50 unassigned. Seven of these
came from Nebraska, thirty-eight came from the two par-
tial composite sequences (marked 0 and VI in Fig. 1), and
five came from stratigraphically floating exposures. These
collections were included in ordination and cluster analy-
ses despite uncertain external information because they
still shed light on taxonomic associations. Although con-
tingency table analysis indicates that collections are not
distributed randomly (Gadj(df 5 24) 5 54.45, P 5 0.00037; So-
kal and Rohlf, 1995), no stratigraphic or geographic trends
are evident in Table 2.
Most of the collections are small with median richness
equal to 5 and median number of specimens equal to 21
(Fig. 3). Although their small sizes suggest that many tax-
onomic lists may not be complete, the large number of col-
lections reduces statistical uncertainty.
Collections were taken from a wide variety of lithologies
and taphonomic states. Some collections came from hard
limestone, from which specimens had to be removed with
chisel and hammer or examined on slab surfaces, whereas
others came from soft mudrocks, which could be easily
bulk sampled or surface collected. Some fossils occurred as
original shell material while others were molds (some-
times in the same collection). In addition, collections most
certainly represent different amounts of time-averaging
(Kidwell and Bosence, 1991; Brett, 1995); some come from
condensed intervals within meter-scale cycles represent-
ing 100’s to 1000’s of years, whereas others come from fa-
cies that represent relatively rapid burial.
Although all these non-uniform factors increase vari-
ability, the collections are reliable as records of fossil taxa
that demonstrably co-occur in specific beds at specific lo-
cations. As such, they provide a great deal of information

TABLE 2—Distribution of collections used in this study by geographic zone and composite sequence. Values are ‘‘all collections’’ with S $ 2
(Table 1). Numbers in parentheses are ‘‘new collections’’, S $ 2. The distributions of Mudge and Yochelson’s (1962) collections and ‘‘new
collections’’ are significantly correlated (r 5 0.738 . r950.05 5 0.334; df 5 33).

Geographic zones
Composite sequences 1 2 3 4 5 6 7 Row totals

V 4 (0) 31 (14) 16 (6) 15 (3) 4 (0) 8 (0) 14 (0) 92 (23)

IV 2 (1) 9 (6) 3 (0) 3 (1) 7 (6) 1 (0) 5 (1) 30 (15)
III 10 (0) 73 (33) 25 (12) 3 (0) 8 (1) 16 (9) 12 (1) 147 (56)
II 3 (0) 21 (7) 16 (2) 7 (0) 3 (0) 6 (0) 5 (2) 61 (11)
I 12 (1) 31 (6) 18 (2) 5 (0) 14 (0) 10 (0) 4 (1) 94 (10)
Column totals 31 (2) 165 (66) 78 (22) 33 (4) 36 (7) 41 (9) 40 (5) 424 (115)
PALEOCOMMUNITY RECURRENCE
FIGURE 3—Relationship between number of specimens and number
of taxa (Richness, S). Plots describe data including all genera (bi-
valves and brachiopods) and all collections with S $ 2. The majority
of collections include fewer than 40 specimens. (A) Number of collec-
tions with a given total abundance (number of specimens). (B) Num-
ber of collections with a given richness. (C) Cross-plot of total abun-
dance versus richness. Each cross represents a collection.
with regard to paleoecological associations. This is not to
say that fossil co-occurrence proves co-occurrence during
life, which is rarely demonstrable in the rock record. How-
ever, a long history of actualistic live-dead comparisons of
shelly, marine macrofauna indicates that death assem-
blages, and ultimately fossil assemblages, faithfully re-
cord compositional differences between different environ-
ments (Kidwell and Bosence, 1991; Martin, 1999).
All analyses presented here were performed using pres-
ence/absence data rather than abundance data for several
reasons. First, actualistic studies and numerical models
indicate that, although relative abundance data in death
assemblages do reflect living abundances to some degree
(Kidwell and Flessa, 1995), they also experience modifica-
tion due to differences in taphonomic susceptibility among
taxa and environments (Cummins et al., 1986; Staff et al.,
1986; Staff and Powell, 1988; Powell et al., 1989; Miller
and Cummins, 1990; Olszewski and West, 1997). Second,
447
the small number of specimens in many collections means
that relative and rank abundances are poorly constrained.
Third, different collecting methods bias samples in differ-
ent ways. For example, large bulk collections yield reliable
abundance data (Bennington and Rutherford, 1999), but
surface collecting often results in samples with elevated
diversity relative to the number of specimens because the
aim in the field was to maximize the number of different
taxa. Finally, presence/absence data are less sensitive to
differences between collections and are, therefore, more
likely to indicate a higher level of similarity between any
two collections than absolute abundances or abundance
rankings (Rahel, 1990). As it turns out, this bias is in op-
position to our findings and strengthens the interpreta-
tion presented below. This is not to say that relative abun-
dance data are inappropriate for paleoecological studies.
In many cases, where sampling procedures are strictly
controlled and taphonomy and lithology are more uniform
than this study, relative abundances can reveal patterns
of ecological or taphonomic significance that are obscured
by presence/absence data (Olszewski and West, 1997).
ANALYSES AND RESULTS
Data were analyzed using a variety of techniques, each
with a different purpose. Exploratory approaches included
cluster analysis (R-mode, Q-mode, and two-way) and or-
dination (correspondence analysis). To test for changes in
paleoecological associations through time, randomization
and resampling approaches were applied (Crowley, 1992),
here dubbed ‘‘recurrence analysis.’’
External information for collections included geograph-
ic location, stratigraphic position, and lithology. External
information for genera included membership in higher
taxonomic groups.
In some cases, collections below a specified richness or
taxa below a specified rarity were discarded (subsets listed
in Table 1). This is because outliers, consisting of a few
small collections or rare taxa, often obscure broader pat-
terns when using exploratory statistical techniques. Anal-
yses were initially performed using all data but were
pared down to see if interpretable patterns became more
clear; the results included here were selected for the pat-
terns they show and presentability. The characteristics of
the data used in each analysis are described in the figure
captions.
Cluster Analysis
Clustering was conducted using the hierarchical, un-
weighted pair group method with arithmetic averaging
(UPGMA). The aim was to examine how collections and
taxa group together and whether a clear classification
emerges from the data. The similarity metric used was the
Dice coefficient:
SD 5 2c/(Ni 1 Nj) (1),
c 5 number of taxonomic co-occurrences between collec-
tions i and j, Ni, Nj 5 total number of occurrences in i and
j, respectively. This definition is in the context of Q-mode
analysis (clustering of collections), but the coefficient also
can be used for R-mode analysis (clustering of taxa). Note
that other names for the same metric are the Sorenson
448 OLSZEWSKI & PATZKOWSKY

and Czekanowski coefficients and that equation (1) is the

binary reduction of the ‘‘percentage similarity’’ and ‘‘Bray-
Curtis measure’’ (Sepkoski, 1974). The Dice coefficient
was chosen for several reasons. First, in its fully numerical
form, this metric has been found to reflect ecological differ-
ences very effectively (Faith et al., 1987). Second, it cor-
rects for sparseness in the data matrix, thereby decreasing
the influence of rare taxa and low-diversity collections in
the overall result (Archer and Maples, 1989). Third, like
the related Jaccard coefficient (S D 5 2Sj/(Sj11), Sj 5 Jac-
card coefficient), it does not count mutual absences, which
can severely skew results using sparse paleoecological
data matrices (Maples and Archer, 1988). Unlike the Jac-
card coefficient, however, the Dice has a wider and more
symmetrical spread of expected values in randomized
tests (Archer and Maples, 1987).
R-mode analysis of brachiopods and bivalves together
(Fig. 4), using only collections with a taxonomic richness of
4 or more, reveals three major groupings—a brachiopod
cluster, a bivalve cluster, and a group of rare taxa. Both
major clusters have a ‘‘chain’’ topology, suggesting that
taxa were added one by one to a single, growing cluster
rather than being segregated into subgroups and then ag-
glomerated. The rare taxa are grouped at very low levels of
similarity, indicating that they do not form a cluster based
on strong association. Within the brachiopods, Crurithyr-
is, Hystriculina, and Lissochonetes form a subcluster.
These are all small taxa typically found in dark gray to
black, pyritic mudrocks containing high organic carbon.
Collections are usually numerically dominated by one spe-
cies. These characteristics indicate a physiologically
stressful environment due to low oxygen concentrations in
the water column (Boardman et al., 1984; Kammer et al.,
1986).
Q-mode and two-way cluster analyses of the same data
used to generate figure 4 yield very large dendrograms
that could not be shown in a single journal figure; hence,
the subset of the data collected as part of this study (‘‘new
collections’’) is presented instead (Fig. 5). Comparison
with analyses of the entire data set, which can be found in
Olszewski (2000), have a similar geographic and strati-
FIGURE 4—R-mode cluster analysis of all genera using collections
graphic distribution (Table 2), and show the same basic
with S $ 4. Clusters are labeled by their dominant taxa: brachiopods,
patterns as described here. bivalves, and rare taxa. Numbers in parentheses are genus identifi-
The Q-mode analysis of collections (Fig. 5) reveals a cation numbers provided for comparison with other figures (see Fig.
large number of clusters. Some, but not all, show strong 7 for key). ‘B’ and ‘C’ signify brachiopod and bivalve, respectively.
associations with lithology. For example, H has a high pro- Other numbers represent the total number of occurrences of each
portion of carbonate collections, and D is entirely siliciclas- genus in this data set. Note the low number of occurrences in the
group of rare taxa and that degree of association within the other
tic. In addition, there is some correspondence between
clusters correlates to rarity. Apparent ‘‘polytomies’’ are an artifact of
clusters and collections from particular composite se- the software used (SPSS) and do not represent unresolved associa-
quences. For example, all the collections from composite tions.
sequence VI are included in cluster A, many collections
from sequence IV occur in D, and many collections from
sequence V are seen in G, H, and I. ly, this is interpreted to reflect an environmental gradient
The two-way cluster analysis (Fig. 5) relates taxonomic from well-to-poorly oxygenated conditions (Boardman et
content to collection clusters. Q-mode clusters G, H, and I al., 1984), although taxa from both ends of the gradient
clearly segregate from the other clusters because they are also can be found together. Note that although bivalves
dominated by bivalves rather than brachiopods. The clus- and brachiopods are clearly segregated, they are not mu-
ters of rare taxa are easily revealed as such. Within the tually exclusive—there are collections in almost all of the
low-diversity brachiopod clusters, a trend can be seen clusters that include representatives of both major taxa.
starting at the Derbyia-Neochonetes-Composita (taxa 9, 23, The last cluster analysis to be presented here includes
and 7) R-mode subcluster and migrating to the previously only bivalves (Fig. 6). The aim was to see if there were any
described dysoxic subcluster (Crurithyris-Hystriculina- patterns within this group that were being swamped by
Lissochonetes; taxa 8, 14, and19). As mentioned previous- including them with the more numerous brachiopod-dom-
PALEOCOMMUNITY RECURRENCE
FIGURE 5—Two-way cluster analysis of ‘‘new collections’’ with S $
2 (key to genera as in Fig. 7); B 5 brachiopod, C 5 bivalve. External
data for collections includes composite sequence (Roman numerals)
and lithology (s 5 siliciclastic, c 5 carbonate).
449
inated collections. In the R-mode, taxa associate in much
the same way as before, and rare taxa are still excluded.
However, the two-way analysis suggests a possible gradi-
ent from Septimyalina (taxon 36) and Schizodus (taxon
35) to Edmondia (taxon 11) and Myalina (Orthomyalina)
(taxon 21).
Ordination Analysis
As a complement to cluster analysis, the data were or-
dinated using correspondence analysis. Ordination helps
to reveal gradients that can be broken up artificially by
clustering, and orders taxa and collections less arbitrarily
than they are depicted by a dendrogram. In addition, or-
dination expresses multidimensional relationships more
effectively than cluster analysis.
Correspondence analysis was chosen for two reasons.
First, Kenkel and Orlóci (1986) found it to be very effective
at extracting known patterns in artificial ecological data
compared to other ordination techniques. Second, corre-
spondence analysis ordinates both taxa and collections in
the same multivariate space (i.e., both R- and Q-mode
analyses are conducted simultaneously), which allows the
two to be directly related (Jongmann et al., 1995).
A disadvantage of correspondence analysis is that it can
produce an ‘‘arch’’ effect. This is the result of compression
at the ends of ordination axes and a systematic, often qua-
dratic, relationship between axes. These problems can be
corrected by non-linear rescaling (Hill and Gauch, 1980),
but we chose not to do so for several reasons. First, non-
linear detrending is a ‘‘brute force’’ re-adjustment of the
pattern that often can lead to loss of ecologically meaning-
ful information (Pielou, 1984). If patterns are readily in-
terpretable, as is the case in the present analyses, there is
no need for adjustment. Second, Minchin (1987) and Ken-
kel and Orlóci (1986) found that non-linear detrending of-
ten does not improve simulated patterns and can even ex-
acerbate distortion of the ordination space. Ter Braak
(1995) provides a more comprehensive discussion of these
issues, including their mathematical basis and recom-
mended solutions.
Analyzing both brachiopods and bivalves together (Fig.
7A) reveals that they are segregated strongly in ordina-
tion space. Almost all brachiopod genera have positive val-
ues, while the bivalves are negative without exception.
Demarcating the R-mode clusters of genera from figure 4
on the ordination shows that results of the two analyses
are quite consistent. Note that, although the cluster
marked ‘‘Rare Taxa’’ appears to sit within the brachiopod
cluster, it is separated in higher ordination dimensions
where it is pulled out of the plane defined by the ‘‘Bivalve
Dominated’’ and ‘‘Brachiopod Dominated’’ clusters. The
slight overlap of bivalve and brachiopod genera involves
the bivalves Wilkingia (taxon 40) and Pteronites (taxon 3),
and the brachiopod Juresania (taxon 16) (the other taxa in
the region of overlap are rare and, therefore, their posi-
tions in ordination space are constrained poorly and pro-
vide limited information on their relationships). This is
consistent with direct observations in the field, where
Wilkingia and Pteronites are often the only bivalves in a
collection otherwise dominated by brachiopods and other
open-marine groups, and where Juresania seems to be
characteristic of nearshore environments otherwise domi-
450
FIGURE 6—Two-way cluster analysis of bivalve genera and collec-
tions with S $ 2 (key to genera as in Fig. 7). External data for collec-
tions includes composite sequence (Roman numerals) and lithology
(s 5 siliciclastic, c 5 carbonate).
OLSZEWSKI & PATZKOWSKY
FIGURE 7—Correspondence analyses of genera. (A) All genera (bi-
valves circled), all collections with S $ 4. Outlined regions are R-mode
clusters from figure 4. Note the gap (indicated) between brachiopod-
and bivalve-dominated biofacies. (B) Brachiopod genera, collections
with S $ 2. (C) Bivalve genera, collections with S $ 3. Percent of
total inertia is shown for each axis. Units are mean standard devia-
tions. 1 5 Anthraconeilopsis, 2 5 Aviculopecten, 3 5 Pteronites, 4 5
Beecheria, 5 5 Cancrinella, 6 5 Clavicosta, 7 5 Composita, 8 5
Crurithyris, 9 5 Derbyia, 10 5 Echinaria, 11 5 Edmondia, 12 5 En-
teletes, 13 5 Hustedia, 14 5 Hystriculina, 15 5 Isogramma, 16 5
Juresania, 17 5 Leptalosia, 18 5 Linoproductus, 19 5 Lissochonetes,
20 5 Meekella, 21 5 Myalina (Orthomyalina), 22 5 Myalina (Myali-
nella), 23 5 Neochonetes, 24 5 Neospirifer, 25 5 Paleyoldia, 26 5
Permophorus, 27 5 Petrocrania, 28 5 Promytilus, 29 5 Pseudomon-
otis, 30 5 Pteria, 31 5 Punctospirifer, 32 5 Retaria, 33 5 Reticulatia,
34 5 Rhipidomella, 35 5 Schizodus, 36 5 Septimyalina, 37 5 Strob-
lochondria, 38 5 Volsellina, 39 5 Wellerella, 40 5 Wilkingia.
PALEOCOMMUNITY RECURRENCE 451

nated by mollusks. The wide gap between most of the bi-

valves and the brachiopods (marked in Fig. 7A) suggests a
strong discontinuity between the two main compositional
groups. The first axis not only separates brachiopods from
bivalves, it also strings the bivalves into a compositional
gradient. Axis 2 extracts a gradient in the brachiopods
perpendicular to the bivalve gradient of axis 1. This or-
thogonal relationship suggests that the gradients within
the two groups are largely independent of one another.
Ordinating the two main taxonomic groups further re-
fines the patterns seen in the large ordination. In the bra-
chiopods (Fig. 7B), the first axis, which is very consistent
with the second axis of figure 7A, segregates Rhipidomella
(34), Crurithyris (8), Wellerella (39), Lissochonetes (19),
Hustedia (13), and Hystriculina (14) from the rest. As de-
scribed above in the cluster analyses, these genera are all
relatively small (,1.5 cm across) and often very abundant
in facies interpreted as dysoxic deposits. The opposite end
of the gradient is dominated primarily by Juresania (16)
and some rare taxa (e.g., Echinaria [10], Cancrinella [5],
Meekella [20]). The second axis further pulls out these rar-
ities but is otherwise uninterpretable.
In the bivalve ordination (Fig. 7C), the gradient along
the first axis separates Pteronites (3), Wilkingia (40), and
Schizodus (35) at the negative extreme, and Edmondia
(11), Pseudomonotis (29), and Myalina (Orthomyalina)
(21) at the other. Although not identical, this is consistent
with the faunal gradient from open-marine to relatively
restricted coastal or lagoonal environments interpreted in
the cluster analyses.
Because correspondence analysis performs both R-mode
and Q-mode analyses simultaneously (Figs. 7 and 8), it is
possible to examine patterns of other important variables
in the same ordination space, thereby directly relating
taxa to environment and time. An important difference be-
tween figures 7A and 8A (R and Q-modes of the same or-
dination space) is the lack of two distinct clumps in the
scatter of collections (Fig. 8A). This suggests that the seg-
regation of brachiopods and bivalves is not an artifact re-
sulting from uneven sampling at the ends of a continuous
gradient—i.e., the correspondence analysis shows the po-
sition of the intermediate collections. Also investigated
was the relationship of taxa to lithology. Comparison of
figures 7A and 8A indicates that both bivalves and bra-
chiopods can be found in either siliciclastic or carbonate
rocks. The contrast between these two main lithologies is
one of the most prominent in the study interval, yet these FIGURE 8—Correspondence analysis of collections. Solid circles 5
analyses suggest that it does not restrict the distribution carbonates; empty circles 5 siliciclastics; two-tone circles 5 mix of
of the two main taxonomic groups. Rather, Olszewski carbonates and siliciclastics. Large circles represent means for the
(2000) recognized complete paralic to open marine gradi- two lithologies and brackets indicate one standard deviation on each
ents based on lithology in both siliciclastic and carbonate side. These are the same ordination spaces as shown in figure 7. (A)
facies, suggesting that the benthic organisms analyzed All genera, collections with S $ 4. (B) Brachiopod genera, S $ 2. (C)
Bivalve genera, S $ 3. Percentages of total inertia are shown for each
here were more sensitive to the onshore-offshore gradient axis. Units are mean standard deviations.
(a gradient-complex reflecting water depth, environmen-
tal energy, sediment grain size, storm frequency, salinity,
etc.) than the mineral composition of the substrate. Previ- val. Using all taxa (Fig. 9), the average value of each cycle
ous work by Miller (1988, 1989) has found that by the late shifts from nearshore, bivalve-dominated assemblages at
Paleozoic, North American bivalves do not seem to favor the base of each composite sequence to open marine, bra-
terrigenous over carbonate environments, as they seem to chiopod-dominated assemblages through the middle of
have earlier in the Paleozoic. each composite sequence.
To investigate change in communities through time, the The same approach using only brachiopods (Fig. 10)
first axis correspondence-analysis values of collections shows an overall trend through the entire study interval
were plotted against meter-scale cycles in the study inter- from well-oxygenated to poorly-oxygenated assemblages
452 OLSZEWSKI & PATZKOWSKY

except for a brief reversal in the lower part of composite se-

quence V. Composite sequences I and II are dominated by
well-oxygenated faunas, sequence III has equal numbers
of low and high oxygen collections, whereas sequences IV
and V are dominated by low oxygen faunas (despite the
switchback in composite sequence V, it also contains many
of the most dysoxic brachiopod assemblages). Note that
dysoxic collections do not increase at the expense of well-
oxygenated collections; rather, increasingly low-oxygen
points are added to a continuous succession of high oxygen
points. Cycle 51 represents a return to well-oxygenated
conditions, which is consistent with findings based on co-
nodonts (Boardman et al., 1995).
Lastly, with respect to paleocommunity recurrence, it is
of interest to compare the faunal gradients of each com-
posite sequence separately (Fig. 11). As mentioned earlier,
each composite sequence represents, in effect, a naturally
replicated experiment in repopulating the Midcontinent
platform after subaerial exposure. In the brachiopod plots,
dysoxic taxa (Rhipidomella [34], Wellerella [39], Crurithyr-
is [8], Hystriculina [14], Lissochonetes [19]) and nearshore
taxa (Juresania [16], Linoproductus [18]) are indicated. In
all sequences, the dysoxic group and the nearshore group
form distinct patches in the ordination space, but they do
not always fall at the ends of the ordination-defined gra-
dients.
The inferred end-members of the bivalve gradient also
are indicated—Wilkingia (40), Pteronites (3), and Schizo-
dus (35) at the open-marine end, and Edmondia (11), Per-
mophorus (26), Pseudomonotis (29), and Promytilus (28) at
the restricted end (Fig. 11). The bivalve collections are far
less numerous than the brachiopod collections; hence,
their plots are less well constrained in terms of faunal re-
lationships. For example, sequence IV, although it shows
the gradient nicely, is based on only five collections (Table
1). Sequences I and III show the two patches segregated,
but sequences II and V show them overlapping. Exploring
higher axes failed to improve the pattern. Also noteworthy
is the distribution of taxa within these groups, which is not
consistent from sequence to sequence. This suggests that
genera in this data set are not tightly restricted to partic-
ular associations. The statistical significance of this sug-
gestion is tested using recurrence analysis.

Recurrence Analysis
In addition to exploratory methods, tests were also con-
ducted for stratigraphic recurrence of paleoecologic asso-
ciations using methods similar to those described by
Clarke and Warwick (1994; see also Ivany and Baumiller,
1998). Taxonomic associations in a set of fossil collections
can be described completely by an R-mode matrix of simi-
larity coefficients between every pair of taxa. Such a ma-

←
FIGURE 9—First axis correspondence analysis values for all collec-
tions (all genera, S $ 4) plotted against stratigraphic position. Crosses
represent individual collections, dots represent mean values for each
meter-scale cycle. Composite sequence boundaries are shown by dot-
ted lines. Gray stripe shows interpreted general trend of stronger bi-
valve influence at the base of each composite sequence followed by
increased brachiopod dominance upward.
PALEOCOMMUNITY RECURRENCE 453

FIGURE 11—Correspondence analyses of brachiopod and bivalve

genera by composite sequence (key to genera as in Fig. 7). Percent
of total inertia is shown for each axis. Units are mean standard devi-
ations. Outlined patches represent inferred gradient end-members
(see text for discussion).

←
FIGURE 10—First-axis correspondence-analysis values for all collec-
tions (brachiopod genera, S $ 2) plotted against stratigraphic position.
Crosses represent individual collections, dots represent mean values
for each meter-scale cycle. Composite sequence boundaries are
shown by dotted lines. Gray stripe shows interpreted general trend of
increasingly dysoxic faunal signatures through study interval.
454 OLSZEWSKI & PATZKOWSKY

trix was built using the Dice coefficient and all the collec-
tions (S$2) from each composite sequence. These are the
same data that were ordinated for each sequence (Fig. 11),
so they can be generally compared with the results of the
exploratory techniques. Such comparisons should not be
taken too literally because correspondence analysis, al-
though it is an effective means of visualizing complex pa-
leoecological patterns, does not make use of the Dice coef-
ficient. Recurrence of faunal associations was measured
by comparing the Dice coefficient matrices for each pair of
composite sequences using Spearman’s rank correlation
coefficient (r). Note that only taxa occurring in both se-
quences can be included using this approach.
The recurrence coefficients obtained in this manner can
be tested for significance, which measures whether the r
based on real data is significantly greater than if taxa
were distributed randomly among collections. Standard
significance tables for Spearman’s rank correlation coeffi-
cient are not valid in this case for two reasons. First, they
assume a null hypothesis of r 5 0, but non-random struc-
ture (i.e., r.0) can occur in similarity matrices (i.e., the R-
mode Dice matrices used here) due to differences in the
number of occurrences of different taxa, regardless of
whether there are any ecological associations. Second, un-
derlying distributions are not normal (Clarke and War-
wick, 1994). To deal with these issues, a randomization
approach was adopted (Crowley, 1992). The occurrences of
each taxon were randomized among all the collections for
each of two composite sequences. By doing so, the number
of occurrences of each genus did not change, just the dis-
tribution of co-occurrences between genera. Occurrences
were shuffled randomly within taxa until all collections
had at least one occurrence. Note that this loosens the re-
striction of at least two occurrences per collection in the
original data matrices and makes the test conservative in
recognizing recurrence by widening the calculated signifi-
cance values. After both data matrices intended for com-
parison were randomized, their similarity matrices and
correlation coefficient were calculated in the same manner
as the original data. This was performed 1000 times to
build a distribution of correlation coefficients, the top and
bottom 2.5% of which were used to determine the 95% sig-
nificance values. If the correlation coefficient from the real
data lies outside this range, the statistical null hypothesis
(i.e., that the degree of similarity between taxonomic as-
sociations in the two sequences being tested would be pos-
sible with just random associations of taxa) can be reject-
ed. In other words, if the recurrence coefficient lies outside
the 95% range, it is reasonably certain that there is a non-
random degree of recurrence of taxonomic associations be-
FIGURE 12—Randomized recurrence significance. Triangles,
tween sequences. squares, and circles indicate measured Spearman Rank Correlation
According to the randomization results (Fig. 12), the de- Coefficient using all genera (A), brachiopods (B), and bivalves (C),
gree of faunal association using all taxa (brachiopods and respectively. I-bars indicate 95% two-sided confidence intervals based
bivalves) is significantly recurrent (except for sequence I on randomized data. Roman numerals indicate composite sequences
versus IV). In contrast, the gradients within each group (Fig. 1). If the measured value falls outside the confidence interval,
then it is outside the expected degree of recurrence if genus associ-
are generally not recurrent. This suggests that the segre-
ations were random.
gation of bivalves and brachiopods is a real aspect of the
data set, while the faunal associations within each group
are weak. ing bivalve and brachiopod associations, the aim is to de-
Whether or not the correlation coefficients differ signif- termine whether one is more recurrent than the other. To
icantly from random associations, they still describe the test this, a bootstrap procedure was used (Efron and Tib-
amount of recurrence in the structure of the data, which shirani, 1991): rather than randomize the data matrices,
may differ between the two taxonomic groups. In compar- they were sampled with replacement by randomly picking
PALEOCOMMUNITY RECURRENCE
FIGURE 13—Bootstrapped confidence intervals (CI’s) for recurrence.
Squares and circles indicate measured Spearman Rank Correlation
Coefficients using brachiopod and bivalve genera, respectively. I-bars
indicate 95% two-sided confidence intervals based on bootstrapped
resampling. Roman numerals indicate composite sequences (Fig. 1).
If any of these cases showed the measured values of each group
falling outside the confidence interval of the other, then that would
indicate that the degree of recurrence was significantly different. This
is not the case in any of these comparisons.
entire collections from the original data set. The difference
between this procedure and the randomization approach
is that the bootstrapped matrix is composed of collections
known to be samples reflecting real species associations in
each composite sequence. (The randomized matrices shuf-
fled occurrences; hence, any non-random species associa-
tions originally present were destroyed.) The bootstrap
procedure addresses the question: if the two sequences
were resampled many times over, retaining any structure
of taxonomic associations present in the collections, what
would be the range of correlation coefficients? Repeating
the bootstrap 1000 times and using the top and bottom
2.5% of the distribution of the correlation coefficients pro-
vides 95% confidence intervals to bracket the true values.
When comparing bivalves and brachiopods for differences
in degree of recurrence, if either of the two true correlation
coefficients falls within the 95% bracket of the other
group, the two are not considered significantly different.
This is a very stringent criterion, but this choice helps
compensate for the tendency of bootstrapping to inherent-
ly underestimate the true range of variation. In addition,
note that bootstrapped occurrence matrices were created
so that every taxon occurred at least once, slightly loosen-
ing the original restrictions of two or more occurrences per
taxon and making the test conservative.
The results of the bootstrapped tests of whether the re-
currence coefficients differ between the two groups are
presented in figure 13. Although there are numerous oc-
currences of r for one biofacies falling outside the range of
the other, because the exclusion was not mutual, they
were not counted as significant. These findings indicate
that given the number and nature of the collections, there
is no consistently documentable difference in the strength
with which bivalve and brachiopod genera adhere to their
respective environmental gradients.
455
Summary of Results
In the Pennsylvanian-Permian Midcontinent, two dis-
tinct biofacies can be distinguished—one dominated by bi-
valves and the other by brachiopods. This is the over-
whelming pattern in both the cluster analyses and ordi-
nations of these data, and is consistent with the findings of
previous workers in the region (Elias, 1937; Mudge and
Yochelson, 1962). Within each of these two biofacies, pre-
viously unrecognized compositional gradients can be re-
solved. In the brachiopods, one end of the gradient is inter-
preted as representing dysoxic environments (dominated
by relatively small taxa like Crurithyris, Rhipidomella,
Wellerella, Lissochonetes, and Hystriculina). The other end
of the gradient is represented by Juresania and Linoprod-
uctus, which are interpreted to have lived in more near-
shore environments. In the bivalve biofacies, Wilkingia
and Pteronites are associated frequently with brachiopods,
suggesting they preferred open marine conditions, while
other bivalves like Edmondia occupied more restricted
nearshore settings.
Ordination of collections from each composite sequence
suggests that these broad gradients recurred each time
marine conditions were re-established on the platform,
but the particular order of specific taxa along the gradi-
ents does not strictly recur. Recurrence analysis using a
randomization approach confirms that bivalves and bra-
chiopods remain segregated in each cycle, but the gradi-
ents within each biofacies do not recur to a degree signifi-
cantly distinguishable from random associations. Boot-
strapping recurrence coefficients indicates that the rigidi-
ty (or lack thereof) of the order of species is not
significantly different between the brachiopod gradient
and bivalve gradient.
At first glance, the lack of gradient recurrence may
seem contradictory with the results of correspondence
analysis, which show interpretable environmental trends
within the two groups. However, this simply indicates that
taxa within the two communities are not strictly limited in
their associations—they may co-occur with certain taxa
more often than others, as depicted by the ordinations, but
their range of tolerances relative to the amount of environ-
mental change along the gradients allows them to occur
with most other taxa in the paleocommunity. This lack of
strictly limited associations means that recurrence is dif-
ficult to detect because the order of taxa observed every
time the gradient is resampled will be quite variable.
DISCUSSION
Nature of Pennsylvanian-Permian Paleocommunities
of the Midcontinent
The approach taken in this investigation is related
closely to gradient analysis as described by Whittaker
(1967). Rather than classifying assemblages into commu-
nity types, he was interested in how species assemblages
change across a landscape in response to a changing envi-
ronmental complex. Springer and Miller (1990) advocated
adopting such an approach in paleoecological studies.
Whittaker’s concept is summarized in figure 14. The
‘‘environmental gradient’’ is the sum of interrelated envi-
ronmental factors that affect the organisms of the com-
456
FIGURE 14—Graphical representation of ecological composition gra-
dients. (A) Taxa showing exclusion and coadaptation. (B) Taxa show-
ing exclusion but no coadaptation. (C) Taxa showing coadaptation but
no exclusion. (D) Taxa showing neither coadaptation nor exclusion.
(E) Exclusion between biofacies, but neither coadaptation nor exclu-
sion within biofacies (after Springer and Miller, 1990; modified from
Whittaker, 1975).
munity. ‘‘Species importance values’’ are typically thought
of as abundances but can be any ecological variable, such
as biomass or frequency of occurrence. When using pres-
ence/absence data, as in this study, the curves can be
thought of as the chance that a species will occur in a col-
lection from a certain point on the gradient. Note that al-
though abundance and commonness (i.e., number of col-
lections in which a taxon occurs) are often related (e.g.,
Fig. 3C), they need not be: some taxa occur in many collec-
tions but rarely in great abundance, and some occur in
profusion but only at a few sites.
There are two related means of understanding how spe-
cies are distributed across an ecological gradient: direct
gradient analysis and indirect gradient analysis. Direct
gradient analysis involves plotting species distributions
against some factor like elevation, moisture, temperature,
water depth, or oxygenation. Such distributions can be fit
with Gaussian curves showing overlapping species distri-
butions in an environment (Fig. 14). Note that the mea-
sured factor need not be the actual control on species dis-
tribution—it can be a proxy for a complex of environmen-
tal factors. In marine paleoecology, biotic gradients often
are associated with water depth when the main control
may actually be a composite of turbidity, substrate tex-
ture, temperature, and oxygenation (Robbins and Bell,
1994).
Indirect gradient analysis arranges samples based on
taxonomic composition. Compositional differences typical-
ly can be related to environmental factors without strict
adherence to a geographic or stratigraphic gradient. That
is to say, collections can be arranged along a gradient (like
OLSZEWSKI & PATZKOWSKY
those in Fig. 14) regardless of their geographic or strati-
graphic position. The Pennsylvanian-Permian Midconti-
nent Platform was a complex mosaic of depositional envi-
ronments disrupted by numerous stratigraphic disconti-
nuities—the outcrop belt rarely shows a simple facies gra-
dient vertically or laterally (Olszewski 2000). However, by
using as many collections as possible, compositional gra-
dients can be well constrained by reducing the gaps be-
tween samples. Correspondence analysis is one approach
to indirect gradient analysis and it provides a maximum
likelihood approximation of Gaussian curves for species
distributions (ter Braak, 1985).
The results presented above—i.e., two biofacies each
with an interpretable but weak gradient (Fig. 7A)—can be
depicted as in figure 14E. Although bivalves and brachio-
pods segregate, there is no pattern indicating competitive
exclusion or coadaptation within either biofacies. This is
more consistent with a Gleasonian model of species inde-
pendence within communities than with a strong Cle-
mentsian interconnection.
However, with regard to species interactions within a
biotic community, Whittaker et al. (1973) very clearly
warned against using the distribution of taxa along a gra-
dient as an indicator of how they divide the habitat at a lo-
cation into niches (see Schoener, 1989 for elaboration).
These patterns each represent a different scale, and the
question of niches (sensu Whittaker et al., 1973) cannot be
addressed using data of the sort used in this study. The
point is that understanding the range of conditions that a
species can occupy along an environmental gradient pro-
vides only limited insight into the nature of interactions
with neighboring species at a single location during life.
Paleocommunity Stasis in the
Pennsylvanian-Permian Midcontinent
Another question of interest to paleoecologists is wheth-
er these paleocommunities are static or not. The present
results indicate that the biofacies-level division into bra-
chiopod-dominated and bivalve-dominated assemblages
in these data is static—that is, it recurs consistently every
time the basinal ecosystem is re-established. On the other
hand, the way taxa fall along faunal gradients (i.e., how
they divide up habitats) within the two biofacies is not
static.
In a comparison of taxonomic turnover in the basin over
12.5 Myr, analysis presented by Olszewski and Patzkows-
ky (in press) found that background turnover in bivalves
and brachiopods was not significantly different, although
their histories of first and last appearance episodes were.
With regard to the relationship between ecological struc-
ture (measured using recurrence) and background turn-
over, the present results are equivocal—neither group dif-
fers from the other in either aspect. If ecological structure
had differed but turnover had not, or vice-versa, then it
could be concluded that one had little influence on the oth-
er. If both aspects were different in both groups then it
could be determined whether stronger ecological structure
fostered or suppressed turnover (depending on whether
the two were positively or negatively correlated).
In contrast to background turnover, the consistent seg-
regation of biofacies suggests that the independence of
turnover episodes (brief periods of elevated first or last ap-
PALEOCOMMUNITY RECURRENCE 457

pearance) in the two groups (Olszewski and Patzkowsky,

in press) may be related to the introduction or elimination
of the habitat conditions required by the taxa composing
each biofacies. For example, the dramatic decrease in bra-
chiopod diversity above the Beattie Limestone Formation
(Olszewski and Patzkowsky, in press) may occur because
some environments favored by the brachiopods were elim-
inated (see the dramatic shift from dysoxic to well-oxygen-
ated brachiopod collections near the top of the study inter-
val in Fig. 10; the base of cycle 51 is the base of the Beattie
Limestone Formation). Habitat destruction (i.e., elimina-
tion of habitable environments) is one of the most efficient
extinction mechanisms in the modern world (Tilman et al.,
1994), and this is consistent with the paleontological re-
sult found in this study.

Comparison with Other Studies

Most recent studies concerning evolution of benthic ma-
rine faunas have focused on the taxonomic composition of
an entire basin over 106 to 107 years (e.g., Brett and Baird,
1995; Tang and Bottjer, 1996; Patzkowsky and Holland,
1997; Olszewski and Patzkowsky, in press), although
some have examined patterns at the finer scale of biofacies
(e.g., Delcourt and Delcourt, 1991; DiMichele and Phillips,
1995, 1996; Westrop, 1996). Some of these studies (Brett
and Baird, 1995) have recognized ecological-evolutionary
(EE) subunits (lasting 3 to 11 Myr) showing a static pat-
tern of taxonomic composition and bounded by brief epi-
sodes of elevated turnover. Morris et al. (1995) suggested
that processes governing ecological interactions might be
responsible for the static patterns within EE subunits.
Such processes would be expected to result in strong re-
currence of biotic gradients between composite sequences
due to strict limitation of the roles and interactions of in-
dividual taxa (however, see comments of Whittaker et al.
[1973] discussed above). In this study, no influence of any
such processes acting within biofacies has been detected.
In addition, no patterns of EE subunit stasis in the faunas
of the Pennsylvanian-Permian Midcontinent platform
have been found (Olszewski and Patzkowsky, in press).
In contrast to long-term, basinal analyses, several stud-
ies of macroinvertebrates are comparable to this one in
terms of duration, resolution, and geographic scale (Ben-
nington and Bambach, 1996; Holterhoff, 1996; Pandolfi,
1996). All of these, like the present one, were based on in-
dividual site collections rather than basinal compilations
and take advantage of stratigraphic cycles to test for con-
sistency of faunas through time. The findings of each are
briefly reviewed here to search for broader similarities
and differences.
Bennington and Bambach (1996) examined fossil as-
semblages from four major marine incursions in Pennsyl-
vanian strata of the Appalachians. These recurred every
0.4 to 2.5 Myr (about the same as the composite sequences
of the Midcontinent) and contain many of the same taxa as
the data presented here. The aim of their study was to de-
termine how similar individual collections had to be before
they could be considered the same. They did this by com-
paring collections from different locations in the same cy-
cle, comparing collections between cycles, and resampling
individual collections to constrain the range of variation.
They found that there were significant differences in
FIGURE 15—Graphic explanation of difference between analysis of
Bennington and Bambach (1996) and this study. Resampling of sites
permitted Bennington and Bambach (1996) to constrain the range of
compositional variation at points along an environmental gradient
(sensu Whittaker, as in Fig. 14). The approach taken here, based on
numerous collections from many points along the environmental gra-
dient, does not allow testing of individual collections for compositional
similarity, but does constrain the expected range of compositional var-
iation along an environmental gradient.
many (but not all) collections from individual cycles, and
in all comparisons between cycles. They found recurrence
only at the level of ‘‘paleocommunity type’’—equivalent to
the biofacies identified in this study.
These results are consistent with the present study, al-
though the scale of analysis is somewhat different. Large-
scale associations between cycles were compared in this
work, whereas Bennington and Bambach (1996) focused
on reproducibility of individual collections (Fig. 15). In
both studies, biofacies or ‘‘paleocommunity types’’ recur
but show a wide range of internal variation that is not
structured in a recurrent manner.
Holterhoff (1996) examined crinoid associations from
three Upper Pennsylvanian cycles of the northern Mid-
continent (just a few million years older than rocks of the
present study). His cycles are similar in structure and du-
ration to the present composite sequences. He found five
paleocommunity types arrayed along an onshore-offshore
gradient. Three of these paleocommunity types recur in
each cycle (the two others are absent in all but one due to
absence of sufficiently offshore depositional environ-
ments). One of these biofacies was unique to the trans-
gressive phase of the cycles and absent from the regressive
portions.
458
These results are also consistent with the present study,
although a ‘‘no-analog’’ assemblage (Overpeck et al., 1992)
unique to only one portion of a composite sequence has not
been recognized. Like the present results and those of
Bennington and Bambach (1996), none of Holterhoff’s
(1996) individual collections appear identical, although
they do fall into distinct associations that recur from cycle
to cycle.
Lastly, Pandolfi (1996) examined Pleistocene coral-reef
assemblages from Papua New Guinea. He examined nine
cycles over a 95-kyr period, comparable in resolution to
the meter-scale cycles of the Midcontinent, but much finer
than Midcontinent composite sequences. At three differ-
ent locations, he measured transects from reef crest to reef
slope and compared assemblages from those two environ-
ments from place to place and through time. He found
greater differences between locations than through time
at any single location; this contrasts to other studies of
Quaternary terrestrial and level-bottom marine commu-
nities. As a result, Pandolfi (1996) suggested that coral-
reef dynamics distinctly differ from these other systems.
An important difference that Pandolfi (1996) points out
between his work and some other studies is the difference
in temporal scale—many other studies either focused on
much shorter ecological time scales or much longer pale-
ontological time scales. This led him to suggest that differ-
ent patterns may be emerging at different scales of study.
Although the temporal resolution used herein is signifi-
cantly coarser than his, similarities are seen between Pan-
dolfi’s (1996) results and the present investigation. Be-
cause it was not possible to resample the same point on an
environmental gradient through time, results of the pre-
sent study only can be compared to his individual tran-
sects through time. However, if his three sites are consid-
ered as points along an environmental gradient (not nec-
essarily parallel to their geographic trend), then that gra-
dient recurs very consistently from cycle to cycle just as
the Midcontinent bivalve-brachiopod gradient does.
Overall, these three studies are consistent with the find-
ing of this study that biofacies or paleocommunity types,
as segments of environmental gradients, do recur consis-
tently, even when they experience severe environmental
disruption and must be reassembled from their constitu-
ent taxa. Individual points on environmental gradients
(collections or local paleocommunities) are never identical
when resampled through time, but often do show a degree
of interpretable consistency. This probably reflects the
preferences and tolerances of individual taxa rather than
some degree of coadaptation or evolutionary integration.
CONCLUSIONS
(1) Paleoecological analysis of a large data set of fossil
occurrences from Pennsylvanian-Permian rocks of the
northern Midcontinent indicates the presence of two bio-
facies, one dominated by brachiopods and the other by bi-
valves.
(2) Within these two biofacies, compositional gradients
can be recognized. The brachiopod gradient is interpreted
to reflect degree of water-column oxygenation. The bivalve
gradient ranges from open-marine conditions to more en-
vironmentally restricted conditions, but the controlling
OLSZEWSKI & PATZKOWSKY
factors (temperature, salinity, oxygenation, etc.) are un-
clear.
(3) Testing for the recurrence of taxonomic associations
each time marine conditions were re-established indicates
that segregation of the biofacies is strongly recurrent, but
the order of species along gradients within the biofacies is
not. The groups of genera defining the compositional gra-
dients described above do roughly recur, but the specific
relationships among individual taxa do not.
(4) With regard to stasis, a pattern of recurrent associa-
tions within biofacies that is statistically distinguishable
from random taxonomic associations cannot be identified.
On the other hand, the segregation of the bivalve and bra-
chiopod biofacies is significantly recurrent, but probably
reflects different environmental preferences between
these two groups rather than coevolutionary integration of
paleocommunities.
ACKNOWLEDGMENTS
This work was completed in partial fulfillment of T.D.
Olszewski’s doctoral dissertation in the Department of
Geosciences, The Pennsylvania State University (avail-
able through the Penn State Electronic Theses and Disser-
tations website at: http://etda.libraries.psu.edu/). Discus-
sions with R.R. West and D.R. Boardman II in the field
provided insights to the stratigraphy and paleoecology of
the Midcontinent. Thanks to A.I. Miller and an anony-
mous reviewer for thoughtful and encouraging comments
that helped to improve this article. Grants from the Na-
tional Geographic Society, the Geological Society of Amer-
ica, Sigma Xi, and the Krynine Fund of the Penn State De-
partment of Geosciences made this work possible.
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ACCEPTED MARCH 30, 2001