Palaeogeography, Palaeoclimatology, Palaeoecology, 63 (1988): 183 199

Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands

183

VERTEBRATE PRESERVATION IN FLUVIAL CHANNELS

A N N A K. B E H R E N S M E Y E R

Department of Paleobiology, NHB-E207 MRC 121, Smithsonian Institution, Washington, DC 20560 (U.S.A.)
(Received July 21, 1987)

Abstract
Behrensmeyer, A. K., 1988. Vertebrate preservation in fluvial channels. Palaeogeogr., Palaeoclimatol., Palaeoecol.,
63:183 199.
Two taphonomic modes for attritional vertebrate assemblages in channels are proposed, based on the sedimentary
context of the vertebrate remains and taphonomic features of the bones themselves. The channel-lag mode includes
bones that are buried with coarse lithologies near the bases of active channels. The channel-fill mode occurs in finegrained to mixed fills of abandoned channels. The extreme for a channel-lag assemblage would be a cluster of
allochthonous, abraded, unidentiflable fragments, and the extreme for a channel-fill assemblage would be a cluster of
autochthonous, unbraded, complete skeletons. Between these extremes there is a broad spectrum of possible
taphonomic histories for bones in channels, but distinct channel-lag vs. channel-fill modes can be recognized in fluvial
deposits in different tectonic and climatic settings throughout the Phanerozoic. Physical and biological processes that
affect the different modes produce different samples of vertebrate paleocommunities, with bones in the channel-lag
mode representing transported remains from a variety of habitats, whereas channel-fill assemblages are more
autochthonous and habitat-specific.
Channel facies, channel pattern, and alluvial architecture are used to develop hypotheses concerning how the
taphonomic modes relate to different scales of fluvial processes. Fluvial systems with numerous abandoned channels
provide more sites for preservation of relatively complete fossil vertebrates in channel-fills, while systems that
continually rework sediments by lateral migration preserve more vertebrate remains as channel-lags. Large-scale
physical controls on channel pattern and fluvial architecture probably have had significant effects on the quality and
quantity of the verrtebrate record throughout the history of land vertebrates.
Taphonomic modes provide a basis for comparing faunas with similar preservational histories throughout the
geologic record, and they can help to minimize biases in important paleobiological parameters such as diversity
estimates and the timing of appearance and extinction events.

Introduction
A s i g n i f i c a n t p a r t of t h e v e r t e b r a t e fossil
record occurs within fluvial channel deposits
and has been affected by sedimentary processes
associated w i t h c h a n n e l f o r m a t i o n a n d deposit i o n . C h a n n e l s c a n be c u t g r a d u a l l y o r i n s t a n t a n e o u s l y , t h e y c a n move l a t e r a l l y or vertically t h r o u g h time, a n d they e v e n t u a l l y are
filled w i t h s e d i m e n t s r a n g i n g f r o m c o a r s e
c o n g l o m e r a t e to m u d a n d p l a n t d e b r i s . T h e s e
p r o c e s s e s r e s u l t i n a w i d e r a n g e of t a p h o n o m i c

histories for bones preserved in channel dep o s i t s . S o m e of t h e p a t t e r n s of p r e s e r v a t i o n i n

0031-0182/88/$03.50

channels recur in different rock sequences

t h r o u g h o u t the h i s t o r y of l a n d vertebrates.
A t a p h o n o m i c m o d e is d e f i n e d h e r e as a
r e c u r r i n g p a t t e r n of p r e s e r v a t i o n of o r g a n i c
remains in a particular sedimentary context,
a c c o m p a n i e d by c h a r a c t e r i s t i c t a p h o n o m i c
f e a t u r e s . T h e g o a l of t h i s p a p e r is to d e v e l o p
the h y p o t h e s i s t h a t two t a p h o n o m i c modes,
" c h a n n e l - l a g " a n d " c h a n n e l - f i l l ''1, o c c u r w i t h

1Channel-fill and channel-lag will be hyphenated when

referring to the taphonomic modes but will not be
hyphenated when used in a more general sedimentological
sense, e.g., "channel filling", "channel lag deposits".

1988 Elsevier Science Publishers B.V.

184

different frequencies in different types of

fluvial regimens. These two modes represent
end points on a range of possible taphonomic
histories from transported attritional bone
assemblages to untransported attritional assemblages. They are not all-inclusive, and
other modes could be defined based on descriptive criteria (e.g., density of fossil material),
inferred causes of death (e.g., catastrophic vs.
attritional), or other characters. The channellag and channel-fill modes (as well as other
modes not discussed in this paper) preserve
different types of biological and ecological
information, resulting in biases that affect
interpretations of vertebrate evolution, extinction, and paleocommunity structure.
V e r t e b r a t e b o n e s in c h a n n e l s
Paleontologists have long recognized the
association between vertebrate fossils and
channel deposits. Rapid burial by energetic
flow in channels is an obvious way to preserve
bones, although it also may damage them and
reduce their value as paleontological specimens. Preservation in channel deposits is often
taken as an indication that carcasses and/or
individual bones experienced substantial
transport prior to burial, and that this period
of transport left its signature on the composition of the bone assemblage. The assumption
that bones in channels usually are transported
also implies that they represent mixtures of
animals from different habitats, and this
affects how the preserved vertebrates are used
in paleoecological reconstructions (Shotwell,
1958; Behrensmeyer, 1982).
A review of literature on fossil vertebrates
found in channels shows that there is wide
variability in lithologies associated with bone
assemblages as well as in their taphonomic
features. Often bones are found as part of~'lag ''
deposits, defined here as winnowed and sorted
residues composed of relatively large or heavy
particles that are above the threshold competence (transporting ability) of local currents,
Bones from sand and gravel channel deposits
include lag assemblages associated with con-

glomeratic lenses (e.g., Olson, 1962; Carroll

et al., 1972; Hlavin, 1972; Behrensmeyer, 1975;
Cross et al., 1979; Salisbury, 1982; Eberth and
Berman, 1983; Badgley, 1986), dispersed, disarticulated remains (e.g., Wolff, 1973; Badam and
Ganjoo, 1986); mass accumulations of bones
(e.g., Lawton, 1977), and isolated skeletons or
partial skeletons (e.g., Olson, 1962; Gradzinski,
1970; Dodson, 1971; Gradzinski and Jerzykiewicz, 1974). Fossils are also found in channels
filled with poorly sorted mixtures of mudclast
conglomerates, sand and mudstone (e.g., Voorhies, 1969; Hunt, 1978; Stewart, 1981; Berman
et al., 1985; Behrensmeyer, 1987) and finertextured sediment including organic debris
(e.g., Hook and Ferm, 1985). The vertebrate
remains found in examples given above range
from abraded fragments to undamaged whole
skeletons.
Obviously processes associated with channels can promote fossil preservation, but the
wide variation in sediment type and bone
condition indicates that such processes do not
always involve energetic currents and rapid
sedimentation rates. Within the overall channel context, there may be a range of taphonomic histories linked to different patterns of
channel formation and filling. Prior generalizations concerning the taphonomy of bones in
channels need reexamination based on theoretical considerations and a range of examples
from the fossil record.
Particular fluvial formations often appear to
have characteristic patterns of vertebrate preservation that persist through significant
periods of time. In some stratigraphic sequences fossils occur in both channel and
overbank lithofacies (e.g., Olson, 1962;
Behrensmeyer, 1975; Dodson et al., 1980; Badgley, 1986); in others remains are found almost
exclusively in overbank deposits (e.g., Smith,
R . M . H . , 1980; Bown and Kraus, 1981; Kraus
et al., 1985) or exclusively in channels (e.g.,
Gradzinski, 1970; Dodson, 1971; Salisbury,
1982). Such patterns suggest large-scale sedimentological and/or taphonomic controls on
how vertebrates are preserved in different
fluvial systems.

185
In the following paper, relationships between fluvial environments and vertebrate
assemblages in the Siwalik sequence of Pakistan and the Permian deposits of Texas will be
used to construct a general hypothesis concerning sedimentological controls on the occurrence of channel-lag and channel-fill modes
in the vertebrate record. Prior to description of
the specific examples from Pakistan and Texas,
the following introduction to channel processes is offered as a basis for understanding
how such processes can affect the occurrence
and frequency of the two different taphonomic
modes,
Channel deposits
Channel deposits bear several classes of
information that pertain to different scales of
processes operating in fluvial environments:
(1) sedimentary textures and structures that
record the rate and mode of local deposition,
(2) evidence for original channel patterns in
sedimentary structures and overall geometry,
which reflect the balance of sediment input,
slope, and other factors at a particular point in
time, (3) evidence for longer-term patterns of
basin subsidence in the preserved shapes of
channel deposits and their occurrence in
stratigraphic sequences. Channel facies, channel patterns, and the geometry or "architecture" of the channel deposits all can be used to
relate fluvial processes to different modes of
vertebrate preservation,
The cutting of a channel by energetic flow
and its subsequent filling with sediment represent two distinct, potentially independent
phases of sedimentary history. The shape of
channel deposit in a stratigraphic sequence
reflects the mode of erosion. Sheet sands result
from sustained bank cutting and deposition by
meandering, braided, or anastomosing streams
in tectonic settings where there is a low rate of
subsidence (Bridge, 1985). Ribbon sands occur
when vertical down-cutting is combined with
lateral erosion within a restricted channel
belt, and shoe-string sands typically result
from down-cutting events with minimal lateral

erosion. The latter are characteristic of chutes,

crevasse splay channels, and some anastamosing channels (Friend et al., 1979; Smith, D. G.
and Smith, N.D., 1980; Smith, D.G. and
Putnam, 1980; Bridge, 1985).
The depositional phase of a channel results
in the sediments and structures used to characterize its flow, although these (including organic remains) may represent only the later
phases of activity when currents are depositing
more than they remove from a particular
reach. Bar structures formed of relatively
coarse sediment are characteristic of laterally
migrating channels with sustained or frequent
high-energy flow, while both coarse and finegrained sediments occur as channel fill deposits in abandoned channels (Fisk, 1944, 1947;
Bridge, 1985). In stratified fluvial deposits, a
sediment package that fills a U-shaped trough
and/or forms a distinct lens indicates an
abandoned channel. Sandy fills are evidence
for gradual abandonment, with the current
maintaining its capacity to transport sand
until filling is complete. In contrast, finegrained channel fills indicate sudden abandonment and either a drastic reduction in the
energy needed to transport coarser sediment or
a barrier (e.g., vegetation) to the supply of
sediment (Bridge, 1985). Mixed fills generally
fine upward and reflect gradual abandonment
with periods of energetic flow (as during floods)
alternating with quiet water deposition or
paleosol formation within the channel. There
is a complete spectrum of channel fills between
the fine and coarse end-members (Fig.l), and a
single abandoned channel segment may have
sand fills at either end and clay fill in its middle
section (Smith, D. G., 1983).
Channel patterns characterize the fluvial
regimen, which ultimately is controlled by
climatic and tectonic processes. Slope, discharge, sediment load and vegetation all are
known to affect channel patterns (Leopold
et al., 1964; Schumm, 1977; Baker, 1978; Rust,
1981). In general, meandering rivers are associated with low slope and low sediment loads
while anastomosed and braided rivers reflect
increased slope and higher sediment loads. All

186
LITHOLOGY

SEDIMENTATION PATTERN

A.

Sapr~l

TIH

B.

Co

D.

Fig.1. A generalized model for channel fills, showing a progression from finer to coarser-grained deposits from A through D.
Hypothetical graphs to the right show the pattern of sedimentation in relation to time, with finer-grained deposits
representing slow, relatively steady deposition. Coarser deposits reflect sporadic erosion and deposition by active currents
and more rapid channel filling overall. Channel fills comparable to B-D (but not A) occur in the Siwalik deposits of Pakistan
and fills comparable to A-D occur in the Lower Permian deposits of Texas. Stippling = sand, gray = sandy to clayey silt,
black = clay or coal.
types of rivers c a n deposit sheet sands w h e n
conditions favor sustained lateral aggradation
(e.g., in stable to slowly subsiding basins)
(Bridge, 1985; K r a u s and Middleton, 1987). In
subsiding basins, the same rivers can form a
different type of fluvial a r c h i t e c t u r e (Allen,
1978), w i t h discrete c h a n n e l belt (i.e., ribbon)
sand bodies dispersed t h r o u g h o v e r b a n k deposits (Bridge, 1985; K r a u s and M i d d l e t o n ,
1987). It a p p e a r s t h a t w h a t e v e r the original
c h a n n e l p a t t e r n , m o r e sheet-like c h a n n e l deposits will be p r e s e r v e d in a r e a s of slow
subsidence, while m o r e d i s c r e t e r i b b o n or
s h o e s t r i n g c h a n n e l bodies will be p r e s e r v e d in
areas of i n c r e a s e d subsidence (e.g., K r a u s and
Middleton, 1987).
A n a s t o m o s i n g c h a n n e l p a t t e r n s often are
associated with rapidly subsiding basins or
o t h e r s i t u a t i o n s w h e r e v e r t i c a l a g g r a d a t i o n is
d o m i n a n t (Friend et al., 1979; Smith, D. G. and
Smith, N . D . , 1980; Smith, N . D . and Cross,

1985; Smith, D . G . , 1986). S u c h systems are

c h a r a c t e r i z e d by f r e q u e n t a v u l s i o n and channel a b a n d o n m e n t , r e s u l t i n g in t r o u g h - s h a p e d
s h o e s t r i n g deposits with c o a r s e to fine-grained
fills. P r e s e r v e d segments of m e a n d e r i n g channels are k n o w n in b o t h fluvial and deltaic
settings (Elliot, 1965; G a r d n e r , 1983; Smith,
R . M . H . , 1987), and in s i t u a t i o n s w h e r e deposit i o n and subsidence were affected by local
s t r u c t u r a l c o n t r o l (Hook and Ferm, this issue).
L o c a l cycles of a v u l s i o n are i n t e r p r e t e d as the
c a u s e of a b a n d o n e d sinuous c h a n n e l s with
mixed to fine-grained fill in fluvial sequences
from areas of m o d e r a t e subsidence (Hopkins,
1985; G o r d o n and Bridge, 1987; B e h r e n s m e y e r ,
1987).
S e d i m e n t o l o g i s t s h a v e t r a d i t i o n a l l y used
v e r t i c a l profiles and s e d i m e n t a r y s t r u c t u r e s to
classify r i v e r p a t t e r n s as m e a n d e r i n g or
braided (Vischer, 1965; Miall, 1978). H o w e v e r ,
it is now a p p a r e n t t h a t similar v e r t i c a l se-

187

quences and sedimentary structures can result

from different kinds of channel patterns (Miall,
1980, 1984). Consequently, there is new emphasis on using lateral control in channel deposits
to establish the characteristics of ancient
rivers (Bridge and Gordon, 1985). Analysis of
fluvial systems also has expanded from the
study of single channels to the interrelationships of multiple channel and overbank sequences, or alluvial architecture (Allen, 1978;
Bridge and Leeder, 1979; Allen and Williams,
1982; Miall, 1987).

laterally discontinuous, fine-grained lithofacies indicating a mosaic of localized depositional settings such as ponds. The sheet sands
are interpreted as channel deposits of a major
river (on the scale of the modern Indus or
Ganges), and the facies along their upper
surfaces as infillings of depressions left after
channel avulsion (Behrensmeyer and Tauxe,
1982; Behrensmeyer, 1987).
Ribbon sands occur within the fine-grained
deposits and vary in frequency throughout the
different formations in the Siwalik Group
(Behrensmeyer, 1987). These sands are usually
single-storied and show little evidence for
point-bar accretion or lateral erosion of floodplain sediments. Their widths range from tens
to hundreds of meters and thicknesses from < 1
to 10 m. They typically have trough-shaped
lower contacts and poorly preserved bedding
structures. Upper contacts are gradational
into silts and clays, often with root traces and
other paleosol features. At the edges of the
trough, the upper part of the channel sand may
pass laterally into sandy levee facies, indicating in situ vertical aggradation. These channels include lenses of carbonate and mudclast
conglomerate, but mud-drapes and other fine-

Siwalik c h a n n e l d e p o s i t s

Types of channels
Throughout the sequence of Miocene formations in northern Pakistan, which spans
approximately 12 m.y., there are two distinctly
different types of sand bodies (Fig.2). Sheet
sands with thicknesses between 6 and 20m
alternate with thicker sequences of finegrained deposits. Internal stratification indicates complex, large-scale bar structures of a
braided or anastomosing, sand-dominated
river. The upper parts of these sheets include
Lower

Chinji

Formation-13.1

m.y.

~,,::::,.:=,:::.:.:.:.:.:.:::~:."

50

0 m.

.
0
[

..:::.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:.:

1.0 k m .
I

~ .......

; - , . , :.:- . . . . . .

Vertebrate

Localities

Channel

Lag Facies

C h a n n e l Fill F a c i e s

Fig.2. Example of fluvial lithofacies in the lower part of the Siwalik deposits of the Potwar Plateau, Pakistan. Vertebrate
localities are concentrated in the middle part of the fill of a large.scale abandoned channel, but also occur in smaller channel
fills, in channel lag facies, and in other contexts in the overbank deposits. Size of bone icon is roughly proportional to
number of fossils at each site. Coarse stippling indicates major sheet sandstones, white represents overbank lithologies.

188
grained sediments are rare. They are interpreted as crevasse-splay channels t h a t were
cut and filled by short-term pulses of overbank
flow which did not cause lateral channel
movement through time. Fossil vertebrates
occur in abundance in lenses of conglomerate,
in the upper, fine-grained parts of the channel,
and in adjacent levee deposits, but almost
never within the channel sands themselves.
Fine-grained and mixed channel fills occur
at a variety of scales, from small trough-shaped
lenses a few meters in width to complex
deposits overlying large-scale, irregular erosional depressions 1 km or more in width,
normal to current direction (Fig.2). The largerscale erosional features are interpreted as
"failed avulsions", in which flow from the
major river eroded very large crevasse channels during floods but then failed to shift
course into these channels (Behrensmeyer,
1987). Analogous large, abandoned channels
occur in braided river systems presently draining the Himalaya Mountains (Gole and Chitale, 1966; Holmes, 1968; Coleman, 1969). The
large Siwalik channel fills have discontinuous
basal conglomeritic lags and some coarse sand
lenses, but the predominant fill is fine sand and
clayey silt, often with complex cut-and-fill
bedding. Thin lenses of mud- and carbonateclast conglomerate within the finer facies
alternate with weakly developed paleosols,
indicating sporadic flow in the abandoned
channel. The degree of bioturbation increases
upward, and the channel fills typically are
capped by silty clays t h a t lack bedding and
have more mature paleosol features. Smaller
scale channel fills are similar, but less complex
internally and finer-grained overall,
Fine-grained channel fills are very similar to
floodplain facies, especially in the large-scale
abandoned channels. Establishing the bottom
and at least one edge of the erosional trough is
the surest way to confirm the existence of an
abandoned channel. In the absence of wellestablished geometry, local textural complexity and well-preserved bedding usually differentiate the channel fill facies from laterally
contiguous floodplain deposits, which are more

homogeneous and lack distinct bedding. Adjacent floodplain deposits also may have a welldeveloped paleosol at the same horizon as the
less mature paleosol capping the channel fill
(Fig.2).

Sedimentary context and taphonomy of

vertebrate remains
Throughout the Siwalik sequence, vertebrate fossils occur in low frequencies in sheet
sands deposited by the major channel belts.
Isolated bones, teeth, and bone pebbles are the
rule, although there are rare occurrences of
partial skeletons or skulls of larger animals.
Typically, bones in this context are fragmentary and abraded. In contrast, some of the
richest fossil localities occur in fine-grained
facies t h a t fill depressions on the tops of the
sands. These assemblages include unabraded
skeletal material from a wide range of taxa and
body sizes, with taphonomic features similar to
those in the channel fills described below.
Fossils can be extraordinarily abundant in
fine-grained channel fills, and there is a
marked pattern of association with the middle
to upper parts of these fills rather than with
basal units (about 75//0 of the channel-fill fossil
occurrences follow this pattern; Behrensmeyer, 1987). Concentrations of bones and
bone fragments occur in thin nodule-clast and
mudclast conglomerates and in bioturbated
units of mixed clay, silt, and fine sand. The
former are often size-sorted and include
abraded material, while the latter are characterized by articulated, undamaged skeletal
parts. Micro- and macro-vertebrates occur
together in both of these facies.
Bones larger than 1 cm (maximum diameter)
occur at densities up to 14/m 2 in excavated
samples from the channel fill deposits. These
show extensive pre-burial breakage and surface scratching indicating the combined effects
of carnivore activity and trampling (Behrensmeyer et al., 1986; Behrensmeyer et al., in
press). The subvertical orientation of a number of the excavated specimens is added
evidence t h a t trampling was an important

189
process in bone modification and burial
(Behrensmeyer et al., 1986; Behrensmeyer et
al., in press). Variable orientations of bones in
the conglomeratic facies do not suggest strong,
unidirectional flow, and the presence of skeletal parts with a wide range of hydraulic
equivalences also implies that the assemblages
were not subjected to extensive current action.
Rather, it appears that bones accumulated
through attritional processes within the abandoned channels; some were transported short
distances, winnowed, and concentrated by
sporadic flow, and others were buried without
transport by the vertical build-up of finegrained sediments and by trampling into soft
substrates,
The greatest number of the paleontologically
important localities in the Potwar region occur
in middle, upper, and occasionally lower parts
of fine-grained channel fills. Vertebrate fossils
can occur in fine-grained floodplain deposits
throughout the Siwalik Group, but they are
relatively uncommon in this context. According
to data assembled by Badgley for the Dhok
Pathan Formation (Badgley, 1982: table6-3),
only 7% of the recovered vertebrate fossils
occur in the floodplain context, while 520//o are
from channels or channel margins. The remainder are associated with mud- or carbonate-clast
conglomerate units which probably represent
crevassesplay sheets or shallow crevassesplay
channels. This pattern appears to hold for the
Siwalik sequence in general in the Potwar
Plateau region (Raza, 1983).
The P e r m i a n d e p o s i t s o f T e x a s
The following observations are based on a
brief survey of notable exposures and vertebrate localities in Seymour County, Texas.
While the data are limited compared to what is
available for the Siwalik vertebrate record,
there are similarities in the patterns of bone
occurrence across approximately 260 million
years, in markedly different biotic and tectonic
settings. The Permian examples also demonstrate the value of comparative taphonomy, in
which general patterns of preservation are

supported by comparing the taphonomy of

organic concentrations from different temporal or physical settings.
Deposits of the Wichita and Clear Fork
Groups in central Texas represent fluviodeltaic environments where vertebrates lived
and died over a considerable span of time in the
Early Permian (Case, 1915; Romer, 1935, 1958;
Olson and Beerbower, 1953; Olson, 1958; Dalquest and Kocurko, 1986). Associations of
faunas and subenvironments in the Clear Fork
Group provided the basis for E.C. Olson's
pioneering work in vertebrate community evolution (Olson, 1952, 1976, 1977; Olson and
Mead, 1982). More recently, paleoecological as
well as taxonomic history has been extended
downward into the Wichita Group by N. Hotton (pers. comm., 1984). The Permian terrestrial facies are predominantly red siltstones
and mudstones, with some formations characterized by increased amounts of sandstone.
There are occasional thin, intercalated units of
marine limestones, indicating that the overall
setting was much closer to deltaic and marine
environments than the Siwalik sequence.
Moreover, the tectonic setting was quite different; Permian fluvial deposits accumulated on a
stable craton whereas the Siwalik sequence
resulted from the collision between India and
Asia. In spite of these different settings, local
sedimentary processes deposited similar fluvial
facies representing coarse and fine-grained
channel fills, floodplains, and levees.
Channel deposits in the Belle Plains and
Arroyo Formations (Wichita Group) include
sheet sands and small-scale ( < 1 0 2 m wide)
abandoned channels with fine-grained to mixed
fills. Sheet sandstones are better exposed and
apparently more abundant in the Arroyo
Formation and are characterized by wellpreserved lateral accretion surfaces indicative
of large-scale (> 102 m wide) meandering channels (Edwards et al., 1983). Vertebrate fossils
are uncommon in the sheet sands, as also noted
for overlying formations (Olson, 1962).
Fine-grained channel fill deposits in the
lower Wichita Group were first reported by
Case (1915). They are often difficult to distin-

190

A.
~1 m.[

~10

rn.

B.
~1 m.[

~I0

m,

Fig.3. Two examples of channel-fill contexts for fossil vertebrates from the Lower Permian deposits of Texas. A. The
Craddock Bone Quarry in the Arroyo Formation, Seymour County, showing fill of channel (probably a chute) cut into pointbar deposits of a major sheet sandstone. Unfossiliferous carbonate nodule conglomerate occurs at the base, and the rest of
the fill consists of red to purple mudstone with pedogenic carbonate nodules and rare lenses of sand. Well-preserved
vertebrate material occurs in the lower 1.5 m of the channel-fill. B. Channel-fill in the Belle Plains Formation, Seymour
County, showing context of size-sorted vertebrate material associated with thin nodule conglomerates within fine sands and
silts. Basal unit is a purple silty clay, with minor sand at the base. Channel is cut into red overbank silts.
g u i s h f r o m s u r r o u n d i n g floodplain deposits
a n d m a y be m o r e a b u n d a n t in t h e r e d b e d
sequences than they appear. Typically the
l o w e r c o n t a c t s c a n be d e t e r m i n e d b y a lag
c o n g l o m e r a t e of m u d c l a s t s a n d c a r b o n a t e nodules, w h i c h defines t h e e r o s i o n a l t r o u g h . Fills
v a r y f r o m d r a b gray, p u r p l e or red m u d s t o n e s
to i n t e r b e d d e d fine s a n d s a n d silts w i t h wellp r e s e r v e d b e d d i n g (Fig.3). T h i n , d i s c o n t i n u o u s
beds of l i m o n i t i c c o n g l o m e r a t e w i t h m u d a n d
c a r b o n a t e c l a s t s o c c u r a t v a r i o u s levels w i t h i n
t h e s e c h a n n e l fills. M u d s t o n e s a r e t y p i c a l l y
b i o t u r b a t e d a n d o c c a s i o n a l l y p r e s e r v e roott r a c e s a n d b u r r o w s i n d i c a t i v e of t h e i n i t i a l
p h a s e s of soil f o r m a t i o n ,
Fossil v e r t e b r a t e s o c c u r in t h e t h i n conglome r a t e s w i t h i n c h a n n e l fills, a n d t h e r e m a i n s
t y p i c a l l y a r e f r a g m e n t a r y , size-sorted, a n d in
v a r y i n g s t a t e s f r o m f r e s h to a b r a d e d . Wellp r e s e r v e d b o n e s o c c u r in s o m e of t h e muds t o n e s a n d a r e c h a r a c t e r i z e d by a r t i c u l a t e d or
a s s o c i a t e d p a r t i a l s k e l e t o n s , a wide r a n g e of
b o d y a n d b o n e sizes, a n d g e n e r a l l y fresh,
u n a b r a d e d b o n e surfaces. P l a n t r e m a i n s also

o c c u r in t h e d r a b b e r m u d s t o n e s a n d m a y be
i n t e r b e d d e d w i t h t h e v e r t e b r a t e - b e a r i n g cong l o m e r a t e s (a r a r e i n s t a n c e of c o - o c c u r r i n g
p l a n t a n d v e r t e b r a t e r e m a i n s in t h e s e strata).
In c o n t r a s t to t h e c o n g l o m e r a t e s w i t h i n t h e
c h a n n e l fills, lags at t h e b a s e s of c h a n n e l s
generally lack vertebrate remains.
V e r t e b r a t e fossils o c c u r in o t h e r c o n t e x t s
w i t h i n t h e o v e r b a n k d e p o s i t s of t h e Belle
Plains and Arroyo Formations, including
" c l u s t e r s " of a r t i c u l a t e d i n d i v i d u a l s of t h e
s a m e t a x o n in floodplain facies a n d t h e occasional p a r t i a l s k e l e t o n or i s o l a t e d b o n e w i t h i n
t h e s h e e t s a n d s t o n e s . T h e p a t t e r n s o b s e r v e d in
t h e s e f o r m a t i o n s a r e b r o a d l y s i m i l a r to O l s o n ' s
d e s c r i p t i o n s of fossil o c c u r r e n c e s in t h e overl y i n g L o w e r P e r m i a n d e p o s i t s of T e x a s a n d
O k l a h o m a (Olson a n d B e e r b o w e r , 1953; Olson,
1958, 1962). T h e a s s o c i a t i o n of b o n e assemb l a g e s w i t h small-scale a b a n d o n e d c h a n n e l
deposits also h a s b e e n d o c u m e n t e d for c o m p a r able s t r a t a on t h e w e s t e r n side of t h e L o w e r
P e r m i a n s e a w a y in N e w M e x i c o ( E b e r t h a n d
B e r m a n , 1983; B e r m a n et al., 1985).

191
Similarities to the Miocene examples of
vertebrate fossil occurrences include: (1) the
tendency for the best-preserved assemblages to
be within channel fills, (2) the pattern of
association with fine-grained deposits above
the basal-lag conglomerate, (3)the occurrence
of size-sorted fragmentary material in thin
nodule and mudclast conglomerates within the
channel fills, and (4) preservation of associated
skeletal material in fine-grained units. Taphonomic features of bones associated with the
finer versus the coarser facies of the channel
fills also appear to be similar in the Permian
and Miocene examples. Although the physical
taphonomic processes associated with these
fluvial facies probably have remained fairly
constant through time, biological processes
(e.g., s c a v e n g i n g ) u n d o u b t e d l y have changed,
Detailed comparisons of patterns of breakage
and other features in the Permian bone assemblages and the excavated Miocene samples
must await further taphonomic work in the
Permian deposits,
P a t t e r n s o f v e r t e b r a t e p r e s e r v a t i o n in
channels

The channel-fill and channel-lag modes of

preservation in channels are based on documentation of vertebrate occurrences in the
Pakistan sequence, with supporting evidence
from the Texas Permian (Table I) and published studies of vertebrate occurrences in
channels. These modes are defined primarily
on overall sedimentary context and secondarily on taphonomic features of the bone assemblages. Sedimentary context appears to differentiate them rather clearly, while there is
considerable overlap in taphonomic features,
This overlap may be due in part to present lack
of detailed information on the taphonomy of
bone assemblages from the different sedimentary contexts. However, it also reflects the fact
that similar processes interact with bones in
both channel-lag or channel-fill contexts. Particular channel environments may shift the
degree to which specific physical and biologi-

cal processes such as transport and trampling

affect vertebrate remains, but overlap in their
taphonomic attributes is to be expected.
The channel-lag mode refers to bones in the
lower part of an erosional channel feature
which are in direct association with coarse
clastic material (TableI). The base of the
channel may be eroded into fine-grained deposits or into previous channel deposits, as in
local scours or multistoried sand bodies. Channels may be of any size but are filled with sand
or coarser material and usually represent the
more continuously active rivers in a drainage
basin. Bones have taphonomic features (e.g.,
abraded edges and processes, size-sorting) indicating that they have experienced sustained
interaction with moving water and sediment. A
model for the formation of this mode in
meandering channels has been proposed previously (Behrensmeyer, 1982).
The channel-fill mode refers to bones preserved in mixed to fine-grained deposits that
fill a channel after it has been abandoned by
sustained, active flow. Usually such deposits
occur in the middle to upper parts of a channel
fill, although they also may be found immediately above the basal lag. Vertebrate material
can be associated with thin beds of coarse
clastics, especially mudclast or nodule conglomerates, or with mudstones and clays. A
characteristic of channel-fill assemblages is
t h a t taphonomic features are highly variable,
but there is a larger component of fresh, wellpreserved, material (e.g., unabraded and unsorted bones, articulated skeletons) than in
channel-lag assemblages.
Potential overlap between these two modes
exists when a sand- or gravel-filled channel
occurs within the overall context of an abandoned channel that has predominantly finegrained fill, or vice-versa. In such cases, the
relative scale of the two modes must be
assessed, along with which depositional context
is most relevant to the genesis of the fossil
occurrences.
The common association of fossils with the
middle parts of the Siwalik Miocene and Texas
Permian channel fills indicates that conditions

192
TABLE I
Characteristics of two taphonomic modes in channel vertebrate assemblages composed of attritional skeletal remains (i.e.,
accumulated gradually over periods of 102-104yr, not due to single-event mass deaths)
Channel-lag

Channel-fill

Large-scale

Lower parts of channels or erosional

troughs

Above basal lags, usually in middle to

upper parts of channels

Small-scale

Basal lag deposits, scour pockets,

channels within channels

Discontinuous, thin, coarse beds,

thicker fine-grained units

Lithology

Sands, gravels, mudclast and nodule

conglomerates

Mudstones, silts, clays, fine sands,

nodule conglomerates

Sorting

Larger, heavier, robust elements more

common (e.g., jaws, teeth); usually
well-sorted

Size-sorting in coarser sediments;

variable to poor sorting otherwise

Abrasion

Edges often rounded, bone pebbles

common

Edges fresh to rounded, usually

fresh in mudstones

Fragmentation

Variable; usually broken parts

Variable; more complete in finer

sediments

Associated Skeletal
Parts

Rare

Variable; more frequent in mudstones

Orientation

Commonly aligned with paleocurrent;

usually horizontal

Variable alignment with paleocurrent;

random in mudstones; often at angles
to a horizontal plane

Body Sizes

Variable; large usually more common

A wide range usually present,

including microfauna

Interpretative notes

Bones usually allochthonous; may

represent large areas of the drainage
basin

Bones at death site or transported

short distances; most are autochthonous
with respect to the local channel

Channels represent active drainages

with recurring energetic flow and
reworking of banks and bedload

Channels are abandoned and have

sporadic, waning flow with minor
reworking of bank and bedload sediments

Sedimentary context

Taphonomic attributes

often f a v o r e d bone c o n c e n t r a t i o n d u r i n g waning r a t h e r t h a n a c t i v e p h a s e s of c h a n n e l

activity. A b s e n c e of b o n e s in b a s a l conglome r a t e s a n d s a n d s implies t h a t t h e y were n o t
being c o n c e n t r a t e d d u r i n g the c u t t i n g or
initial filling stages of the c h a n n e l . Circumstances that could promote bone concentration
a n d b u r i a l in a b a n d o n e d c h a n n e l fills include:
(1) t o p o g r a p h i c lows likely to receive a n d
p r o t e c t o r g a n i c remains, (2) c o n c e n t r a t i o n s of
a n i m a l s n e a r a b a n d o n e d c h a n n e l s due to
localized a v a i l a b i l i t y o f food a n d water, particu l a r l y d u r i n g times of r e s t r i c t e d r e s o u r c e s o n

the alluvial plain ( J a r m a n , 1972; G. H a y n e s ,

pers. comm., 1986), (3) localized t r a n s p o r t ,
sorting, a n d h y d r a u l i c c o n c e n t r a t i o n of bones,
s u c h as m i g h t o c c u r d u r i n g s p o r a d i c floods, (4)
sedimentation
patterns
characterized
by
periods of n o n - d e p o s i t i o n w h e n a t t r i t i o n a l
remains could accumulate, alternating with
periods of r e l a t i v e l y r a p i d d e p o s i t i o n t h a t
w o u l d b u r y bone-rich horizons, (5) low r a t e s of
d e s t r u c t i o n by soil o r g a n i s m s a n d c h e m i c a l
d i s s o l u t i o n b e c a u s e of t h e c o m b i n e d effects of
r a p i d b u r i a l a n d (perhaps) a n a e r o b i c conditions. B o n e p r e s e r v a t i o n m i g h t also be en-

193

hanced by local concentrations of nutrients

such as Ca and P from increased biotic activity
in this environment,
In modern environments, inactive channels
often harbor bodies of standing water and
dense patches of vegetation (Fisk, 1947; Jarman, 1972; Gagliano and Howard, 1983), making them attractive places for herbivores and
predators. Attritional bone assemblages would
be expected in such situations, and periods of
low sedimentation could result in high bone
densities on and in soils developed within the
channel fill. Trampling by animals within the
abandoned channels would contribute to disarticulation and breakage of bones but also
would enhance burial in soft substrates,
Abandoned channels have a pattern of
clastic sedimentation t h a t might be more
conducive to vertebrate preservation than
other fluvial sub-environments, at least in
some systems (for a contrasting situation with
low clastic input, see Hook and Ferm, this
issue). Given constant input of bones, there
must be a balance between sedimentation and
rates of bone dispersal and decomposition to
create a bone concentration (Fig.4). If sedimentation is too slow, bones will decompose faster
than they can be buried, or they will be
destroyed after burial as soils mature on stable
ASSUMECONSTANTBONEINPUT:

k Bo,~
TooO~.,.0,
.... TooO........
~
~ . - ~

T
,at. o,
Sediment
Accumulation

Oof
......
B......
Bone
Input
a,,S.....

Bo
D
ecnoernspD
o~
eceoBrnopnoe~se

~asterThanTl~ey~eBuried
Quality and Quantity
of P . . . . . . .

d B. . . . ~

Fig.4. Hypothetical relationship between rate of sediment

accumulation and the quality of the vertebrate record for
attritional assemblages in channel deposits. Rate on the yaxis refers to individual sedimentary units t h a t preserve
bones; absolute values for optimal rates of sediment
accumulation for bone preservation would depend on bone
input and on bone sizes. (See text for further explanation.)

land surfaces. If sedimentation is too rapid,

there will be insufficient time for bones to
accumulate, and they are likely to be dispersed
or damaged by repeated interaction with other
sedimentary particles and high-energy currents. The timing of sedimentation events also
is important, and there may be an ~'on-off"
periodicity at particular stages of channel
filling which is optimal for bone preservation.
All of the factors enumerated above can
occur in non-channel environments, but apparently at least some of them were more frequently associated with abandoned channels
in the Miocene and Permian fluvial deposits.
For these examples, it is not yet possible to say
which factors were most important in creating
the observed taphonomic patterns. However,
the scarcity of bones in paleosols that cap
channel deposits and in floodplain paleosols in
Miocene and Permian examples suggests that
they were less likely to be preserved where the
more mature soils developed in these fluvial
systems. This implies that rate and mode of
sedimentation is one of the most critical
factors promoting preservation in channel fills.
The channel-fill mode of fossil occurrence
persists throughout the Miocene sequence in
northern Pakistan and transcends major
changes in the fluvial systems during this
period of time (Behrensmeyer and Tauxe, 1982;
Behrensmeyer, 1987). Therefore it appears that
conditions favoring bone preservation are
linked more strongly to local processes associated with abandoned channels than to the
overall fluvial regimen. This also is supported
by the occurrence of the channel-fill mode in
the Permian deposits of Texas, which were
formed by a different fluvial system in a
different tectonic (and probably a different
climatic) setting. However, the overall vertebrate record in the Siwalik sequence is also
affected by the number of a b a n d o n e d c h a n n e l s
t h a t occur at different stratigraphic levels
(Behrensmeyer, 1987), and this appears to be
related more directly to the fluvial system and
its tectonic setting.
Paleoecological implications of the channelfill and channel-lag modes can be inferred from

194
their differing context and taphonomic features. In channel-lag assemblages, attritional
vertebrate remains may be derived from various sources (e.g., channel banks, upstream
drainages) (Behrensmeyer, 1982) but in general
are allochthonous with respect to the depositional site. Multiple cycles of channel erosion
and deposition can be represented in the lag
assemblage, depending on the pattern of channel migration within the fluvial system. In
some cases, this mode also can record a single
event of erosion, transport, and concentration
of attritional bones from a restricted source
area. The species in the assemblage may
represent members of the community or communities inhabiting different environments in
the drainage basin, time-averaged over
102-104 yr (Behrensmeyer, 1982). In channelfill assemblages, the vertebrate remains are
derived from a smaller area within the channel
or from adjacent overbank environments, and
they are autochthonous with respect to the
abandoned channel environment. Time-averaging depends on the rate of filling of the
channel, which can occur in < 102 yr based on
modern examples (Gagliano and Howard,
1983).
General implications for the vertebrate
record
The channel-lag mode has been recognized
for some time and characterizes what many
paleontologists and sedimentologists visualize
when they think of bones in channels. The
channel-fill mode also has been noted previously (e.g., Boy, 1977; Eberth and Berman,
1983), and evidence from the Miocene and
Permian examples discussed above supports
the recurring association of vertebrate remains with channel fills in widely different
time periods and alluvial settings. The broader
significance of the channel fill context in the
vertebrate record probably has been underestimated because it is more difficult to discern in
outcrop than the channel-lag context,
Patterns of occurrence of the channel-lag
and channel-fill modes should be tested further

in fluvial deposits other than the Siwalik

Miocene and Texas Permian sequences. However, these two modes appear to be generally
represented i n t h e v e r t e b r a t e r e c o r d . Processes
associated with channels affect fossil preservation at three discernable levels: (1) local
circumstances that control whether bones are
preserved in channel-lags or channel-fills (e.g.,
rates and modes of depositional events), (2)
characteristics of the fluvial system that affect
the frequency of different types of channels,
(3) local to basin-scale rates of subsidence that
affect fluvial regimen and the ultimate preservation of channel deposits.
In both channel-fill and channel-lag contexts, local channel processes have a major
effect on the balance of bone input versus
destruction or dispersal in creating a fossil
assemblage (Fig.4). Channel pattern (e.g.,
braided, meandering,
anastomosing) and
change through time (lateral migration, avulsion) affect the degree to which bones are
transported, winnowed, abraded, or left undisturbed. Highly sinuous channels are subject to
neck cut-off, and resulting oxbow lakes typically have low clastic input compared to less
sinuous chutes and braid channels (Fisk, 1947;
Allen, 1965; Hook and Ferm, this issue).
Depositional processes affecting bone concentrations in channel fills thus are controlled in
part by channel sinuousity. Braided and anastomosing systems are characterized by multiple channels that repeatedly form and reform
around bars or islands (Leopold and Wolman,
1957; Schumm, 1963), while meandering systerns progressively rework their own deposits
by lateral erosion and deposition, usually over
longer time periods. Channel-lag assemblages
would be subjected to different degrees of
short- versus long-term reworking and concentration in fluvial systems with different channel patterns.
Given local conditions favorable to bone
concentration in the channel-lag and/or the
channel-fill mode, fluvial systems with active,
migrating channels in well-established channel belts would be likely to generate more
channel-lag assemblages, while systems with

195

A.

B.
~ !

S
"
/fl
~

/~
~

[
A___2A ~ '
,~ ~

~~.~-------~,--,~
~'~:~:=~. . . . .
~-"~-"~~':'~

~ ,
~
~i.~
~ ~ ' ~ ~ f ~
~
K ~
.~
A'

B'

~
'

Fig.5. Comparison of a meandering, laterally migrating

fluvial system within a restricted channel belt (A) and an

anastomosing, avulsion-dominated system that is not

confined to a restricted c h a n n e l belt (B), showing the
resulting alluvial architecture and the different types of
channeldeposits. The laterally migrating system generates
sheet sandstones and would be expected to have more

channel-lag vertebrate assemblages than the avulsiondominated system, which would have more channel-fill
assemblages,

frequent avulsion and channel-belt abandonment would have more instances of preservation in the channel-fill mode (Fig.5). Based on
observations in modern environments, abandoned channel environments would be expectedin most river systems. Because of lateral
erosion of a river within its channel belt,
however, deposits in these environments are
repeatedly destroyed until the river avulses to
a new position on the alluvial plain,
The frequencies and scales of different kinds
of secondary channels (e.g., crevasse splay,
chute, and tributary) also would affect the
number of opportunities for the channel-fill

and channel-lag modes, with more secondary

channels probably favoring the former. Both
suspended load (meandering) and bedload
(braided and anastomosing) river systems have
crevassesplays and/or secondary floodplain
channels, with number and scale depending on
overall rates of aggradation and local climatic
conditions that affect flood cycles and floodplain drainage.
There is presently little direct information
on how tectonics and climate control the
frequency of abandoned versus active channels
at a given point in time in different fluvial
settings. It does appear that avulsion-dominated systems frequently characterize areas o f
high sediment input and rapid subsidence
(Smith, 1986), resulting in alluvial architecture
with ribbon or shoe-string sands isolated
within the floodplain deposits (Fig.hb).
Laterally migrating systems that deposit multistoried ribbon and sheet sands are more
typical of lower rates of subsidence and
sediment input (e.g., Kraus and Middleton,
1987). Thus, areas of rapid subsidence would
generate and preserve more instances of the
channel-fill mode, while channel-lag a s s e m blages would occur more frequently in areas of
lower subsidence. The vertebrate record in the
first case would be biased toward well-preserved samples from a particular habitat while
in the second it would represent a broader
habitat spectrum with less complete fossil
material.

Ultimately, the preservation of whole sequences of vertebrate-bearing strata depends

on the large-scale tectonic setting. As suggested by R. Hook (pers. comm., 1986), it is
possible that fluvial deposits at colliding continental margins (e.g., the Siwaliks) contribute
significantly only to the latest part of the
vertebrate record, since older rocks from this
setting have been tectonically deformed and
subducted. The more long-lived deposits are
likely to occur on trailing margins, rift basins,
and stable cratons. Thus, the fossil record of
land vertebrate evolution and paleoecology
may be derived from significantly different
continental settings through the Phanerozoic.

196
Conclusion

The channel-lag vs. channel-fill modes are a

way of classifying vertebrate fossil assemblages according to sedimentary context and
taphonomy and are analogous to "taphofacies"
as defined for marine invertebrate assemblages
(see Speyer and Brett, this issue). The extreme
case of a channel-lag assemblage would be a
cluster of rounded, unidentifiable bone pebbles
in a conglomerate within a channel deposit,
while the opposite extreme would be complete,
articulated skeletons preserved in a sapropel
in a channel-fill. Most channel vertebrate
assemblages lie between these extremes; in
some fluvial systems they will tend toward the
channel-lag mode while in others they will
more commonly occur in channel fills. Both
modes should be present in most fluvial deposits that preserve vertebrates in channels,
but in different frequencies depending on the
fluvial regimen. The relative proportion of
fossil occurrences in these modes can be tested
further to determine if there is a general
correlation with the type of fluvial system and
a link to broader climatic and tectonic controls,
Taphonomic modes provide a basis for comparing faunas with similar preservational histories throughout the geological record,
thereby helping to moderate the effects of
taphonomic biases in diversity estimates, timing of appearance and extinction events, and
other important paleobiological parameters,
Traditionally, paleontologists have combined
samples with different preservational histories
to make the most of anatomical, biostratigraphic, and paleogeographic information in
developing an overview of vertebrate evolution and paleoecology. The recognition of
taphonomic modes can help to refine this
approach because different kinds of samples
from in the fossil record can be matched with
more specific evolutionary and paleoecological
questions. For example, vertebrate fossils from
attritional channel-lag assemblages should
provide the most homogeneous sample of the
overall paleocommunity for the analysis of

basin-scale faunal change through long

periods of time. This also would be a good
taphonomic mode for determining the stratigraphic position of immigration and extinction
events since the appearance or disappearance
of a taxon could be affected by which habitats
are being sampled in a fossil assemblage. The
channel-lag mode would represent different
habitats on the alluvial plain, while other
types of samples from channel fills or floodplain paleosols might be more habitat-specific
(see also Badgley and Gingerich, this issue).
Since channel-fill assemblages provide samples
of smaller areas and shorter time periods, they
are better suited for analyses of habitatspecific paleocommunities as well as anatomical studies of populations from similar environments.
Acknowledgments
The ideas in this paper have benefitted from
productive discussions with A. Aslan, C. Badgley, J. Barry, J. Bridge, G. Haynes, R. Hook,
N. Hotton, L. McRae, H. Sues and S. Wing.
Helpful comments on the manuscript were
provided by C. Badgley, M. Kraus, R. Hook,
and H. Sues. I have greatly appreciated assistance in the field from I. Khan, K. Sheikh and
continuing support from Dr. Ibrahim Shah of
the Geological Survey of Pakistan. The field
work has been supported by Smithsonian
Foreign Currency Program grants to D. Pilbeam and J. Barry (Harvard University) with
additional funding from the Smithsonian Research Opportunities Program.
References
Allen, J. R. L., 1965. A review of the origin and
characteristics of recent alluvial sediments. Sedimentology, 5: 89-191.
Allen, J. R. L., 1970. Physical Processes of Sedimentation.
Allen and Unwin, London, 248 pp.
Allen, J. R. L., 1978.Studies in fluviatile sedimentation: an
exploratory quantitative model for the architecture of
avulsion-controlledalluvial suites. Sediment. Geol., 21:
129-147.
Allen, J. R. L. and Williams, B. P. J., 1982.The architecture
of an alluvial suite: rocks between the Townsend Tuff

197
and Pickard Bay Tuff beds (early Devonian), southwest
Wales. Philos. Trans. R. Soc. Lond. B, 297:51 89.
Badam, G. L. and Ganjoo, R. K., 1986. Preliminary
taphonomicalstudiesofsome Pleistocene f a u n a f r o m t h e
central Narmada Valley, Madhya Pradesh, India.
Palaeogeogr., Palaeoclimatol., Palaeoecol., 53: 335-348.
Badgley, C. E., 1982. Community Reconstruction of a
Siwalik Mammalian Assemblage. Thesis, Yale Univ.,
New Haven, Conn., 364 pp. (unpublished).
Badgley, C. E., 1986. The taphonomy of mammalian fossil
remains from Siwalik rocks of Pakistan. Paleobiology,
12: 119-142.
Badgley, C. E. and Behrensmeyer, A. K., 1980. PaleoecologyofMiddleSiwalik sediment and faunas o f t h e P o t w a r
Plateau. Palaeogeogr., Palaeoclimatol., Palaeoecol., 30:
133-155.
Baker, V. R., 1978. Adjustment of fluvial systems to climate
and source terrain in tropical and subtropical environments. In: A. D. Miall (Editor), Fluvial Sedimentology.
Can. Soc. Pet. Geol., Calgary, pp. 211-230.
Behrensmeyer, A. K., 1975. The taphonomy and paleoecology of Plio-Pleistocene vertebrate assemblages of Lake
Rudolf, Kenya. Bull. Mus. Comp. Zool., 146:473 578.
Behrensmeyer, A. K., 1978. Taphonomic and ecologic
information from bone weathering. Paleobiology, 4:
150-162.
Behrensmeyer, A. K., 1982. Time resolution in fluvial
vertebrate assemblages. Paleobiology, 8:211 227.
Behrensmeyer, A. K., 1987. Miocene fluvial facies and
vertebrate taphonomy in northern Pakistan. In: F.G.
Ethridge, R. M. Flores, M. D. Harvey (Editors), Recent
Developments in Fluvial Sedimentology. Soc. Econ.
Paleontol. Mineral. Spec. Publ., 39:169 176.
Behrensmeyer, A. K. and Kidwell, S. M , 1985. Taphonomy's contributions to paleobiology. Paleobiology, 11:
105 109.
Behrensmeyer, A. K. and Tauxe, L., 1982. Isochronous
fluvial systems in Miocene deposits of northern Pakistan. Sedimentology, 29: 331-352.
Behrensmeyer, A.K., Gordon, K.D. and Yanagi, G.T.,
1986. Trampling as a cause of bone surface damage and
pseudo-cutmarks. Nature, 319: 768-771.
Behrensmeyer, A. K., Gordon, K. D. and Yanagi, G. T., in
press. Non-human bone modification in Miocene fossils
from Pakistan. In: R. Bonnichsen and M. Sorg (Editors),
Bone Modification. Center for the Study of Early Man,
Orono, ME.
Berman, D.S., Reisz, R.R. and Eberth, D.A., 1985.
Ecolsonia cutlerensis, an Early Permian dissorophid
amphibian from the Cutler Formation of north-central
New Mexico. N.M. Bur. Mines Miner. Resour. Circ., 191:
1 31.
Bown, T. M. and Kraus, M. J., 1981. Vertebrate fossilbearing alluvial paleosols (Willwood Formation, northwest Wyoming, U.S.A.): implications for taphonomy,
biostratigraphy, and assemblage analysis. Palaeogeogr.,
Palaeoclimatol., Palaeoecol., 43: 95-128.
Boy, J. A., 1977. Typen und Genese Jungpal~iozoischer
Tetrapoden-Lagerst~itten. Palaeontographica Abt. A,
156:111 167.

Bridge, J. S., 1985. Paleochannel patterns inferred from

alluvial deposits: a critical evaluation. J. Sediment.
Petrol., 55: 579-589.
Bridge, J. S. and Gordon, E. A., 1985. Quantitative
reconstructions of ancient river systems in the Oneonta
Formation, Catskill Magnafacies. Geol. Soc. Am. Spec.
Pap., 201: 163-181.
Bridge, J. S. and Leeder, M. R., 1979. A simulation model of
alluvial stratigraphy. Sedimentology, 26:617 644.
Campbell, K. E. and Frailey, D., 1984. Holocene flooding
and species diversity in southwestern Amazonia. Quat.
Res., 21: 369-375.
Carroll, R.L., Belt, E.S., Dineley, D.L., Baird, D. and
McGregor, D.C., 1972. Vertebrate paleontology of
Eastern Canada. In: Field Guidebook A59, 24th Int. Geol.
Congr., Montreal, 113 pp.
Case, E. C., 1915. The Permo-Carboniferous beds of North
America and their vertebrate fauna. Carnegie Inst.
Wash. Publ., 207:1 176.
Coleman, J. M., 1969. Brahmaputra River: Channel processes and sedimentation. Sediment. Geol., 3(2/3):
129 239.
Cross, A. T., Prouty, C. E. and Bain, R. J., 1979. The
sedimentological and paleontological features of Late
Pennsylvanian and Early Permian rocks of Southeastern Ohio and western West Virginia. Field Guide,
Great Lakes Section, Soc. Econ. Paleontol. Mineral.
Michigan State Univ., East Lansing, Mich., 111 pp.
Dalquest, W. W. and Kocurko, M. J., 1986. Geology and
vertebrate paleontology o f a lower Permian delta margin
in Baylor County, Texas. Southwest. Nat., 31: 477-492.
Dodson, P., 1971. Sedimentology and taphonomy of the
Oldman Formation (Campanian), Dinosaur Provincial
Park, Alberta (Canada). Palaeogeogr., Palaeoclimatol.,
Palaeoecol., 10:21 74.
Dodson, P., Behrensmeyer, A.K., Bakker, R.T. and
McIntosh, J.S., 1980. Taphonomy and paleoecology of
the dinosaur beds of the Jurassic Morrison Formation.
Paleobiology, 6:208 232.
Eberth, D. A. and Berman, D. S., 1983. Sedimentology and
paleontology of Lower Permian fluvial redbeds of northcentral New Mexico - - preliminary report. N.M. Geol.,
5:21 25.
Edwards, M.B., Eriksson, K.A. and Kier, R.S., 1983.
Paleochannel geometry and flow patterns determined
from exhumed Permian point bars in north-central
Texas. J. Sediment. Petrol., 53:1261 1270.
Elliot, R. E., 1965. Swilleys in the coal measures of
Nottinghamshire interpreted as paleo-river courses.
Mercian Geol., 1(2): 133 142.
Fisk, H. N., 1944. Geological Investigation of the Alluvial
Valley of the Lower Mississippi River. Miss. River
Comm., Vicksburg, 69 pp.
Fisk, H. N., 1947. Fine-grained alluvial deposits and their
effect on Mississippi River activity. Miss. River Comm.,
Vicksburg, 82 pp.
Friend, P.F., Slater, M . J . and Williams, R.C., 1979.
Vertical and lateral building of river sandstone bodies,
Ebro Basin, Spain. J. Geol. Soc. Lond., 136: 39-46.
Gagliano, S. M. and Howard, P. C., 1983. The neck cutoff

198
oxbow lake cycle along the lower Mississippi River. In:
Proc. Conf. Rivers '83, Waterway, Port, Coastal and
Ocean Division, ASCE, New Orleans, pp.147-158.
Gardner, T. W., 1983. Paleohydrology and paleomorphology of a Carboniferous, meandering, fluvial sandstone,
J. Sediment. Petrol., 53: 991-1005.
Gole, C. V. and Chitale, S. V., 1966. Inland delta building
activity of the Kosi River. J. Hydraul. Div., Proc. Am.
Soc. Civil Eng., 92: 111-126.
Gordon, E. A. and Bridge, J. S., 1987. Evolution of Catskill
(Upper Devonian) river systems: intra- and extra-basinal
controls. J. Sediment. Petrol., 57: 234-249.
Gradzinski, R., 1970. Sedimentation of dinosaur-bearing
Upper Cretaceous deposits of the Nemegt Basin, Gobi
Desert. Palaeontol. Pol., 21: 147-229.
Gradzinski, R. and Jerzykiewicz, T., 1974. Sedimentation of
the Barun Goyot Formation. Palaeontol. Pol., 30:
111-146.
Hlavin, W. J., 1972. Early Permian vertebrates from the
upper Washington Formation at Belpre, Ohio. In:
T. Arkle (Editor), I.C. White Mem. Symp. Field Trip,
W. Va. Geol. Surv., Morgantown, W. Va., pp. 30-51.
Holmes, D. A., 1968. The recent history of the Indus. Geogr.
J. R. Geogr. Soc. Lond., 134: 367-382.
Hook, R. W. and Ferm, J. C., 1985. A depositional model for
the Linton tetrapod assemblage (Westphalian D, Upper
Carboniferous) and its palaeoenvironmental significance. Philos. Trans. R. Soc. Lond. B, 311:101 109.
Hopkins, J. C., 1985. Channel-fill deposits formed by
aggradation in deeply scoured, superimposed distributaries of the Lower Kootenai Formation (Cretaceous). J.
Sediment. Petrol., 55: 42-52.
Hunt, R. M., 1978. Depositional setting of a Miocene
mammal assemblage, Sioux County, Nebraska (U.S.A.).
Palaeogeogr., Palaeoclimatol., Palaeoecol., 24: 1-52.
Jarman, P. J., 1972. The use of drinking sites, wallows and
salt licks by herbivores in the flooded Middle Zambezi
Valley. E. Afr. Wildl. J., 10: 193-209.
Kidwell, S. M. and Jablonski, D., 1983. Taphonomic
feedback: ecological consequences of shell accumulation. In: J. S. Tevesz and P. L. McCall (Editors), Cycles
in Sedimentation and Evolution. Plenum, New York,
N.Y., pp.195 248.
Kraus, M. J. and Middleton, L. T., 1984. Dissected
paleotopography and baselevel fluctuations in an
ancient fluvial sequence. Geol. Soc. Am. Abstr. with
Program, 16: 227.
Kraus, M. J. and Middleton, L. T., 1987. Contrasting
architecture of two alluvial suites in different structural
settings. In: F. G. Ethridge, R. M. Flores, M. D. Harvey
(Editors), Recent Developments in Fluvial Sedimentology. Soc. Econ. Paleontol. Mineral. Spec. Publ., 39:
253 262.
Kraus, M. J., Bown, T. M., Kvale, E. P. and Vondra, C.F.,
1985. Upper Jurassic/Lower Cretaceous and Paleogene
alluvial sediments of the Bighorn Basin, northwest
Wyoming. In: R.M. Flores and M. Harvey (Editors),
Field Guidebook to Modern and Ancient Fluvial Systems in the United States, Third Int. Fluvial Sedimentol.
Conf., Fort Collins, Colo., pp.19-44,

Lawton, R., 1977. Taphonomy of the dinosaur quarry,

Dinosaur National Monument. Contrib. Geol., Univ.
Wyo., 15: 119-126.
Leopold, L. B. and Wolman, M. G., 1957. River channel
patterns: braided, meandering and straight. U.S. Geol.
Surv. Prof. Pap., 282-B: 39-73.
Leopold, L. B., Wolman, M. G. and Miller, J. P., 1964.
Fluvial Processes in Geomorphology. Freeman, San
Francisco, Calif., 522 pp.
Miall, A. D., 1978. Lithofacies types and verrtical profile
models in braided river deposits: a summary. In: A. D.
Miall (Editor), Fluvial Sedimentology. Can. Soc. Pet.
Geol. Mem., 5: 597-604.
Miall, A. D., 1980. Cyclicity and the facies model concept
in fluvial deposits. Bull. Can. Pet. Geol., 28: 59-80.
Miall, A. D., 1984. Principles of Sedimentary Basin
Analysis. Springer, New York, N.Y., 490pp.
Miall, A. D., 1987. Recent developments in the study of
fluvial facies models. In: F.G. Ethridge, R.M. Flores,
M.D. Harvey (Editors), Recent Developments in Fluvial
Sedimentology. Soc. Econ. Paleontol. Mineral. Spec.
Publ., 39: 1-9.
Olson, E. C., 1952. The evolution of a Permian vertebrate
chronofauna. Evolution, 6: 181-196.
Olson, E. C., 1958. Fauna of the Vale and Choza, 14.
Summary, review, and integration of the geology and
faunas. Fieldiana Geol., 10: 379-448.
Olson, E. C., 1962. Late Permian terrestrial vertebrates,
U.S.A. and U.S.S.R. Trans. Am. Philos. Soc., 52: 1-224,
Olson, E. C., 1976. The exploitation of land by early
tetrapods. In: Morphology and Biology of Reptiles. Linn.
Soc. Symp. Ser., 3: 1-30.
Olson, E. C. and Beerbower, J. R., 1953. The San Angelo
Formation, Permian of Texas, and its vertebrates. J.
Geol., 61: 389-423.
Olson, E. C. and Mead, J. G., 1982. The Vale Formation
(Lower Permian): is vertebrates and paleoecology. Tex.
Mem. Mus. Bull., 29: 1-46.
Reineck, H. E. and Singh, I. B., 1975. Depositional
Sedimentary Environments. Springer, New York, N.Y.,
439 pp.
Raza, S. M., 1983. Taphonomy and paleoecology of Middle
Miocene vertebrate assemblages, southern Potwar Plateau, Pakistan. Thesis. Yale Univ., New Haven, Conn.,
414pp. (unpublished).
Romer, A. S., 1935. Early history of Texas redbeds
vertebrates. Bull. Geol. Soc. Am., 46: 1597-1658.
Romer, A. S., 1958. The Texas Permian redbeds and their
vertebrate fauna. In: T. S. Westoll (Editor), Studies on
Fossil Vertebrates. Atholone Press, London, pp.157-179.
Rust, B. R., 1981. Sedimentation in an arid-zone anastomosing fluvial system: Cooper's Creek, central Australia. J.
Sediment. Petrol., 51: 745-755.
Salisbury, J. P., 1982. Fluvial sedimentology and associated bone distribution in the northwestern corner
(Steveville Region), Dinosaur Provincial Park, Alberta,
Canada. Thesis. Wichita State Univ., Wichita, Kans.,
135 pp. (unpublished).
Schumm, S. A., 1963. A tentative classification of alluvial
river channels. U.S. Geol. Surv. Circ., 477, 10 pp.

199
Schumm, S. A., 1977. The Fluvial System. Wiley, New York,
N.Y., 338 pp.
Shotwell, J. A., 1958. Inter-community relationships in
Hemphillian (mid-Pliocene) mammals. Ecology, 39:
271 282.
Smith, D. G., 1983. Anastomosed fluvial deposits: modern
examples from Western Canada. In: J. D. Collinson and
J. Lewin (Editors), Modern and Ancient Fluvial Systerns. Spec. Publ. Int. Assoc. Sedimentol., 6:155 168.
Smith, D. G., 1986. Anastomosing river deposits, sedimentation rates and basin subsidence, Magdalena River,
northwestern Columbia, South America. Sediment.
Geol., 46:177 196.
Smith, D. G. and Putnam, P. E., 1980. Anastomosed river
deposits: modern and ancient examples in Alberta,
Canada. Can. J. E a r t h Sci., 17: 1396-1406.
Smith, D. G. and Smith, N. D., 1980. Sedimentation in
anastomosed river systems: examples from alluvial
valleys near Banff, Alberta, J. Sediment. Petrol., 50:
157 164.
Smith, N. D. and Cross, T. A., 1985. Avulsion-controlled
fluvial evolution: the Cumberland Marshes, east-central
Saskatchewan. In: Abstr. Third Int. Fluvial Sedimentology Conf., Fort Collins, Colo., p. 36.

Smith, R. M. H., 1980. The lithology, sedimentology and

taphonomy of flood-plain deposits of the Lower Beaufort
(Adelaide Subgroup) strata near Beaufort West. Trans.
Geol. Soc. S. Afr., 83:399 413.
Smith, R. M. H., 1987. Morphology and depositional
history of exhumed Permian point bars in the southwestern Karoo, South Africa. J. Sediment Petrol., 57:19 29.
Stewart, D. J., 1981. A field guide to the Wealden Group of
the Hastings Area and Isle of Wight. In: T. Elliot
(Editor), Field Guides to Modern and Ancient Fluvial
Systems in Britain and Spain, Proc. Second Int. Conf,
Fluvial Sedimentology, Univ. Keele, U.K., 3; pp. 1 30.
Vischer, G. S., 1965. Fluvial processes as interpreted from
ancient and recent fluvial deposits. In: G. V. Middleton
(Editor), Primary Sedimentary Structures and their
Hydrodynamic Interpretation. Soc. Econ. Paleontol.
Mineral. Spec. Publ., 12:84 115.
Voorhies, M. R., 1969. Taphonomy and population dynamics of an early Pliocene vertebrate fauna, Knox County,
Nebraska. Univ. Wyo. Contrib. Geol., Spec. Pap., 1, 69 pp.
Wolff, R. G., 1973. Hydrodynamic sorting and ecology of a
Pleistocene mammalian assemblage from California
(U.S.A.). Palaeogeogr., Palaeoclimatol., Palaeoecol., 13:
91 101.