Beruflich Dokumente
Kultur Dokumente
183
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
184
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
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,
187
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).
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
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
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
Small-scale
Lithology
Sorting
Abrasion
Fragmentation
Associated Skeletal
Parts
Rare
Orientation
Body Sizes
Interpretative notes
Sedimentary context
Taphonomic attributes
193
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. . . . ~
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
195
A.
B.
~ !
S
"
/fl
~
/~
~
[
A___2A ~ '
,~ ~
~~.~-------~,--,~
~'~:~:=~. . . . .
~-"~-"~~':'~
~ ,
~
~i.~
~ ~ ' ~ ~ f ~
~
K ~
.~
A'
B'
~
'
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
196
Conclusion
197
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