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

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

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

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 A. SEDIMENTATION PATTERN

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). 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.

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-

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Vertebrate

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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

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~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 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

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-

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
Sedimentary context

Channel-fill Above basal lags, usually in middle to upper parts of channels Discontinuous, thin, coarse beds, thicker fine-grained units Mudstones, silts, clays, fine sands, nodule conglomerates Size-sorting in coarser sediments; variable to poor sorting otherwise Edges fresh to rounded, usually fresh in mudstones Variable; more complete in finer sediments Variable; more frequent in mudstones Variable alignment with paleocurrent; random in mudstones; often at angles to a horizontal plane A wide range usually present, including microfauna Bones at death site or transported short distances; most are autochthonous with respect to the local channel Channels are abandoned and have sporadic, waning flow with minor reworking of bank and bedload sediments

Large-scale Small-scale Lithology

Taphonomic attributes

Lower parts of channels or erosional troughs Basal lag deposits, scour pockets, channels within channels Sands, gravels, mudclast and nodule conglomerates Larger, heavier, robust elements more common (e.g., jaws, teeth); usually well-sorted Edges often rounded, bone pebbles common Variable; usually broken parts Rare Commonly aligned with paleocurrent; usually horizontal Variable; large usually more common Bones usually allochthonous; may represent large areas of the drainage basin Channels represent active drainages with recurring energetic flow and reworking of banks and bedload

Sorting

Abrasion Fragmentation Associated Skeletal Parts Orientation

Body Sizes
Interpretative notes

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
A S S U M E C O N S T A N T B O N E INPUT :

T ,at. o,
Sediment Accumulation

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

~ /

O ...... B ...... of Bone Input a,,S.....

B D o e c n o e r n s p D o ~ e c e o B r n o p n o e ~ s e
Quality and Quantity of P . . . . . . .

~asterThanTl~ey~eBuried
dB .... ~

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___2 A~' ,~ ~ ~ , ~ ~i.~ ~ ~ ' ~ ~ f ~ ~ K ~ .~

A' B 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

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.

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

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-controlled alluvial 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. 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