A New Locality of Fossiliferous Harding Sandstone: Evidence for Freshwater Ordovician

Vertebrates
Author(s): Greg Graffin
Source: Journal of Vertebrate Paleontology, Vol. 12, No. 1 (Mar. 6, 1992), pp. 1-10
Published by: Taylor & Francis, Ltd. on behalf of The Society of Vertebrate Paleontology
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2 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 12, NO. 1, 1992
Sandstone, where mixed assemblages did occur, the
vertebrate fossils were always disarticulated and worn,
indicating that the remains must have been washed
into the sea from streams and rivers. This scenario was
supported by osmoregulatory physiological studies of
Smith (1932).
Denison (1956) cited a general lack of positive ev-
idence supporting the freshwater theory and main-
tained that freshwater deposits were lacking at the old-
est known vertebrate localities. From evidence at the
type locality, he was convinced that the Harding Sand-
stone was deposited in the sea and that the widespread
vertebrate fragments could not have been introduced
by fluvial processes. Mouths of streams and rivers are
locally restricted, he maintained, and yet the bone frag-
ments of the Harding Sandstone are found over a geo-
graphic expanse that stretches from central Colorado
to northern Wyoming. Also, he stressed that the bone
fragments were much larger than the largest sand grains,
which implied that the two could not have been de-
posited simultaneously. The "ostracoderms" must have
fallen onto the sandy sea substratum when they died.
Spjeldnaes (1967) attempted a paleoecological anal-
ysis of the Harding Sandstone and concluded the ver-
tebrate-bearing units were deposited in a restricted ma-
rine environment. This was based on invertebrate
associations at the type locality and also on boron anal-
ysis of certain clays in the sediments. However, the
salinity values obtainable by boron analysis are no
longer accepted as indicating a marine environment
because many terrestrial sediments can yield higher
salinity values than normal marine sediments.
Based primarily on the studies of Denison and
Spjeldnaes, it became generally agreed among pale-
ontologists that the vertebrates from the Harding Sand-
stone were marine animals. But virtually all previous
paleoecological studies of the Harding Sandstone have
suffered from methodological flaws to such an extent
that they have yielded unreliable results. These were,
first, the reliance on paleontological associations with-
out a clear knowledge of the facies relationships, and,
second, the collection of data at or near only one out-
crop, the type section.
METHODOLOGY
A pervasive methodology in paleontology is the de-
termination of past environments based on fossil as-
sociations. The basis for the concept is the fact that
certain fossil taxa consistently co-occur in strata, some-
times correlatable over large geographic distances. It
is possible to get a rough idea of the biotic relationships
of a fossil species in such instances, but it is not nec-
essarily true that the organisms lived together. It is not
uncommon to find references to associations of fossils
from an entire formation. A single formation, however,
is usually composed of multiple strata deposited in
various depositional environments, all of which may
contain fossils. The only paleoenvironmentally mean-
ingful associations thus are those from the same de-
positional environment.
A methodology for determining depositional envi-
ronments has been developing over the past 25 years.
It is rooted in principles put forth in past centuries. In
1894, Johannes Walther stated that "only those facies
... can be superimposed, primarily, that can be ob-
served beside each other at the present time" (Mid-
dleton, 1973). This, called "Walther's Law," allows us
to interpret a sequence of strata as originally juxtaposed
environments of deposition. The documentation of
modern sedimentary processes constitutes the most
important step in the development of the "modern
revolution in stratigraphy" (Miall, 1984). This has led
to the erection of sedimentary facies models (Walker,
1984) that provide the foundation upon which sedi-
mentological interpretations are built. Presently, most
types of modern depositional environments have been
documented and are characterized by particular assem-
blages of sedimentary structures. Sedimentary facies
are defined based on such structures. A given depo-
sitional system, such as a prograding delta, is composed
of many environments, including proximal channels,
distal channels, overbank areas, abandoned delta lobes
and delta front that are recognizable as different facies.
It is clear that within one depositional system many
types of organisms, from vastly different habitats, may
be preserved. For instance, a deltaic system may bury
terrestrial vertebrates in proximal areas and marine
fishes in distal channels. In stratigraphic sections such
disparate forms may appear within the same formation
separated vertically by only a short interval. Without
facies analysis this situation could lead to an incorrect
assessment of biotic relationships. The fossils might
be considered in association, even though the organ-
isms were living in completely different environments.
Although a stratigraphic description of the Harding
Sandstone was published when it was named (Walcott
1892), and later studies revealed its widespread oc-
currence (Behre and Johnson, 1933; Sweet, 1954; Ger-
hard; 1967), the present study is the first published
facies analysis of this important formation. Unfortu-
nately, the type locality near Canon City, Colorado,
where most previous paleontological work has been
done, is a quarry that has been worked for nearly 100
years. It is very difficult to find a complete stratigraphic
section, and many of the fossiliferous strata have been
obliterated. There is, however, a previously unde-
scribed locality about 50 miles west of Canon City that
permits a complete facies analysis.
THE BUSHNELL LAKES LOCALITY
On the upper eastern slope in the northern Sangre
De Cristo range of central Colorado is a cirque basin
that contains the Bushnell Lakes. It is located in San
Isabel National Forest, in Fremont County (Fig. 1),
about seven statute miles (11 km) southwest of Howard
(which is about 18 miles (29 km) southeast of Salida,
Colorado along U.S. Highway 50). The Bushnell Lakes
 GRAFFIN-HARDING SANDSTONE 3

VICINITY
MAP
DENVER
Glenwood Springs

1O Us 6

A0pen Colorado
BUSHNELL LAKES REGION
Springs
SSa

Walsenburg Arkansas

Alamos C mpgrou
"

San Isabel

( (

Forest
GNational
-
1Mile
Scale:

(fN COALDALE
Q0Sn

O
Hayden

Creek.

Scale: 1 MilePeak

(1.6 km) Hayden Pass

FIGURE 1. Location map of the Bushnell Lakes. The region is within San Isabel National Forest. The locality is accessible
by a marked trail from the Hayden Creek campground along county road 249 (lower right of map). Canon City, Colorado is
36 miles (58 km) east along US Highway 50, and Salida, Colorado is 18 miles (29 km) northwest of Howard along Highway 50.
4 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 12, NO. 1, 1992
basin is a glacially carved valley, generally devoid of
vegetation, near the crest of the Sangre De Cristo range.
Three lakes occur in series at progressively lower ele-
vations (pater noster lakes). The third lake in the series
is surrounded by steep canyon walls composed of an
uninterrupted sequence of Ordovician sedimentary
rocks and is the site of the vertebrate locality. Its el-
evation is at about 11,200 feet (3,405 meters). Because
this is above treeline, very little vegetation is present.
This provides excellent exposure of the strata and sub-
tle sedimentary structures. All of its qualities render it
an ideal locality for a facies analysis.
Ordovician sedimentary rocks form a conformable
sequence about 110 meters thick. The sequence is di-
vided into three units, the Lower Ordovician Manitou
Formation, the Middle Ordovician Harding Forma-
tion (=Harding Sandstone), and the Upper Ordovician
Fremont Formation. Both the Manitou and Fremont
formations are primarily composed of dolomite. The
former contains some quartzitic units. Near the base
of the Manitou Formation, the lithoclastic quartz con-
tent of the carbonate rock is never more than seven
percent. The quartz percentage increases upward in the
section until, at about five meters below the contact
with the Harding Sandstone, the carbonate content of
the rock is only about 40 percent, and quartz grains
make up about 60 percent. The Harding Sandstone is
comprised entirely of detrital clastic rocks such as
mudstones, very fine-grained wackes, fine and medi-
um-grained arenites, and highly fossiliferous pebble
conglomerates. The entire Ordovician sequence lies
nonconformably upon Precambrian gneiss.
The contact between the Manitou Formation and
the Harding Sandstone has been defined as an uncon-
formity (Litsey, 1958). At Bushnell Lakes, however,
the sequence appears to be conformable from the base
of the Manitou Formation to the very top of the Fre-
mont Formation. Because the Harding Sandstone is
thus a part of a depositional continuum, it is necessary
to view it in relation to the sedimentary system that
operated during the formation of all three Ordovician
units. This requires a facies analysis of the entire Or-
dovician sequence.
FACIES TYPES
There are four basic types of sedimentary structures
preserved in the Ordovician rocks: 1) parallel lami-
nations in fine-grained units; 2) burrows and other ev-
idence of bioturbation; 3) very low-angle wedge-set
cross-bedding in fine to medium grained units; and 4)
epsilon and trough cross-beds. These, and associated
sedimentological features, can be used to define the
different types of facies that occur at Bushnell Lakes
(Fig. 2).
Laminations occur throughout the Manitou For-
mation and in the upper part of the Fremont Forma-
tion. They are commonly faint, usually about 5 to 10
millimeters thick, and are present in both micritic and
sparitic units. Laminae can be traced laterally many
tens of meters before becoming truncated or obscured.
Units with these characteristics are interpreted as sub-
tidal deposits that were laid down below wave base,
below the critical velocity of ripple formation. The
units that show uninterrupted laminations that extend
laterally for 10 to 100 meters are interpreted as deposits
of the offshore zone (Elliott, 1986).
Almost invariably, laminae are associated with ev-
idence of biological sediment disruption, which is found
in all three formations. Units with laminations grade
upward into zones with worm burrows and other ev-
idence of bioturbation. Often this zone grades upward
into another laminated unit. Extreme bioturbation
tends to erase all sedimentary features, because organ-
isms thoroughly disrupt sediments before lithification.
Churned bedding often results and is recognizable by
the presence of mottling in the sediments and also by
remnant patches of parallel laminations, areas not af-
fected by the biological disruption (Conybeare and
Crook, 1968). Burrows are sometimes preserved with-
in mottled beds in the Manitou Formation.
Due to the high degree of bioturbation, it is often
difficult to find any distinctive sedimentary structures
preserved in the shoreface zone (Elliott, 1986). Thus
determination of shoreface deposits is based primarily
upon churned bedding and secondarily on the position
within a facies sequence. Units that show laminations
interrupted by bioturbation characterize the offshore-
shoreface transition zone.
Very low-angle cross-beds (wedge sets) are found in
all three formations at Bushnell Lakes. They consist
of sandy laminae with slightly discordant contacts. The
scale of bedding is about five to ten centimeters. Bed
sets are 0.5 to 4 meters thick and can be traced laterally
for many tens of meters. These structures are only
found in units with fine or medium granularity that
generally lack mud. They are invariably overlain or
underlain by units showing bioturbation; however, nei-
ther burrows nor churned bedding occurs in units with
low-angle wedge-set cross-beds.
Wedge-sets are very distinctive sedimentary struc-
tures and characterize deposits of the upper foreshore
or beach (Reineck and Singh, 1986). The laminations
are caused by plane-bed deposition during the swash
and backwash of wave action (Elliott, 1986). Upon
handlens inspection, they are seen to be composed of
minor grain size differences between layers. The dis-
cordances in the beds reflect changes in the inclination
of the beach face toward the sea. This results from
differential deposition/erosion rates at different times.
Theoretically, this ratio could change due to season-
ality (stormy and calm seasons) or changing climate
over time.
The fourth type of sedimentary structure found in
the Ordovician rocks at Bushnell Lakes comprises ep-
silon and trough cross-beds. They are found only in
the coarsest units of the Harding Sandstone. Epsilon
cross-beds are characterized by inclined sets of lentic-
6 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 12, NO. 1, 1992
ular sandy beds that form erosive bases. Trough (or
festoon) cross-beds are broadly concave and also form
erosive bases. Often the two types are found within the
same sand body. The scale of bedding is variable but
the largest individual epsilon cross-beds are about one
meter deep. The smallest troughs are only about 10
cm in depth and about 50 cm wide. Some of the epsilon
and trough cross-bedded units contain pebble con-
glomerates that extend laterally for less than 10 meters
and are about one meter thick.
Epsilon cross-beds (Allen, 1963) are features formed
by lateral accumulation of sediment. They are a superb
guide to fluvial facies because each individual bedding
surface represents a point bar from a meandering stream
(Collinson, 1986). The dip indicates the direction of
migration of the deposit through time. This is normal
to the direction of current flow. The nature of stream-
flow causes lateral migration of fluvial channels and
associated deposits. This tendency causes erosion of
underlying sediments and is seen as scoured bedding
surfaces in stratigraphic sections.
The trough cross-beds are also interpreted as chan-
nels. They do not indicate a laterally migrating stream
but represent channel scours that have become infilled
with coarse sediment (Conybeare and Crook, 1968).
They are commonly found on point bars or as indi-
vidual channels within a braided stream system.
The coarse conglomerates are interpreted as channel
"lag" deposits of the lower point bar and stream bed
(Reineck and Singh, 1986). In general, conglomerates
are key indicators of fluvial channel deposits (Collin-
son, 1986) but may not always be present in a fluvial
sequence. They are rather restricted in distribution at
the Bushnell Lakes. Where present, they are found at
the base of, and grade into, sandy units that represent
higher point bar deposits. It is within these units that
the greatest concentration of fossil vertebrate material
is found.
FACIES SEQUENCE
Facies succession at Bushnell Lakes is cyclic. Each
cycle consists of a general upward pattern of shallowing
deposits. The lowest unit is an offshore, subtidal de-
posit characterized by laminations. Overlying that is
a bioturbated shoreface deposit, which, in turn, is over-
lain by an intertidal foreshore (beach) deposit. Rarely,
this is overlain by supratidal fluvial and related facies.
Thus a generalized unit sequence, or facies model, can
be discerned (Fig. 3) that shows such an upward-shal-
lowing trend. A similar facies model was erected as a
generalization for many Paleozoic carbonate sequences
(James, 1984). This model was strictly for carbonates,
but the Bushnell Lakes sequence shows that the model
can be consistent with a mixed carbonate and clastic
succession as well.
The facies relationships suggest that the Harding
Sandstone represents a terrigenous clastic phase of de-
position within a shallowing-upward, carbonate-dom-
inated marine system. It was deposited atop a gently
sloping carbonate ramp (sensu Ahr, 1973), and it de-
fines the zone of clastic influx in such a model (Wilson,
1975). Much of the Harding Sandstone is marine as
suggested by past authors (Denison, 1956; Spjeldnaes,
1967). But its upper portions represent supratidal ter-
restrial deposition, the only occurrence of such in the
entire Ordovician sequence. Detrital quartz began to
enter the system in small quantities during deposition
of the Manitou Formation. Gradually, quartz influx
increased, culminating in progradation of terrestrial
environments over the intertidal. After deposition of
the Harding Sandstone there ceased to be any quartz
introduced. The Fremont Formation is purely carbon-
ate and was laid down in successively deeper water.
FOSSIL DISTRIBUTION
As reported at other localities (Johnson, 1945), the
Manitou Formation at Bushnell Lakes contains poorly
preserved invertebrate fossils that are few in number.
Virtually all are crinoid fragments that occur in the
offshore facies within five meters of the contact with
the Harding Sandstone.
Invertebrate fossils are more abundant and varied
in the Fremont Formation than in the Manitou For-
mation. The remains comprise fragments of brachio-
pods, solitary rugose corals, crinoids, and coralline al-
gae. These occur from the base to five meters above
the upper contact of the Harding Sandstone, within the
shoreface facies. The abundance of fossils in the Fre-
mont Formation is not great, similar to that described
at other localities (Sweet, 1954; Johnson, 1945).
Vertebrate and invertebrate fossils have been re-
ported from the same strata at the type locality of the
Harding Sandstone (Walcott, 1892; Denison, 1956;
Spjeldnaes, 1967). Such "faunal mixing" has played
an important role in bolstering the view that the ver-
tebrates of the Harding Sandstone were marine ani-
mals. At the Bushnell Lakes, however, there are no
invertebrate fossils found in any of the units that con-
tain abundant vertebrate fossils. Furthermore, the
greatest accumulations of vertebrate fossils are restrict-
ed to the supratidal facies (Fig. 4) where they occur in
coarse conglomeratic channel deposits. These rocks are
composed almost exclusively of vertebrate fragments
and rounded quartz granules (Fig. 5). Intertidal units
of the Harding Sandstone contain occasional rare ver-
tebrate fossils. Invertebrate body fossils are absent from
the Harding Sandstone. Although bioturbated sedi-
ments within the Harding Sandstone provide evidence
that invertebrates were present, vertebrate fossils do
not occur in such units.
The data at hand suggest that there is no basis for
assuming the vertebrates of the Harding Sandstone
inhabited the sea. Rather, it is most parsimonious to
assume that they were freshwater animals that lived
in rivers and streams. Presumably, after death, the
elements forming their exoskeletons (tesserae) became
disconnected and were incorporated into the bedload
of streams.
 GRAFFIN-HARDING SANDSTONE 7

SEDIMENTARY
SHALLOWING MODEL
- UPWARD STRUCTURES INTERPRETATION

Terrestrial Epsilonand festoon cross-bedswith Progradational

granularconglomeratechannellags Fluvial System
(highly fossiliferous) interbedded
Supratidal
with laminatedand rippledmudstone.

Low-anglewedge sets and rippled

Intertidal calcareniteor fine to medium Foreshore
sandstone.

Bioturbated(churnedbedding) Sh f
micriticcarbonateor fine sandstone.
Subtidal
Laminated micritic carbonate or very Offshore-
fine sandstone. Shoreface
Transition

FIGURE 3. Shallowing-upwardmodel in a mixed carbonate-clasticsetting.A completesequenceof shoalingdeposits includes

offshore-shorefacetransition,shoreface,foreshore,and fluvial sediments.Thereare five shoalingcycles in the Ordovicianrocks
at Bushnell Lakes (see Fig. 2); only one includes all four types of deposits. The fluvial facies are only found in the Harding
Sandstone.Similar sedimentarystructuresmay be found in both carbonateand clastic deposits. Partiallyadaptedfrom James
(1984).

60- DISCUSSION
The fact that vertebrate and invertebrate fossils gen-
erally do not occur together at Bushnell Lakes is pa-
S40 II Vertebrates leoenvironmentally significant. It implies either that
vertebrates and invertebrates lived together and were
separated after death or that the two groups never lived
20 together and were preserved separately, in different
environments of deposition. Although it is difficult to
prove, the fact that vertebrate fossils are most abun-
dant in the granular conglomeratic inferred fluvial fa-
subtidal intertidal fluvial cies (units that lack any invertebrate fossils) strongly
suggests that the latter scenario is most parsimonious.
Depositional enviroment
Arguments that the vertebrates of the Harding Sand-
stone were marine require one of two unlikely condi-
tions: 1) the vertebrate fragments were washed up-
60
stream from the mouths of streams and came to rest
in fluvial environments, or 2) the fluvial channels rec-

40
~ Inverteb rates

20 FIGURE 4. Histogramof fossil types and distributionsin

the fossiliferous Ordovician sediments at Bushnell Lakes.
The depositional environments were determined by facies
0-
analysis. Although vertebrates are present in some of the
intertidalunits, they are much more abundantin fluvial fa-
subtidal intertidal fluvial cies, suggestingthat they lived in fresh water and that some
remains probably got washed in to the nearshoreenviron-
Depositional enviroment ment. The fossiliferousunits are indicated in Figure 2.
 GRAFFIN- HARDING SANDSTONE
vertebrate fragments in the supratidal deposits, their
paucity in intertidal, and absence in subtidal deposits
is suggestive of a freshwater habitation.
As mentioned above, there are certain intertidal units
within the Harding Sandstone that contain small
amounts of bone (less than 7% of the total rock vol-
ume). These fragments were probably washed into the
intertidal zone from the stream channels, somewhat
analogous to the way the quartz grains of the Manitou
Formation were washed into the intertidal zones. Both
bone and quartz grains likely had a continental prov-
enance. Some units at the type locality of the Harding
Sandstone contain both invertebrate and vertebrate
fossils. It is likely these represent intertidal units that
have received vertebrate material from stream effluent.
Such was the conclusion of Romer and Grove (1935)
but these authors had no evidence that fluvial channels
existed in the Harding Sandstone.
The possibility remains that the vertebrates could
have migrated between fresh water and the sea (diad-
romy) (Griffith, 1987). Theoretically, if one could find
articulated fish skeletons in units that contained abun-
dant marine invertebrates and in freshwater deposits,
diadromy might be a parsimonious conclusion. How-
ever, no such conditions occur at the Bushnell Lakes
and given the distribution of vertebrate fossils there it
is less conservative to assume that they represent diad-
romous animals. More work should be undertaken at
other Ordovician localities to determine whether the
fossil record can help to support the work of Griffith
(1987).
What light does this information shed on the study
of the earliest vertebrate environment? It is safe to say
that the currently held view among most paleontolo-
gists, and in recent textbooks, is that the original ver-
tebrate was a marine animal (McFarland et al., 1979;
Darby, 1982; Boucot and Janis, 1983; Jeffries, 1986;
Keeton and Gould, 1986; Carroll, 1988; Steam and
Carroll, 1989). It is also safe to say that, until the
taxonomic status of the pre Middle Ordovician prob-
lematic fossils (see above) is resolved, the Harding
Sandstone and the Stairway Sandstone (Middle Or-
dovician of Australia; Ritchie and Gilbert-Tomlinson,
1977) contain the oldest uncontested vertebrates (El-
liott et al., 1991). Based on this study it is conceivable
that the oldest North American vertebrates lived in
freshwater.
Many other localities of Ordovician vertebrate fau-
nas similar to the Harding Sandstone have been re-
ported in North America. They show that vertebrate
fragments occur in carbonate rocks (Eliuk, 1973; Leh-
tola, 1973) as well as in detrital sediments (Orvig, 1958).
Also, articulated Ordovician vertebrates have been
found at sites in Australia (Bockelie and Fortey, 1976)
and South America (Gagnier et al., 1986). These im-
portant localities have not been studied paleoenviron-
mentally. Before we can understand the large-scale
habitation patterns of the earliest vertebrates, paleoen-
vironmental analyses of Ordovician vertebrate local-
ities must be made.
9
ACKNOWLEDGMENTS
I thank P. P. Vaughn and W. E. Reed for advice,
direction, and feedback throughout this project. For
discussions on various topics relevant to this study I
thank D. K. Elliott, E. C. Olson, A. R. McCune, R. V.
Ingersoll, and G. Oertel. Fieldwork was supported by
grants from Sigma Xi, the Society of Vertebrate Pa-
leontology, and the Department of Earth and Space
Sciences at UCLA. Preparation of the manuscript was
supported, in part, by NSF grant #BSR-707500 to A.
R. McCune.
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Received 7 December 1990; accepted 2 April 1991.