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 Stable URL: http://www.jstor.org/stable/4523416 Accessed: 02/10/2010 12:43
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indicating that the remains must have been washed into the sea from streams and rivers. Although a stratigraphic description of the Harding Sandstone was published when it was named (Walcott 1892). abandoned delta lobes and delta front that are recognizable as different facies. most types of modern depositional environments have been documented and are characterized by particular assemblages of sedimentary structures. the collection of data at or near only one outcrop. It is clear that within one depositional system many types of organisms. METHODOLOGY A pervasive methodology in paleontology is the determination of past environments based on fossil associations. This. overbank areas. sometimes correlatable over large geographic distances. Colorado.S. 1. Sweet. It is not uncommon to find references to associations of fossils from an entire formation. may be preserved. where mixed assemblages did occur. 12. Presently. and later studies revealed its widespread occurrence (Behre and Johnson. and. Based primarily on the studies of Denison and Spjeldnaes. This has led to the erection of sedimentary facies models (Walker. It is possible to get a rough idea of the biotic relationships of a fossil species in such instances. This was based on invertebrate associations at the type locality and also on boron analysis of certain clays in the sediments. called "Walther's Law. which implied that the two could not have been deposited simultaneously.. primarily. The basis for the concept is the fact that certain fossil taxa consistently co-occur in strata.2 JOURNAL OF VERTEBRATE PALEONTOLOGY. Denison (1956) cited a general lack of positive evidence supporting the freshwater theory and maintained that freshwater deposits were lacking at the oldest known vertebrate localities. the type section. is usually composed of multiple strata deposited in various depositional environments. such as a prograding delta. Unfortunately. From evidence at the type locality. Colorado along U. VOL. A given depositional system. Also. The Bushnell Lakes . 1933. the type locality near Canon City. he was convinced that the Harding Sandstone was deposited in the sea and that the widespread vertebrate fragments could not have been introduced by fluvial processes. NO. Highway 50). a previously undescribed locality about 50 miles west of Canon City that permits a complete facies analysis. all of which may contain fossils. It is very difficult to find a complete stratigraphic section. distal channels. 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. can be superimposed. even though the organisms were living in completely different environments. he stressed that the bone fragments were much larger than the largest sand grains. This scenario was supported by osmoregulatory physiological studies of Smith (1932). 1). Johannes Walther stated that "only those facies . from vastly different habitats. is a quarry that has been worked for nearly 100 years. a deltaic system may bury terrestrial vertebrates in proximal areas and marine fishes in distal channels. that can be observed beside each other at the present time" (Middleton. Mouths of streams and rivers are locally restricted. and many of the fossiliferous strata have been obliterated. In stratigraphic sections such disparate forms may appear within the same formation separated vertically by only a short interval. There is. he maintained. It is rooted in principles put forth in past centuries. The only paleoenvironmentally mean- ingful associations thus are those from the same depositional environment. Spjeldnaes (1967) attempted a paleoecological analysis of the Harding Sandstone and concluded the vertebrate-bearing units were deposited in a restricted marine environment. second. where most previous paleontological work has been done. but it is not necessarily true that the organisms lived together. 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. 1967). including proximal channels. Sedimentary facies are defined based on such structures. the present study is the first published facies analysis of this important formation. It is located in San Isabel National Forest. However. about seven statute miles (11 km) southwest of Howard (which is about 18 miles (29 km) southeast of Salida. however. In 1894. 1984)." allows us to interpret a sequence of strata as originally juxtaposed environments of deposition. is composed of many environments. and yet the bone fragments of the Harding Sandstone are found over a geographic expanse that stretches from central Colorado to northern Wyoming. it became generally agreed among paleontologists that the vertebrates from the Harding Sandstone were marine animals. 1984) that provide the foundation upon which sedimentological interpretations are built. 1992 Sandstone. A single formation. the reliance on paleontological associations without a clear knowledge of the facies relationships. in Fremont County (Fig. however. 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. The documentation of modern sedimentary processes constitutes the most important step in the development of the "modern revolution in stratigraphy" (Miall. The "ostracoderms" must have fallen onto the sandy sea substratum when they died. 1954. first. Gerhard. Without facies analysis this situation could lead to an incorrect assessment of biotic relationships. The fossils might be considered in association. For instance.. the vertebrate fossils were always disarticulated and worn. 1973). A methodology for determining depositional environments has been developing over the past 25 years.

Colorado is 36 miles (58 km) east along US Highway 50.6 km) Hayden Pass FIGURE 1. Scale: 1 MilePeak (1. The locality is accessible by a marked trail from the Hayden Creek campground along county road 249 (lower right of map). and Salida.GRAFFIN-HARDING SANDSTONE 3 VICINITY MAP DENVER Glenwood Springs 1O Us 6 A0pen Colorado BUSHNELL Springs LAKES REGION SSa Walsenburg Arkansas Alamos " C mpgrou San Isabel ( ( Forest GNational 1Mile Scale: - (fN Q0Sn COALDALE O Hayden Creek. Colorado is 18 miles (29 km) northwest of Howard along Highway 50. Canon City. . The region is within San Isabel National Forest. Location map of the Bushnell Lakes.

1986).5 to 4 meters thick and can be traced laterally for many tens of meters. Because this is above treeline. Extreme bioturbation tends to erase all sedimentary features. 1958). Three lakes occur in series at progressively lower elevations (pater noster lakes). The contact between the Manitou Formation and the Harding Sandstone has been defined as an unconformity (Litsey. At Bushnell Lakes. The discordances in the beds reflect changes in the inclination of the beach face toward the sea. which is found in all three formations. 1986). the Middle Ordovician Harding Formation (=Harding Sandstone). fine and medium-grained arenites. 3) very low-angle wedge-set cross-bedding in fine to medium grained units. it is necessary to view it in relation to the sedimentary system that operated during the formation of all three Ordovician units. They consist of sandy laminae with slightly discordant contacts. Thus determination of shoreface deposits is based primarily upon churned bedding and secondarily on the position within a facies sequence. it is often difficult to find any distinctive sedimentary structures preserved in the shoreface zone (Elliott. Units that show laminations interrupted by bioturbation characterize the offshoreshoreface transition zone. below the critical velocity of ripple formation. VOL. Ordovician sedimentary rocks form a conformable sequence about 110 meters thick. Upon handlens inspection. and highly fossiliferous pebble conglomerates. The entire Ordovician sequence lies nonconformably upon Precambrian gneiss. usually about 5 to 10 millimeters thick. areas not affected by the biological disruption (Conybeare and Crook. All of its qualities render it an ideal locality for a facies analysis. The sequence is divided into three units. Churned bedding often results and is recognizable by the presence of mottling in the sediments and also by remnant patches of parallel laminations. Due to the high degree of bioturbation. near the crest of the Sangre De Cristo range. 12. the sequence appears to be conformable from the base of the Manitou Formation to the very top of the Fremont Formation. 1992 basin is a glacially carved valley. 2). They are commonly faint. and quartz grains make up about 60 percent. 1. Very low-angle cross-beds (wedge sets) are found in all three formations at Bushnell Lakes. very little vegetation is present. The former contains some quartzitic units. The quartz percentage increases upward in the section until. This results from differential deposition/erosion rates at different times. Theoretically. however. This requires a facies analysis of the entire Ordovician sequence. the Lower Ordovician Manitou Formation. this ratio could change due to seasonality (stormy and calm seasons) or changing climate over time. Laminations occur throughout the Manitou Formation and in the upper part of the Fremont Formation. Laminae can be traced laterally many tens of meters before becoming truncated or obscured. however. and 4) epsilon and trough cross-beds. They are invariably overlain or underlain by units showing bioturbation. they are seen to be composed of minor grain size differences between layers. NO. because organisms thoroughly disrupt sediments before lithification. the lithoclastic quartz content of the carbonate rock is never more than seven percent. Epsilon cross-beds are characterized by inclined sets of lentic- . 1986). 2) burrows and other evidence of bioturbation. 1968). Near the base of the Manitou Formation. Both the Manitou and Fremont formations are primarily composed of dolomite. laminae are associated with evidence of biological sediment disruption. at about five meters below the contact with the Harding Sandstone.4 JOURNAL OF VERTEBRATE PALEONTOLOGY. The fourth type of sedimentary structure found in the Ordovician rocks at Bushnell Lakes comprises epsilon and trough cross-beds. can be used to define the different types of facies that occur at Bushnell Lakes (Fig. and associated sedimentological features. Often this zone grades upward into another laminated unit. 1986). neither burrows nor churned bedding occurs in units with low-angle wedge-set cross-beds.200 feet (3. and are present in both micritic and sparitic units. Because the Harding Sandstone is thus a part of a depositional continuum. Units with laminations grade upward into zones with worm burrows and other evidence of bioturbation. generally devoid of vegetation.405 meters). 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. Burrows are sometimes preserved within mottled beds in the Manitou Formation. Units with these characteristics are interpreted as subtidal deposits that were laid down below wave base. very fine-grained wackes. The laminations are caused by plane-bed deposition during the swash and backwash of wave action (Elliott. The units that show uninterrupted laminations that extend laterally for 10 to 100 meters are interpreted as deposits of the offshore zone (Elliott. Bed sets are 0. They are found only in the coarsest units of the Harding Sandstone. These structures are only found in units with fine or medium granularity that generally lack mud. These. Its elevation is at about 11. FACIES TYPES There are four basic types of sedimentary structures preserved in the Ordovician rocks: 1) parallel laminations in fine-grained units. and the Upper Ordovician Fremont Formation. Wedge-sets are very distinctive sedimentary structures and characterize deposits of the upper foreshore or beach (Reineck and Singh. The scale of bedding is about five to ten centimeters. The Harding Sandstone is comprised entirely of detrital clastic rocks such as mudstones. This provides excellent exposure of the strata and subtle sedimentary structures. the carbonate content of the rock is only about 40 percent. Almost invariably.

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is overlain by an intertidal foreshore (beach) deposit. Invertebrate body fossils are absent from the Harding Sandstone. It is within these units that the greatest concentration of fossil vertebrate material is found. Such "faunal mixing" has played an important role in bolstering the view that the vertebrates of the Harding Sandstone were marine animals. Trough (or festoon) cross-beds are broadly concave and also form erosive bases. vertebrate fossils do not occur in such units. 1986) but may not always be present in a fluvial sequence. This is normal to the direction of current flow. however. 1956. and it defines the zone of clastic influx in such a model (Wilson. 1968). Johnson. 1967). after death. 1973). 1954. Where present. quartz influx increased. FOSSIL DISTRIBUTION As reported at other localities (Johnson. Often the two types are found within the same sand body. 1992 ular sandy beds that form erosive bases. 1967). solitary rugose corals. similar to that described at other localities (Sweet. subtidal deposit characterized by laminations. which. The dip indicates the direction of migration of the deposit through time. 1956. crinoids. it is most parsimonious to assume that they were freshwater animals that lived in rivers and streams. Furthermore. Much of the Harding Sandstone is marine as suggested by past authors (Denison. These occur from the base to five meters above the upper contact of the Harding Sandstone. These rocks are composed almost exclusively of vertebrate fragments and rounded quartz granules (Fig. FACIES SEQUENCE Facies succession at Bushnell Lakes is cyclic. but the Bushnell Lakes sequence shows that the model can be consistent with a mixed carbonate and clastic succession as well. Detrital quartz began to enter the system in small quantities during deposition of the Manitou Formation. They are rather restricted in distribution at the Bushnell Lakes. 1986). The Fremont Formation is purely carbonate and was laid down in successively deeper water. 1. the only occurrence of such in the entire Ordovician sequence. they are found at the base of. Although bioturbated sediments within the Harding Sandstone provide evidence that invertebrates were present. conglomerates are key indicators of fluvial channel deposits (Collinson. Presumably. 1984). It was deposited atop a gently sloping carbonate ramp (sensu Ahr. in turn. Thus a generalized unit sequence. Gradually. NO. They do not indicate a laterally migrating stream but represent channel scours that have become infilled with coarse sediment (Conybeare and Crook. 1945). Spjeldnaes. or facies model. At the Bushnell Lakes. But its upper portions represent supratidal terrestrial deposition. 3) that shows such an upward-shallowing trend. In general. 5). Intertidal units of the Harding Sandstone contain occasional rare vertebrate fossils. Rarely. The nature of streamflow causes lateral migration of fluvial channels and associated deposits. culminating in progradation of terrestrial environments over the intertidal. sandy units that represent higher point bar deposits. the Manitou Formation at Bushnell Lakes contains poorly preserved invertebrate fossils that are few in number. After deposition of the Harding Sandstone there ceased to be any quartz introduced. Overlying that is a bioturbated shoreface deposit. VOL. Rather. The data at hand suggest that there is no basis for assuming the vertebrates of the Harding Sandstone inhabited the sea. Vertebrate and invertebrate fossils have been reported from the same strata at the type locality of the Harding Sandstone (Walcott. 1945). The abundance of fossils in the Fremont Formation is not great. Virtually all are crinoid fragments that occur in the offshore facies within five meters of the contact with the Harding Sandstone. 12. Denison. Epsilon cross-beds (Allen. The smallest troughs are only about 10 cm in depth and about 50 cm wide. carbonate-dominated marine system. They are commonly found on point bars or as individual channels within a braided stream system. 1963) are features formed by lateral accumulation of sediment. 1975). The trough cross-beds are also interpreted as channels. Invertebrate fossils are more abundant and varied in the Fremont Formation than in the Manitou Formation. Spjeldnaes. The facies relationships suggest that the Harding Sandstone represents a terrigenous clastic phase of deposition within a shallowing-upward. and coralline algae. The coarse conglomerates are interpreted as channel "lag" deposits of the lower point bar and stream bed (Reineck and Singh. the elements forming their exoskeletons (tesserae) became disconnected and were incorporated into the bedload of streams. 4) where they occur in coarse conglomeratic channel deposits. The lowest unit is an offshore. This model was strictly for carbonates. and grade into. This tendency causes erosion of underlying sediments and is seen as scoured bedding surfaces in stratigraphic sections. there are no invertebrate fossils found in any of the units that contain abundant vertebrate fossils.6 JOURNAL OF VERTEBRATE PALEONTOLOGY. 1986). . the greatest accumulations of vertebrate fossils are restricted to the supratidal facies (Fig. within the shoreface facies. 1892. The scale of bedding is variable but the largest individual epsilon cross-beds are about one meter deep. this is overlain by supratidal fluvial and related facies. Each cycle consists of a general upward pattern of shallowing deposits. A similar facies model was erected as a generalization for many Paleozoic carbonate sequences (James. Some of the epsilon and trough cross-bedded units contain pebble conglomerates that extend laterally for less than 10 meters and are about one meter thick. They are a superb guide to fluvial facies because each individual bedding surface represents a point bar from a meandering stream (Collinson. The remains comprise fragments of brachiopods. can be discerned (Fig.

. Arguments that the vertebrates of the Harding Sandstone were marine require one of two unlikely conditions: 1) the vertebrate fragments were washed upstream from the mouths of streams and came to rest in fluvial environments. Although vertebrates are present in some of the intertidalunits. It implies either that vertebrates and invertebrates lived together and were separated after death or that the two groups never lived together and were preserved separately.Similar sedimentarystructuresmay be found in both carbonateand clastic deposits. The fossiliferousunits are indicated in Figure 2. 2). Histogramof fossil types and distributionsin the fossiliferous Ordovician sediments at Bushnell Lakes. or 2) the fluvial channels rec- S40 II Vertebrates 20 subtidal intertidal fluvial Depositional enviroment 60 40 ~ Inverteb rates 20 0subtidal intertidal fluvial Depositional enviroment FIGURE 4. suggestingthat they lived in fresh water and that some remains probably got washed in to the nearshoreenvironment. Offshore- Shoreface Transition model in a mixed carbonate-clastic FIGURE 3. Partiallyadaptedfrom James (1984).A completesequenceof shoalingdeposits includes offshore-shoreface foreshore. Shallowing-upward setting. in different environments of deposition. Although it is difficult to prove. only one includes all four types of deposits. the fact that vertebrate fossils are most abundant in the granular conglomeratic inferred fluvial facies (units that lack any invertebrate fossils) strongly suggests that the latter scenario is most parsimonious.GRAFFIN-HARDING SANDSTONE SEDIMENTARY STRUCTURES 7 MODEL SHALLOWING . wedge sets and rippled Low-angle Intertidal calcareniteor fine to medium sandstone. or Laminated micritic carbonate or very fine sandstone.and fluvial sediments. Bioturbated (churnedbedding) Foreshore Sh f Subtidal micriticcarbonate fine sandstone.Thereare five shoalingcycles in the Ordovicianrocks transition. at Bushnell Lakes (see Fig.shoreface. The fluvial facies are only found in the Harding Sandstone.UPWARD Terrestrial Supratidal INTERPRETATION Epsilonand festoon cross-bedswith granular conglomeratechannel lags (highly fossiliferous) interbedded Progradational Fluvial System with laminatedand rippledmudstone. they are much more abundantin fluvial facies. The depositional environments were determined by facies analysis. 60- DISCUSSION The fact that vertebrate and invertebrate fossils generally do not occur together at Bushnell Lakes is paleoenvironmentally significant.

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1956.Colorado. Manual of Sedimentary Structures. and feedback throughout this project. R. R. J. A. Gagnier. ACKNOWLEDGMENTS I thank P. Allen.Science237:190192. V.in C. K. Before we can understand the large-scale habitation patterns of the earliest vertebrates. 1987. Barnes (ed. and C. Boucot and Janis. S. L. Darby. direction. 1986. and G.. As mentioned above. A. Siliciclasticshorelines.pp. Geology 16:131-192. Based on this study it is conceivable that the oldest North American vertebrates lived in freshwater. These important localities have not been studied paleoenvironmentally. Keeton and Gould.. C. is that the original vertebrate was a marine animal (McFarland et al. St. 698 pp. 1982. E.-Y. B. . Darby. Blackwell. diadromy might be a parsimonious conclusion. Elliott..). Ordovician vertebrates. 1977) contain the oldest uncontested vertebrates (Elliott et al. Reading (ed. there are certain intertidal units within the Harding Sandstone that contain small amounts of bone (less than 7% of the total rock volume). . Newfoundland. R. Griffith. Sedimentology 2:93114. A. R. Bureau of Mineral Resources. G. Preparation of the manuscript was supported. Journal of Paleontology 56:1187-1196. 1973) as well as in detrital sediments (Orvig. Reed for advice. 1973. Blackwell. 1933. Denison. Geobios 19:629-634. and J. L. New York. Also. LITERATURE CITED Ahr. Middle Ordovician fish-bearingbeds from the St. 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 diadromous animals.Fieldiana. 327 pp.HARDING SANDSTONE 9 vertebrate fragments in the supratidal deposits. 1967.). L.L. Blieck. Fieldiana. Some units at the type locality of the Harding Sandstone contain both invertebrate and vertebrate fossils. These fragments were probably washed into the intertidal zone from the stream channels. G. T. Paleoecology 41:251-287. 1986. 1988. the Society of Vertebrate Paleontology. John's.. and absence in subtidal deposits is suggestive of a freshwater habitation. R. 1973. The classification of cross-stratified units. Ritchie and Gilbert-Tomlinson. D.. 1982. Steam and Carroll. Alluvial sediments.R. Geology and Geophysics. J. 1988. 1976) and South America (Gagnier et al. Ordovicianvertebratesfrom the westernUnited States. the oldest North AmericanVertebrate. D. However. Paleozoic geologic development of Canon City embayment. 1991). VertebratePaleontology and Evolution.Oxford. Carroll. 1986. Boucot. AmericanJournal of Science 5:477-486. 1986). The possibility remains that the vertebrates could have migrated between fresh water and the sea (diadromy) (Griffith. Freshwateror marine origin of the ver- . 1968. 1991. 1967.GRAFFIN. 1963. It is likely these represent intertidal units that have received vertebrate material from stream effluent. until the taxonomic status of the pre Middle Ordovician problematic fossils (see above) is resolved. 1958). Oertel.Paleoclimatology. 155-188 in H. Olson. and K.Oxford. Geology 11:359-457. K. 1976. habitat based on a lithologic association. and in recent textbooks. 20-62 in H. R. in part. Gerhard. S. A reassessmentof Astraspisdesiderata. P-Y. by NSF grant #BSR-707500 to A. Many other localities of Ordovician vertebrate faunas similar to the Harding Sandstone have been reported in North America. 1986. For discussions on various topics relevant to this study I thank D. 1973. 1987. Sedimentary Environments and Facies. More work should be undertaken at other Ordovician localities to determine whether the fossil record can help to support the work of Griffith (1987). Environmentof the early Paleozoic vertebrates. D. The carbonateramp: an alternative to the shelf model. 1983. Sedimentary Environments and Facies. First Ordovician vertebratefrom South America. the Harding Sandstone and the Stairway Sandstone (Middle Ordovician of Australia. H. Canadian Journalof EarthScience 10:954-960. LawrenceLowlands of Quebec. 1986. Eliuk. Both bone and quartz grains likely had a continental provenance. A. Reading (ed. Behre. Fieldwork was supported by grants from Sigma Xi. Ingersoll. T. Blieck. and P. 1979. paleoenvironmental analyses of Ordovician vertebrate localities must be made. Such was the conclusion of Romer and Grove (1935) but these authors had no evidence that fluvial channels existed in the Harding Sandstone. Conybeare. Fortey. J. Johnson. 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 paleontologists. Gagnier. Theoretically.pp. A. Department of National Development. Carroll. H. 1989). An early Ordovician vertebrate. Lehtola. 1983. 5th InternationalSymposium on the Ordovician System. if one could find articulated fish skeletons in units that contained abundant marine invertebrates and in freshwater deposits..Nature 260:36-38. Ordovician and Devonian fish horizons in Colorado. E.). E. their paucity in intertidal. They show that vertebrate fragments occur in carbonate rocks (Eliuk.Bulletinof the American Association of Petroleum Geologists 51:2260-2280. Transactionsof the Gulf Coast Association Geological Society 23:221-225. Paleogeography. and G. Janis. Elliott. Freeman and Company. R. M. with notes on their origin. The early vertebrate Astraspis. Jeffries. articulated Ordovician vertebrates have been found at sites in Australia (Bockelie and Fortey.. somewhat analogous to the way the quartz grains of the Manitou Formation were washed into the intertidal zones. P. W. Rodrigo. It is also safe to say that. Collinson. Crook. McCune. H. and R. 1987). and the Department of Earth and Space Sciences at UCLA. Bockelie. A review of the habitatof the earliest vertebrates. Elliott.C. Vaughn and W. C. McCune..

Middleton. Science 200:529-532. H. N. Quarterly of the Colorado School of Mines 40:1-109. V. J. (ed. W.-E. 1984. Facies Models. Lehtola. Quarterly Review of Biology 7:1-26. new species. Wiley. and J. Received 7 December 1990. 1984. K. Water regulation and its evolution in the fishes. T. British Museum (Natural History). 1986. Walker. Springer-Verlag. N. Gilbert-Tomlinson. 1984. R. A. G. Johannes Walther's law of the correlation of facies. 1973.. and J. J. pp. and R. R. Rapport Gronlands Geologiske Undersogelse 91:111-115. U. L. T. Macmillan.10 JOURNAL OF VERTEBRATE PALEONTOLOGY. Walker. Vertebrate Life. The Ancestry of the Vertebrates. D. 531 pp. Litsey. Bulletin of the Geological Society of America 3:153-172. 213-228 in R. the Record of Life. Carroll. Colorado. Environment of the earliest vertebrates. Shallowing-upward sequences in carbonates. University of Michigan 24:23-30. 1979. S.S. 1958. G. London. 1892. Walcott. L. New York. Norton. and J. 1954. a new ostracoderm from the upper Ordovician of North America. Bulletin of the American Association of Petroleum Geologists 38:284-305. Bulletin of the Geological Society of America 84:979-988. 376 pp. The palaeoecology of the Ordovician vertebrates of the Harding Formation (Colorado. 1977. Proceedings of the United States National Museum 108:1-23. Steam. Johnson. Peel. First Ordovician vertebrates from the Southern Hemisphere. Carbonate Facies in Geologic History. H. L. 1945. Orvig. New York. . New York. Stratigraphy and structure of the northern Sangre De Cristo mountains. Geosciences Canada. 490 pp. McFarland. Springer-Verlag. Romer. B. A. 453 pp. Facies Models. L.).. 1958. J. Harding and Fremont Formations. James. Biological Science. S. new genus. F. N. R. American Midland Naturalist 16: 805-856. 12. A resume of the paleozoic stratigraphy of Colorado. Paleontology. P. E. H. J. Contributions from the Museum of Paleontology. 1992 Reineck. Anatolepis from the Early Ordovician of East Greenland-not a fishy tail. 317 pp. Principles of Sedimentary Basin Analysis.. 1978. Pycnaspis splendens. accepted 2 April 1991. 1935. Alcheringa 1:351-368. Grove. Depositional Environments and Facies. A. S. 471 pp. Springer-Verlag. VOL. T.. 1989. 875 pp. W. Sweet.A. 1975. Wilson. W. New York. Singh. H. C. B. Heiser. NO.). Colorado. Keeton. 1979. and I. Spjeldnaes. W. Repetski. J. 1. and B. 1973. J. Gould. Collogues Internationaux du Centre National de la Recherche Scientifique 163:11-20. Smith. Ritchie. Bulletin of the Geological Society of America 69:1143-1178. A fish from the Upper Cambrian of North America. A. 1266 pp. W. Reprint Series 1. 1967. C. G. 1986. Geosciences Canada. Miall. 1932. Ordovician vertebrates from Ontario. 1986. Notes on the discovery of a vertebrate fauna in Silurian (Ordovician) strata. New York. New York. Pough.. Cade. tebrates? Comparative Biochemical Physiology 87A:523531. Jeffries.