Paleoecology is the study of ancient biotic communities and their relationship to the abiotic environment. The conclusions of paleoecology are reached by studying in detail the fossils and sediments and whatever relationships exist between them. Its ultimate aim is to build a picture of the climate, flora, fauna, and topographic setting of an area. For this reason it borrows heavily from paleontology, paleobotany, sedimentology, and climatology.
From studying modern environments, it is observed that each has its own type of sediment. When certain sediments are found in the rock record, it is generally assumed that they represent environments similar to modern ones that produce similar deposits. For example, limestone is forming today in warm, shallow, well-lighted, well-aerated water. When limestone is found in a rock sequence, it is usually assumed that the same or similar environmental conditions occurred in that area in the past. Fossils can further refine the interpretation.
As can be interpreted from foregoing discussions, the sediments deposited in the Fossil Basin vary from stream and flood-plain fluvial to lacustrine. At times, material eroded from surrounding uplands was carried by streams throughout the basin. These rocks are exemplified by the Evanston and parts of the Wasatch and Fowkes formations. When the lake appeared, lacustrine sediments, marlstone, and shale were deposited in the lake, while around the periphery of the basin, fluvial sediments continued to accumulate.
Plants found in the lake sediments of Fossil Basin tell us much about the climatic conditions that prevailed during the Eocene. The flora is quite similar to that now existing in the southeastern United States, reflecting a warm and humid climate. One of the striking examples is that of huge palm fronds that occasionally are found in the fish quarries at Fossil Butte. These seem to be fronds that blew from the trees into the lake. Soon they became water-logged and settled to the bottom to be preserved.
The land animals indicate that a rather wide range of ecologic niches existed over the basin before the lake came into being, around the lake during its presence, and again all over the basin after the lake disappeared. Many of the smaller mammals, some of the rodents, and most of the primates were almost certainly arboreal. The large mammals such as uintatheres, pantodonts, and tapiroids may have been stream-side or marsh dwellers. Probably inhabiting the forest floor and feeding on low bushes and undergrowth were such forms as the condylarths, horses, artiodactyls, and some of the rodents. Feeding on the rodents and other smaller mammals were the creodonts and miacids. Tiny shrew-like forms scampered about the undergrowth and fed upon worms and insects.
Bradley (1963) has estimated possible temperature and precipitation levels in southwestern Wyoming during the Eocene. Bradley’s modern analogues were the Gulf Coast and Great Lakes regions. From a series of calculations, an average annual temperature of 65°F is postulated. This could have fluctuated greatly in the inland setting of Fossil Basin. Precipitation amounts were possibly on the order of 30-43 inches annually. The amount of annual evaporation was also possibly in the range of 30-43 inches.
It is believed that Fossil Lake was thermally stratified—that is, with colder, denser water at depth (hypolimnion) and warmer, less dense water (epilimnion) nearer the surface. The deeper waters probably would have been devoid of oxygen, hence essentially uninhabited other than by anaerobic bacteria that survive without oxygen. If the bottom had been oxygenated, many types of life would have burrowed into the sediment thus destroying the delicate varves (Fig. 28). The lake was probably deep enough that wind and wave action did not roil the bottom sediments.
Fig. 28. A section of shales from the fish beds showing thin laminae that are interpreted as varves. Enlarged six times.
There have been many attempts to interpret the taphonomy (see glossary) of concentrations of fish at Fossil Butte. Bradley (1948) interpreted the cause of death and the reason for their preservation as follows:
In this basin (Fossil, Wyo.) hundreds of thousands of beautifully preserved fish are entombed in the varved sediments. Even the delicate fin and tail rays and other bones originally held in place only by tissue are virtually undisturbed, and even the scales are in place almost completely undisturbed. It seems to me that the picture of this lake as a thermally stratified water body provides nearly all the necessary information to account for the excellent preservation of these fish. Only in the stagnant hypolimnion could they have escaped being torn to pieces by scavengers or distorted by bottom feeders. It is significant that all the well preserved fish are in varved sediments. Those in non-varved sediments are a disordered mass of broken and chewed up bones.
The only part of the story lacking now is how the fish died and got into the hypolimnion. Limnology offers two possible explanations. Sometimes when the surface of a lake gets excessively warm, fish will plunge into deep water and might thus penetrate the hypolimnion, be overcome by hydrogen sulphide, and also have the gas in their swim bladders chilled so that they sank at once to the bottom. Once there, only anaerobic bacteria would attack them. The other hypothesis is that the thermally stratified lake was suddenly chilled so that it overturned more rapidly than the hydrogen sulphide could be oxydized and so killed off large numbers of fish. This seems a little more probable as the fossil fish are of all ages and sizes.
The fish in these quarries have been collected since the 1870s but mainly by commercial collectors. Most museum collections were purchased from these commercial collectors, hence they consist almost entirely of perfectly preserved fish, as poorly preserved specimens would be discarded by the collectors as of no monetary value. The result has been that most people, including Bradley, were misled into believing all of the fish in the varved sediments of the quarry were perfectly preserved. As nearly as can be determined, the first attempt to systematically collect and study this concentration of fish was that by paleontologists from the University of Wyoming (McGrew 1974). This work threw much new light on the occurrence and made available much new data that permit new interpretations.
Shales of the Green River Formation in general have been described by Bradley (1931) and he specifically mentioned those of the fish layer in the Fossil Basin as follows: “Plate 1 shows a thin section of the varved marlstone in the small, unnamed Green River Lake west of Gosiute Lake, where the varves are better developed. Each varve or annual deposit, consists of a layer of microgranular carbonates and a thinner, dark layer of organic matter” (Bradley 1948:645). The X-ray diffraction analyses performed by John Ward Smith of the Laramie Energy Research Center showed that the shales of the “fish layer” consist predominantly of calcite (roughly 60%), aragonite and dolomite (approximately 20%), and quartz (up to 10%).
Although fish are numerous throughout the thickness of this “fish layer,” there are three laminae that contain so many fish that it is almost certain that they represent catastrophic mass mortalities. Two are made up primarily of Priscacara and one consists almost exclusively of Knightia.
X-ray photos show that a rather high percentage of the fish in the shales are not perfectly preserved but have undergone varying amounts of disarticulation. There appears to be an orderly sequence of stages of decomposition—from essentially perfect articulation to total disarticulation. Disarticulation first appears in the most anterior vertebrae. From there it rapidly proceeds anteriorly into the head region and appears to do so posteriorly at a slower rate. In many specimens the head and anterior half of the body are completely disarticulated, while the posterior part of the body shows no disarticulation whatever.
It is assumed that after the dead fish settled to the bottom of the lake, external anaerobic bacteria found access to the “innards” of the fish via the opercular opening. This would account for the first sign of decomposition and disarticulation being just back of the head.
In the blocks of shale covered by X-ray there were 385 fish. For convenience, these were classified into six groups, group I showing no discernable disarticulation and group VI (Fig. 29) showing total disarticulation. The number and percentage of fish in each group are as follows:
| Group I | 223 fish | 58% |
| Group II | 38 fish | 10% |
| Group III | 14 fish | 4% |
| Group IV | 27 fish | 7% |
| Group V | 24 fish | 6% |
| Group VI | 59 fish | 15% |
Fig. 29. A partially disarticulated skeleton of a large Priscacara, Group IV.
Because of the predominance of completely articulated fish throughout the quarry and the fact that the fish involved in the mass mortalities show no disarticulation, it seems probable that some connection might exist between the death of the fish and conditions of the lake bottom that would cause their perfect preservation. It would seem that rapid burial might be the most obvious reason for excellent preservation. Thus any factor or combination of factors that would cause rapid precipitation of carbonates and also would cause mortality of fish would satisfy our requirements. One obvious factor that could, at least theoretically, fulfill these requirements would be an annual bloom of blue-green algae that are known to be toxic to fish (Prescott 1948). Such blooms usually occur during the warmest part of the summer when CaCO₃ is least soluble. By extracting CO₂ from the water, these algae are known to cause precipitation of CaCO₃. Thus we have a possible cause for some annual fish kill and perhaps an occasional superbloom that would bring about a catastrophic mass mortality. The highest mortality of fish then might have occurred during late summer algal blooms and at this time also would occur the most rapid precipitation of CaCO₃.
It is not known how much deposition of CaCO₃ would be required to protect a fish from disarticulation. It may be that a very thin layer, especially if mixed with organic ooze, might provide an effective seal to slow decomposition and prevent disarticulation. If sufficient CaCO₃ was precipitated and deposited to cover the fish that died during this period, one might expect perfect preservation. Such fish would fit into group I. Those fish that died just after this period might lie exposed on the lake bottom for most of a year and be subject to disarticulation. If little or no deposition took place during the rest of the year, fish that died just after the period of deposition should be the most completely disarticulated and fit into group VI. During the fall, winter, and spring, fish that died of attritional mortality would be disarticulated according to the length of time they lay on the bottom prior to the next period of deposition.
If the foregoing is true, one should be able to determine the approximate time of year fish died by the degree of disarticulation. One might assume that blooms of blue-green algae and hence precipitation of CaCO₃ would take place sometime during August and/or September. Thus we should expect the most fish and those most perfectly preserved to have died during this time period (group I). Those fish that died in October and/or November should be the most completely disarticulated (group VI), and those that died in June or July should show only a slight degree of disarticulation (group II). The distribution of the stages of disarticulation seems to fit almost exactly the pattern that one would expect if this interpretation is correct.
Gunter (1947) has shown that annual periods of excess salinity in Texas lagoons cause an annual increase in the death of fish and occasionally a catastrophic mass mortality. Because Lake Gosiute is known to have been saline, it might be assumed that somewhat similar chemical conditions prevailed in the Fossil Lake. It is not known definitely that the two lakes were ever connected but if they were, it was most probably a narrow connection near the southern end of Fossil Lake and probably a rather temporary connection.
That a rather long period of aridity occurred in the general region is demonstrated by various depositional features, primary structures, and salt deposition in Gosiute Lake in a part of the Green River Formation. These deposits appear to have settled down during Lostcabinian (late to early Eocene) times. Because the Wasatch Formation immediately underlying the Fossil Butte Member of the Green River Formation in the Fossil Syncline Basin is Lysitean (mid to early Eocene), it is probable that the fish deposits are of Lostcabinian age. Thus it may well be that the long period of aridity occurred during the deposition of the fish beds. While the Fossil Lake probably never reached the high degree of salinity present in Lake Gosiute, it was probably sufficiently saline that periods of excessive evaporation could increase the salinity enough to contribute to the mortality of fish and occasionally cause a catastrophic mass mortality such as those described by Gunter. The presence of aragonite and dolomite in the shales suggests that at least some of the carbonate deposition might well have been because of excessive evaporation and high concentrations of carbonates (Smith 1974, pers. comm.).
At the University of Wyoming, an attempt is being made to interpret scales of fish from the quarries. In a number of specimens annuli can be observed and circuli counted. Although removal of scales for study is extremely difficult, a number has been removed and photographed with a scanning electron microscope. In Lake George, Florida, black crappie develop annuli during January, February, March, and April (Huish 1954). Climatic conditions in north central Florida may be similar to those that existed in western Wyoming during the early Eocene. Thus annuli may have developed at the same time of year. By counting the number of circuli between the last annulus and the edge of the scale, it should be possible to determine the approximate time of death of a fossil fish. If this should correlate with the degree of disarticulation, a check on this interpretation should be possible. Sufficient data are not yet available, however, for the results to be conclusive.
It is not intended to suggest that the conditions outlined above could account for all of the fish concentrations in the Green River Shales. In certain shales in the Green River Basin, for example, the concentrations of Knightia in nonvarved shales appear to have been deposited in quite shallow water. These fish are extremely well preserved but were obviously laid down under conditions quite different from those at Fossil Butte. Much study of these occurrences will be necessary before interpretations are possible.
The story, as told here, should make it clear that the earth is ever-changing. What was once a beautiful, deep lake in an area of lush tropical forests is now a dry, sagebrush desert. What was once an area in which sediments were accumulating is now an area of erosion, occasioned by a broad uplift of the region near the end of Tertiary time.
It is hoped that this brief story of the geologic history of Fossil Butte National Monument will give the reader some appreciation of the geologic complexity not only of the monument itself but of the surrounding area as well. Certainly such knowledge will add to the enjoyment of a visit to the monument.
It may seem that geologists have all of the answers. This, however, is not so. Interpretations are made on the basis of available evidence. Each time a geologist studies an area more geologic data and more evidence become available and our interpretations become a little more accurate. It will be many years before all details of this fascinating history will be known.
This report, which was prepared under National Park Service Contract CX 6000-3-0076, leans heavily on the publications of those many people who have studied various geologic aspects of the Fossil Basin. We would like to cite particularly Doctors Steven Oriel and Joshua Tracy of the United States Geological Survey, whose studies have clarified the stratigraphic relationships of the sediments involved. Dr. D. L. Blackstone, Jr., of the University of Wyoming supplied information from his yet unpublished interpretations of the complicated structure of the Idaho-Wyoming thrust belt. Mr. Tom Bown helped with the structural part of the report and critically reviewed other parts. The drafting, except for Figs. 3 and 9, was done by Mr. Evan Groutage. Mrs. Elaine Hertzfeldt, secretary for the Department of Geology, aided greatly in preparing this manuscript.
Paul O. McGrew Michael Casilliano