terrestrial biota in coastal marine deposits: fossil-lagerstätten in the pennsylvanian of kansas,...

E L S E V I E R Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-273

PAIAEO

Terrestrial biota in coastal marine deposits: fossil-Lagerst itten in the Pennsylvanian of Kansas, USA

Hans-Peter Schultze Institut fiir Paldontologie, Museum fiir Naturkunde der Humboldt Universitdt, Invalidenstr. 43, D-lOll5 Berlin, Germany

Received 22 february 1994; revised and accepted 8 March 1995

Abstract

The fossil animal assemblages of three Pennsylvanian marine Lagerst~itten of Kansas are compared. Detailed taphonomic investigations and comparison of total fossil assemblages are the most reliable for palaeoenvironment interpretation. Functional morphology, taxonomic uniformitarianism, behavior, absence of marine fossils (for freshwater), and palaeogeography are weak criteria of a hypothesis of freshwater depositional environment based on the assumption that closely related taxa have similar environment preferences, functions, and behaviors. In general, invertebrates, being less mobile, are better indicators of their general environment than vertebrates. Terrestrial vertebrates can dominate the fossil vertebrate assemblage of a clearly marine depositional palaeoenvironment. As long as acceptable indicators of Palaeozoic freshwater environment are not found, the hypothesis of freshwater environment cannot stand the test, only easily falsified by the discovery of marine fossils. Fishes may indicate marine connectedness, not necessarily a marine depositional environment.

1. In trod uc t ion

"People are credulous creatures who f i nd it very easy to believe and very difficult to doubt. In fact, believing is so easy, and perhaps so inevitable, that it may be more like involuntary comprehension than it is like rational assessment." D.T. Gilbert (1991) " How Mental Systems Believe". Am. Psychol., 46(2), p. 117.

Fish fossils occur frequently together with plant and ar thropod remains; normal-marine, shelly fossils are missing. These assemblages are often interpreted as freshwater because marine shelly fossils are rare or absent, even though it may only be a preservational bias towards phosphatic and organic fossils. As example f rom my own experience, at the Tithonian locality at Nusplingen

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in southern Germany, vertebrates, plants, and crustaceans are well preserved in Solnhofen-like platy limestones, but shelly fossils are absent. Occasionally one finds the faint impression of an ammonite, even though they were very common judging f rom the common occurrence of aptychi. Only the originally organic parts of the aptychi are preserved like the crustaceans, the calcareous part, like the shells of the ammonites, is absent. Turbidites (lithographic stones) are intercalated between the platy limestones; these turbidites con- tain shelly fossils. This is only one example to show that one should avoid using negative arguments (absence of...) as an indication of environment; nevertheless negative arguments are commonly the basis for recognizing freshwater palaeoenvironments (see Gray, 1988, p. 20).

256 H.-P. Schultze/Palaeogeography, Palaeoclimatolo~,9,, Palaeoecology 119 (1995) 255 273

Another approach used in recognizing freshwater palaeoenvironments is the principle ofactual- ism; the environment of extant genera or species is applied to that of fossil forms. "The reliability of the analogy approach to paleoecology depends on the accuracy of the underlying systematics and functional morphology. Inferences become less certain when older periods of geologic time are involved..." (Elder and Smith, 1988, p. 579). This uncertainty is demonstrated by the co-occurrence of extant freshwater and marine forms in ancient strata; the co-occurrence of Darwinula and Spirorbis in Pennsylvanian rocks may serve as an example. The ostracode Darwinula occurs today in the freshwater environment and is consequently interpreted as a freshwater indicator in the fossil record (Kaesler, 1989). It occurs together with the polychaete Spirorbis in Pennsylvanian rocks (Cunningham, 1993a). Extant Spirorbis is known only from marine environments (Kelber, 1987). The co-occurrence of these two forms can be interpreted in two ways:

(1) Either one has changed its environmental preference. Commonly it is accepted that Spirorbis occurs in freshwater during the Carboniferous (Trueman, 1942: Spirorbis on freshwater bivalves Naiadites, Carbonicola, Anthraconauta, and rare on Anthracomya) because preference in interpre- ting the palaeoenvironment is given to other forms, such as Darwinula or to the absence of other marine forms. Trueman (1942) even suggested generic separation based on interpretation of depositional environment without differences of charac- ters; Carboniferous freshwater spirorbids be separated generically from marine Carboniferous spirorbids (i.e., marine cementstone of Great Britain with Spirorbis: Belt et al., 1967, p. 717; Spirorbis on marine Posidonomya: Prestwich, 1840; Firket, 1878). Freshwater spirorbids have been described only from the Carboniferous (Beckmann, 1954). Spirorbids occur in older rocks including marine Devonian even on plants (Jux, 1964) and younger marine Triassic rocks on plants (Kelber, 1987) in marine environments like those of the present.

(2) One or the other may have been swept into the deposit. That can be demonstrated by Darwinula from the Pennsylvanian rocks near

Hamilton, Kansas. In different layers, but together on the same layer of the Hamilton Lagerstfitte Spirorbis and DarwinuIa are a very common component of the assemblage. Their co-occurrence in one layer is explained as a change from marine to freshwater depositional laminae. Darwinula is more abundant in lighter laminae, which may represent deposits that resulted from draining of freshwater into the Hamilton palaeochannel (Cunningham, 1993a: in addition other marine ostracods, fora- minifers and macrofossils).

Some fossil forms are used as indicators of freshwater for different historical reasons (e.g., they occur in intermontane basins, or they were once misidentified and compared with recent freshwater forms such as the Old Red Sandstone cepha- laspids identified as Lepi~'osteus by Cuvier) even though we know today that they occur in freshwater and marine environments. Another example are the xenacanths, the so-called freshwater sharks of the Palaeozoic, which occur in marine (Bardack, 1979; Schultze, 1985) and freshwater environments (Zangerl, 1981). Nevertheless, their occurrence is used to interpret a palaeoenvironment as freshwater even though there is no other evidence (Masson and Rust, 1984), and in turn Masson and Rust's (1984) palaeoenvironmental interpretation was used to argue that xenacanths were freshwater forms (Gray, 1988, p. 105).

In this paper I will comment on the criteria noted by Gray (1988) as useful in the interpretation of palaeoenvironments, specifically freshwater environments. I will then use three Pennsylvanian localities to demonstrate that faunal composition is crucial in inferring the palaeoenvironment and that the use of preconceived ideas about palaeoenvironmental preference of one kind of fossil can seriously affect the interpretation. Invertebrates are often better indicators of palaeoenvironments because vertebrates as nektonic organisms are generally more mobile than invertebrates.

2. Gray's criteria for recognition of palaeoenvironments

Gray (1988) presented eight criteria for recognizing palaeoenvironments, specifically freshwater habitats, together with examples.

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-2 73 257

(a) Depositional environment: Gray (1988) listed it as a criterion but it represents the question to be answered about the animal " i f it can be demonstrated that it is the environment in which the organism lived" (Gray, 1988: p. 11). The Joggins locality in Pennsylvanian rocks of Nova Scotia provides "unequivocal evidence of terrestrial habitat" (Gray, 1988, p. 11). Early reptiles found in upright tree stumps may have lived in a terrestrial habitat based on their morphology and thus are used to draw a conventional picture of reptiles roaming around tall trees (Carroll, 1970). Nevertheless the trees had to be destroyed first and the area inundated (Dawson, 1863, by the sea) before the tree stumps could form traps for animals. After all this, tetrapods may have ventured out onto the tidal flats as indicated by tracks and ripple marks as described by Dawson (1863, p. 9: "... they haunted tidal flats and muddy shores..."). Within the stumps is a mixture of animals. Millipedes and spiders, like reptiles and amphibians, have lived in a terrestrial environment at least most of the time (see criterion (b) below). Co-occurring palaeoniscoid, rhizodont and megalichthyid fish scales indicate an aquatic environment, as does Spirorbis (Dawson, 1854, 1855). Spirorbis occurs attached to tree trunks and leaves; and Dawson (1855, p. 147) compared it with "those now found adhering to the seaweeds of the shore (the common Spirorbis spirillium)". Dawson (1863, p. 10) argued at length that the trees were not marine: "remarkable absence at least of open sea animals", and that the environment of the growing trees may have been brackish to fresh. All this is circumstantial evidence based on the supposed life style of the past animals, not on depositional arguments. Taphonomy of the trees (see criterion (f) below) in the deposits at Joggins has not been investigated. The reptiles may have been truly terrestrial, but the evidence suggests deposition of the sediments on tidal flats. Gray (1988) cited a long list of Holocene examples where marine forms were deposited with freshwater forms, as well as Pleistocene examples, especially from estuaries. In conclusion she stated that physical and sedimentological criteria are inadequate to identify freshwater deposits.

(b) Functional morphology: This is sometimes a

useful criterion as long as the fossil animal is not too different from its extant analogue, and it fits best if extant and fossil forms are closely related. In contrast, Lauder (1992) asked for great caution in inferring function from the morphology of extinct taxa based on his experience with extant taxa. Gray (1988) was also cautious and cited examples of Tertiary mammals having a combination of features that defy any reasonable explana- tion of their function when compared to extant forms.

(c) Taxonomic uniformitarianism: This "means the identification of fossils as modern taxa.., at any taxonomic level" (Gray, 1988, p. 19). A basic requirement is the existence of an accepted phylog- eny so that the habitat of closely related forms is compared. Still one has to be very careful as noted by Gray (1988), who cited many exceptions, namely where the ancestors of recent freshwater forms are marine. Again the criterion is best suited for geologically young and closely related forms. Still, considering the habitat breadth of many recent genera, one may be able to pick and choose the environment desired.

(d) Behavior: Organisms only distantly related at ordinal or class level are assumed to have behaved similarly. Gray (1988) considered this a weak argument. For example, Maisey (1989) compared the behavior of the hybodont Hamiltonichthys from the Pennsylvanian near Hamilton, Kansas (see below), with that of very distantly related sting rays in the Amazon River. Neonate sting rays migrate downstream to higher salinity where they grow before they return upstream. Compared to the large Amazon River, the sea was near the site of the Hamilton Lagerst~itte (Schultze et al., 1993), and thus the hybodonts could migrate only a few kilometers into the tidal channel. Thus, there is no way to test such a scenario. Another example where the same palaeoenvironment cannot be deduced from the same behaviour even in (superficially) similar cases follows. Two lungfish genera form flask-like burrows, the recent Protopterus and the late Palaeozoic Gnathorhiza. The burrows of the latter were interpreted as aestivation burrows like those of Protopterus (Romer and Olson, 1954; McAllister, 1992). Gnathorhiza occurred in marine

258 H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-273

(see below and Berman, 1970; Schultze, 1985; Olson, 1989) and possibly freshwater palaeoenvironments (see McAllister, 1992, who omitted the data of Brown, 1970; Schultze, 1985 and Olson, 1989); its burrows occured--hundreds of millions of years ago in the intertidal or supratidal coastal region (Schultze, 1985) close to the coast line and not far inland, unlike Protopterus. Schultze (1985, p. 13) interpreted the burrows of Gnathorhiza as "retreats during tidal changes."

(e) Absence of marine fossils': This is a negative argument and as such is logically very weak. From the hypothesis that rocks of a specific locality were formed in freshwater, the prediction follows that typical marine fossils are absent. Even if we find no marine fossils (the test meets the prediction), we can assume only support but not test for the hypothesis, because we could have just missed the marine fossils; or chemical differences in preservation may have destroyed the calcitic shells typical of marine forms. Of course, discovery of marine fossils falsifies the hypothesis.

(f) Taphonomy: Here Gray (1988, p. 23) was cautious to put much importance on this criterion because she referred only to general investigations, ("Extreme care must be exercised..."), whereas careful taphonomic studies give the most detailed data for interpretation of the palaeoenvironment (Zangerl and Richardson, 1963). Such studies are rare; still the criterion is one of the most powerful for inferring palaeoenvironments.

(g) Community analysis: Gray (1988) accepted such an analysis only in combination with relative frequencies of taxa and taphonomic information. Comparison of communities at the same time level should be a better indicator of palaeoenvironment than the comparison with the environment of extant forms (criterion (a)) or the function (criterion (b)) and behavior (criterion (d)) of related forms. Data on which such a hypothesis is based can be checked and the hypothesis tested. Dineley and Williams (1968) argued that the Devonian fish fauna of Miguasha, eastern Quebec, is domi- nantly freshwater. This was easy falsified by sur- veying the distribution of Devonian vertebrates (Schultze, 1972; Schultze and Cloutier, in press). A community analysis that compares contemporaneous localities (performed as cluster analysis)

results in a hypothesis of palaeoenvironment which can be tested using other criteria (e.g., taphonomy, occurrence of additional taxa, geochemical data).

(h) Palaeogeography: Gray (1988) suggested mapping a shoreline over a large area to distinguish between marine and freshwater environments. This method stacks one interpretation on another (the palaeogeographic position on an assumed distinc- tion of marine from freshwater, the questionned palaeoenvironment) as can be shown in the examples given by Gray (1988, p. 24). Based on personal communications with A.J. Boucot, she argued against Thomson's (1980) interpretation of the Lower Devonian Beartooth Butte Formation as coastal marine deposits because it "is inconsistent with its paleogeographic situation several hundred kilometers away from any contemporary shoreline". Early dipnoans and those of the Water Canyon and Beartooth Butte formations occur in marine deposits (Thomson, 1980; Campbell and Barwick, 1988), and the agnathans of the Beartooth Butte Formation are widely distributed in marine deposits of northern Canada (Elliott, 1993). Thus, the most parsimonious interpretation indicates a marine environment (Janvier and Blieck, 1993). That confirms earlier investigations. Doff (1934) refers to estuarine channel-fill deposits, Sandberg (1961, p. 1308) to marginal marine depositional environment for the fossils bearing deposits and Johnson et al. (1988, p. 167) to brackish water and, possibly, freshwater channel- fill deposits. In the meantime, Ilyes and Elliott (1992) have shown that outcrops in Wyoming form a continuous shoreline with occurrences of similar forms in Idaho, Utah and Nevada and south to Death Valley, California. The Cottonwood Canyon locality of the Beartooth Butte Formation may be older than the other localities (Elliott and Ilyes, in press), even though the same dipnoan occurs in the Water Canyon Formation of Idaho and the Cottonwood Canyon locality of Wyoming. Basically, the geological data base for Gray's (1988) statement was inadequate.

Gray's second example is the lowermost Upper Devonian Escuminac Formation of Miguasha, Quebec; she referred again to personal communications from A.J. Boucot. The occurrence of the Escuminac Formation is limited to an intermon-

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255~73 259

tane basin (Gray, 1988, p. 24) or, better, limited by tectonic features. There is no evidence for a freshwater environment except the present-day dis- tance to the nearest contemporaneous marine Devonian deposits, but the role of tectonics and erosion were not considered. The fish assemblage (criterion (g)) indicates a marine environment (Schultze, 1972; Schultze and Arsenault, 1985 and V6zina, 1991; in contrast to Dineley and Williams, 1968). The sedimentary structures (criterion (f)) "appear identical with those described by Sorauf (1965) in the Upper Devonian of south-central New York State" (Dineley and Williams, 1968, p. 252); and the Upper Devonian rocks to which Dineley and Williams (1968) referred are marine as indicated by their invertebrate content. Also, chemical analyses of the rocks (V6zina, 1991) and of bones of Bothriolepis (Schmitz et al., 1991), discovery of marine trace (contra criterion (e)) fossils (Maples, in press) and occurrence of acritarchs (Cloutier et al., in press) support the interpretation of a nearshore marine palaeoenvironment for the Escuminac Formation, perhaps a silled estuary (Hesse and Sawh, 1992). This requires only a different palaeogeographic recon- struction of the tectonically dislocated Acadian area (e.g., a connection through Scotland or England with the Devonian sea in the North Sea area; see Ziegler, 1982, map 8).

In conclusion, only two of Gray's (1988) criteria are strong: taphonomy and community analysis. The other criteria are weak and based on the assumption that similar taxa have similar environmental preferences, functions, and behavior. Presence of marine fossils suggests a marine palaeoenvironment and is positive evidence rather than negative as stated by Gray. In some instances, geochemical analyses may support taphonomic or community analysis.

3. Comparison of three Pennsylvanian localities

The three Pennsylvanian localities are loca- ted along a northeast-southwest outcrop of Stephanian rocks in eastern Kansas (Fig. 1). The Robinson locality in northeastern Kansas, is the youngest locality (Soldier Creek Shale Member,

Canada

United States

KANSAS

/

o % 0 500km I ~1

!

Fig. 1. Geographic location of the three Stephanian vertebrate Lagerst~tten in Kansas: Robinson in Brown County, Hamilton in Greenwood County, Garnett in Anderson County.

Bern Limestone, Wabaunsee Group, Virgilian, Stephanian C, Gzelian; Fig. 2). Hamilton is older (Topeka Limestone, Shawnee Group, Virgilian, Stephanian B, Gzelian; Fig. 2), even though the exact age (supposedly corresponding to the lower part of the Topeka Limestone) is unknown because overlying deposits are not preserved. Garnett is the oldest of the three localities (Rock Lake Shale Member, Stanton Limestone, Lansing Group, Missourian, Stephanian B, Kasimovian; Fig. 2).

Of the three localities Robinson is the most

260 H. -P. Schuhze/Palaeogeography, Palaeoclimatology, Palaeoecology l 19 (1995) 255 273

, . ._

~°~°mS~e i i R o ~ n ~ o n ) a ~ , u ~ a Ls Mb, ,

Sold e Creek Sh Mbr . . . . . . . . . . . . . I

~ ± : Scranton Shale

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Fig. 2. Generalised section of Stephanian of Kansas (modified from Zeller, 1968) with the stratigraphic position of the three vertebrate Lagerst~itten on the left.

aquatic, and Garnett is the most terrestrial. This interpretation is supported by the lithologies (carbonate at Robinson and Hamilton versus ternge-

nous at Garnett) and associated animal (Schultze and Chorn, 1989) and plant (stromatolites and oncolites at Robinson versus conifers at Hamilton and Garnett) fossils at each locality.

3.1. Robinson locality

Beede (1899) described the stromatolites as sponges (Somphospongia multiformis) from shales with "typical marine Coal Measures fossils... immediately associated with Lingula, Productus and a few pelecypods and gastropods" (p. 129). Johnson (1946) assigned Somphospongia to the marine algae within the Burlingame Limestone to which member within the Bern Limestone Formation the Robinson locality was placed. Nevertheless the vertebrate palaeontologist Baird (1966) referred to "a fresh-water limestone member of the Burlingame Limestone," probably because he assigned the lungfish Sagenodus and the co-occurring amphibian to freshwater by analogy with occurrences of extant lungfish and amphibians. Chorn and Conley (1978) and Conley and Chorn (1979) recorded a mixture of marine invertebrates and freshwater vertebrates, including the marine sharks Cladodus, Hybodus, and Petalodus. They interpreted the environment as deposition in a bay or lagoon with a nearby source area (Nemaha anticline) for terrigenous silt. Sawin et al. (1985) published the first detailed analysis of the stromatolites. They considered "the stromatolite biostromes probably.., a local feature in an intertidal to slightly subtidal environment associated with a coastal embayment" (p. 361) and they placed the stromatolites like Beede (1899) into a shale, the Soldier Creek Shale Member of the Bern Limestone. Like Conley and Chorn (1979), they inferred a nearby land area to the north-northwest (Nemaha anticline) as the source of the presumed freshwater organisms. Comparisons by Maples and Schultze (1989) and Schultze and Maples (1992) placed the Robinson locality at the most marine end of the spectrum of late Carboniferous vertebrate localities including Kinney Brick Company Quarry, New Mexico; Hamilton, Robinson, and Garnett, Kansas; Essex and Braidwood (Mazon Creek), Illinois; Linton, Ohio; Montceau, France; and Nyrany, Czech Republic.

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-273 261

The Robinson locality represents a Concentrat- Lagerst~itte (subtype "concentration traps" by algal encrustation, but not formed in cavities) in the sense of Seilacher (1970). The fauna (Table 1) is taxonomically and possibly volumetrically dominated by marine invertebrates; terrestrial invertebrates are missing in contrast to the other two localities; and only one form (Lingula) is tolerant of brackish-water conditions. In addition, teeth of unquestionable marine elasmobranchs (Petalodus, hybodontid, cladodont, deltodontid teeth) have been recovered. Most of the vertebrates are aquatic, only a few isolated bones of terrestrially adapted vertebrates (Cricotus, protorothyridid, pelycosaur) have been found. Vertebrate bones are stained dark brown to black and are easily recognized. They are prepared using acetic acid, which commonly destroys calcitic skeletons of invertebrates. Therefore, invertebrates are underrepre- sented in vertebrate preparations. By counting specimens in peels of polished sections, Sawin et al. (1985) have shown that invertebrates comprise 70-80% of the skeletal components of the assemblage; only in a few stromatolites do vertebrates account for more than 30% of the assemblage, and in one instance vertebrates comprise 85.7% of the assemblage.

Most of the vertebrates are aquatic or amphibi- ous and are often used as freshwater indicators (see also Sawin et al., 1985) even though they occur in marine rocks sometimes more frequently (as teeth) than in rocks that were deposited in freshwater environment. This is particularly true of xenacanths, acanthodians, palaeoniscoids, actinistians, rhipidistians, and dipnoans (Schultze, 1985). Gnathorhiza is the only known Palaeozoic dipnoan that burrows. It is usually compared with the recent Protopterus from Africa, the only extant genus that forms aestivation burrows, and is therefore interpreted as a freshwater organism (see above). Schultze (1985) argued for the possibility that Gnathorhiza developed the habit as an adaptation to intertidal to supratidal environments. At the Robinson locality Gnathorhiza forms a major component of the vertebrate assemblage; it occurs like Sagenodus, the other dipnoan of the Robinson locality, as disarticulated specimens within the stromatolites. Burrows are unknown at the local-

ity. Because it is a major component of the assemblage it is unlikely that its remains were selectively transported into the marine environment. One has to accept that Gnathorhiza is euryhaline like other fishes at this locality. Because there is always a degree of uncertainty, most fish and the amphibians are listed as freshwater to marine in Table 1. Fish remains are disarticulated teeth, scales and spines, their common mode of occurrence is in marine rocks (Schultze, 1985). Saltwater tolerance is the exception in extant amphibians, but that was not necessarily so for Palaeozoic forms. The Devonian ancestors were at least tolerant of saltwater. Tulerpeton, the Late Devonian tetrapod from Russia, occurs in marine sediments with marine fishes (Lebedev and Clack, 1993). Thomson (1980) argued that Devonian tetrapods from east Greenland were tolerant of saltwater. The time between the deposition of the Devonian tetrapods and their Carboniferous descendants is shorter than the time from the Carboniferous to the Holocene, so a comparison of environmental adaptation with Devonian forms is more reasonable than the comparison with extant forms; although such a comparison is not conclusive. Also Schultze (1985) and Milner (1987) argued that some Carboniferous amphibians are tolerant of saltwater. "An intertidal to shallow subtidal environment" (Sawin et al., 1985, p. 370) is accepted for the Robinson locality based on its invertebrate fauna; but a nearby land mass which is unknown because the Nemaha anticline is not close and is not known, for certain, to have been emergent at this time, is not necessary to explain the occurrence of the vertebrates except for the few bones of rare terrestrial amniotes.

3.2. Hamilton locality

Whereas the Robinson locality has been known since the last century, the Hamilton locality was discovered in 1964 by an oil man, W. Lockard (Bridge and Mapes, 1989). Lockard collected the well-preserved part of the assemblage, vertebrates (tetrapods, fishes and arthropods) and plants (mostly conifers), but also a few invertebrates such as brachiopods (Bridge et al., 1972); the locality was interpreted as a freshwater deposit. Zidek

262 H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-273

Table 1

Environmental distribution of animal taxa in the Robinson Concentrat-Lagerstfitte showing interpreted environmental preferences

and relative abundances. Arrows between environments denote ranges in environment or uncertainty about environmental preference

Terrestrial Freshwater Brackish Marine

TETRAPODA ANTHRACOSAURIA Cricotus CAPTORHINOMORPHA pro lo ro thyr id id sp.

PELYCOSAURIA i nde te rminan t

ELASMOBRANCHII X E N A C A N T H I D A

Orthacanthus A C A N T H O D I I

A c a n t h o d e s

A C T I N O P T E R Y G I I

palaeoniscoids

A C T I N I S T I A

indeterminant RHIPIDISTIA megal ich thy id

DIPNOI Gnathorh&a Sagenodus TETRAPODA NECTRIDEA Diplocaulus LYSOROPHIA indet.

TEMNOSPONDYLI Platyhystrix t r ime ro rhacho i d

FORAMINIFERA fusulinids

encrusting foraminifera BRYOZOA fenestrate and ramose BRACHIOPODA Lingula discinid

Crurithyris Derbyia ?Dielasma Neochonetes Punctospirif er product id , indet. BIVALVIA Aviculopeeten ?Nuculopsis Permophorus Phestia Septomyalina myal in id

incertae sedis

+

+ +

+

+

+

+ +

+

+

+

+

2>

2>

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2>

2>

2>

-2>

2>

-2>

2>

2>

+ -2>

+

+ +

+

+

+

+ ÷

+ + +

+

+

+

+

÷

÷

+

+

+

+ + q-

+

+

+

+

+

+

+

+

+

+

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995)255 273

Table 1 (continued)

263


G A S T R O P O D A ?Euconospira + Euphemites + Glabrocingulum + Straparollus + Strobeus + ? Trepospira + Worthenia + O S T R A C O D A Bairdia + healdiacean + hollinellid + geisinid + TRILOBITA undetermined + E C H I N O D E R M A T A Crinoids + Echinoids + E L A S M O B R A N C H I I P E T A L O D O N T I D A Petalodus + EUSELACHII hybodontid indet. + U N D E T E R M I N E D cladodont + H O L O C E P H A L I deltodontid +

(1976, p. 3) regarded it as having been deposited in the "low energy part of a stream." That was the consensus of the contributors at the annual meeting of the Kansas Academy of Science in 1976 in Emporia, Kansas, even though Andersen cited brachiopods as components of the assemblage. As the freshwater environment seemed to be supported by upland plants (Leisman, 1971, 1976; Leisman et al., 1989). At a second symposium on the Hamilton site (1988 annual meeting of the South- Central Section of the Geological Society of America in Lawrence, Kansas), the controversy between freshwater and marine interpretations began. Components of the assemblage, including animals (Bridge, 1989; Mapes and Maples, 1989; Maples and Mapes, 1989; Kues, 1989; Kaesler, 1989; Zidek, 1989) and plants (Bridge, 1989; Leisman et al., 1989), were interpreted as freshwater indicators; whereas French et al. (1989), Busch et al. (1989), Schultze and Chorn (1989), and

Maples and Schultze (1989) placed the locality in a coastal marine setting based on geological criteria and total assemblage of fossil animals. Mapping and new excavations supported the latter interpretation (detailed taphonomic studies by Cunningham, 1993b; Cunningham et al., 1993; Feldman et al., 1993; Schultze et al., 1994). Rhythmic pattern of lamination thickness variation in the limestones and mudstones indicate that the vertebrate bearing layers were deposited in a tidal environment. Especially enlightning is the list of fossil animals by Cunningham (1993a). Besides such marine invertebrates as bryozoans, articulate brachiopods and echinoderms he reported marine ostracodes in addition to the very common Darwinula and the common occurrence of foraminiferids.

The Hamilton locality is a Conservat- Lagerst~itte (subtype "obrution deposits") with so called skin preservation of acanthodians and dis- sorophoids due to microbially induced early dia-

264 H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255 273

Table 2 Environmental distribution of animal taxa in the Hamilton Conservat-Lagerst/itte showing interpreted environmental preferences and relative abundances. Arrows between environments denote ranges in environment or uncertainty about environmental preference


A R A C H N I D A Prothelyphonus M Y R I A P O D A euphoberiid ?juliform milliped INSECTA BLATTARIA P A L A E O D I C T Y O P T E R A P R O T O R T H O P T E R A T E T R A P O D A C A P T O R H I N O M O R P H A protorothyridid sp. DIAPSIDA araeosceloid P E L Y C O S A U R I A edaphosaur ophiacodont BIVALVIA Anthraconia A N N E L I D A Spirorbis O S T R A C O D A Carbonita Darwinula Geisina S Y N C A R I D A palaeocaridids E U R Y P T E R I D A Adelophtalmus E L A S M O B R A N C H I I X E N A C A N T H I D A Expleuracanthus Orthacanthus Xenacanthus EUSELACHII Hamiltonichthys Palaeoxyris (egg capsule) A C A N T H O D I I Acanthodes breeding OSTEICHTHYES A C T I N O P T E R Y G I I palaeoniscoids RHIPIDISTIA megalichthyid DIPNOI Gnathorhiza Sagenodus T E T R A P O D A T E M N O S P O N D Y L I dissorophid breeding eryopoid tr imerorhachoid

÷ ÷

+ ÷ + ÷ ÷

÷

÷

+ +

+ ÷ + + ÷ + +

+ + +

+ ÷ + +

+ ÷ ÷

+ ÷ + +

÷

÷

÷

~> ~>

~>

~>

~>

÷ ÷ +

+ +

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255-273

Table 2 (continued)

265


+ - > + R E P T I L E indet. F O R A M I N I F E R A

fusulinids ( Triticites) Globivalvulina encrusting foraminiferids PORIFERA sponge spicules BRYOZOA fenestrate and ramose B R A C H I O P O D A

Antiquatonia Juresania Koziowskia Neochonetes Neospirifer Punctospirif er productid indet. B I V A L V I A

Myalinella Permophorus Phestia Schizodus G A S T R O P O D A

?Bellerophon Euphemites A N N E L I D A

Serpula O S T R A C O D A

Amphissites Bairdia Gutschickia Pseudobythocypris E C H I N O D E R M A T A

crinoids echinoids

-k - ~ -

+ + + +

+

+

+

+

+

+

+

+ + +

+

+ + +

+

+

+

+

genesis. The assemblage (Table 2) is dominated volumetrically by freshwater and freshwater to marine forms; marine fossils form taxonomically a slight majority. Historically, such terrestrial "upland" plants, myriapods, and insects and such freshwater to marine forms as eurypterids, acanthodians, palaeoniscoids, lungfish, and temnospon- dyl amphibians were preferentially collected. In contrast, layer by layer excavations in 1988 and 1989 and careful recording of all animal and plant components revealed an abundant marine component throughout the Lagerst~tte sequence (Cunningham, 1993a). Terrestrial forms, including

arachnids, myriapods, and insects, and "upland" plants do not indicate the palaeoenvironment; these forms are washed, or blown (or "fall") into the deposit. Frequent occurrence of fossil charcoal was suggested as evidence for forest fires (Maples et al., 1989), which are commonly associated with strong winds. Terrestrial amniotes are disarticulated (captorhinomorphs; Cunningham and Reisz, in press) or represented by only one element (edaphosaur: two isolated spines; ophiacodont: a max- illa); an exception is a nearly complete araeosceloid (Reisz, 1989). Preservation of these terrestrial amniotes indicates that they are an allochthonous

266 H.-P. Schultze/Palaeogeography, Palaeoclirnatology, Palaeoecology 119 (1995) 255 273

element of the fauna. In different layers of the Lagerst~itte Spirorbis and Darwinula are a very common component of the assemblage, the former suggesting a brackish to marine environment and the latter a freshwater environment. Their co-occurrence in one layer is explained as change from marine to freshwater depositional laminae (Cunningham, 1993a). Darwinula is more abundant in lighter, carbonate-rich laminae that may represent deposits due to runoff following rains. An estuarine environment may have been a preferred environment for such bivalves as Anthraconia; whereas the ostracode Geisina, the syncarids, and eurypterids are placed between freshwater and marine (Table 2), because they are found in rocks that have been formed in both environments. A range from freshwater to marine is accepted for all the fishes based on their Carboniferous record; none of the cited genera can be used to characterize a specific palaeoenvironment. Acanthodians and dissorophid amphibians occur as small, immature to nearly adult forms indicating that the area could have been the breeding ground for the two species; presence of a Palaeoxyris egg capsule, presumably from a hybodont shark, supports this inference. Many extant marine fish enter estuaries to deposit their eggs (Gunter, 1961). Developmental stages of the dissorophid amphibian indicates that it tolerated some degree of salinity. The eryopoid (one shoulder girdle) and trimerorhachoid (a segment of verte- brae) amphibian are allochthonous elements; whereas the aquatic reptile (Reisz, 1989), a single specimen, may be an autochthonous component.

The taxonomically rich marine invertebrate assemblage indicates a special marine palaeoenvironment, an estuary that is close to terrestrial and freshwater effects (Schultze et al., 1993). That interpretation is supported by shape of the outcrop (Fahrer, 1991) and the presence of tidal deposits (Feldman et al., 1993).

3.3. Garnett locality

Mapping Andersen County, Kansas, Norman D. Newell discovered plant and vertebrate fossils at a locality north of Garnett in 1931. The associated invertebrates indicate a marine origin for the deposit even though the plants were characterized

as an "upland" flora (Moore et al., 1936). The locality became famous for terrestrial vertebrates and plants; nevertheless the deposits were still interpreted as having come from a nearshore marine environment (Lane, 1945). Peabody ( 1952, 1954; Moore, 1966) proposed a more detailed model with a river entering a lagoon cut off from the open sea by a barrier; the river transported terrestrial animals and plants into the lagoon. Gymnosperms dominated the marginal coastal flora, and tracks indicated subaerially exposed parts of the lagoon. Reisz et al. (1982) modified Peabody's model; they described a sequence start- ing with a terrestrial environment that was suc- ceeded by tidal mudflats deposited in an estuarine channel finding no indication of an entering river. Woodruff (1984) did a detailed taphonomic study; he recognized eight facies within the channel begin- ning with an estuarine facies containing both marine and terrestrial fossils followed by lagoonal and tidal-flat deposits with intercalated storm deposits. All these environments occur in a coastal marine setting.

The Garnett Conservat-Lagerst~itte (Table 3) contains more terrestrial forms than the other two Kansas localities; nevertheless there are still more marine than terrestrial taxa. Vertebrates and bryozoans and brachiopods occur in the middle part of the Rock Lake Shale (facies V of Woodruff, 1984, which is the only portion considered by Reisz et al., 1982). Body fossils of terrestrial vertebrates are more common in the lower part of facies V, which is interpreted as having been deposited in a subtidal to intertidal environment. Trackways are more often encountered in the upper part interpreted to be of intertidal to supratidal origin. Actinistian fish occur throughout facies V. To have been preserved in this setting, Petrolacosaurus and the pelycosaurs must have ventured out onto the tidal flat; however, the trackways were not produced by these tetrapods, the trackways are assigned to other tetrapod genera. Such additional terrestrial forms as insects and scorpions are extremely rare. Along with the abundant plant remains they may have been blown into the lagoon by strong winds associated with forest fires. As with the Hamilton locality described above, the Garnett locality is rich in fossil charcoal.

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995) 255~73 267

Table 3 Environmental distribution of animal taxa in the Garnett Conservat-Lagerst~Rte showing interpreted environmental preferences and relative abundances. Arrows between environments denote ranges in environment or uncertainty about environmental preference


I N S E C T A Blattaria indet. Megasecoptera indet. Euchoroptera longipennis Parabrodia carbonaria SCORPIONIDA Garnettius TETRAPODA CAPTORHINOMORPHA Notalacerta t r a c k w a y s DIADECTIDAE Megabaropus trackways DIAPSIDA Petrolacosaurus P E L Y C O S A U R I A Haptodus Xyrospondylus Ianthacosaurus o p h i a c o d o n t n.gen, et sp. O S T R A C O D A indet . A C T I N I S T I A Synaptotylus AMPHIBIA DISSOROPHOID A ctiobates I N D E T . Hesperoherpeton FORAMINIFERA fusul in ids PORIFERA sponge spicules C O E L E N T E R A T A R U G O S A ?Lophophyllidium BIVALVIA Myalinella Sedgwickia Yoldia BRACHIOPODA Lingula Composita Neospirif er BRYOZOA Fenestella Polypora Rhombopora A N N E L I D A serpulid ARTHROPODA t r i lob i te ECHINODERMATA crinoids echinoids ELASMOBRANCHII h y b o d o n t i d " C l a d o d u s "

÷ ÷ ÷ ÷

+

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÷

+ ÷

÷ + + ÷ ÷

÷

÷

÷

+

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+

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÷

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÷

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+

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268

GARNETT

Tot a I ( ~ Assemblage ~

H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995)255 273

HAMILTON ROBINSON

Invertebrata

Vertebrata

Terrestrial ~ Freshwater -~ marine [ ] Marine

Fig. 3. Distribution of marine, freshwater to marine, and terrestrial genera at the three Stephanian localities in Kansas (data base see Tables 1 -3).

Fishes are rare. Several actinistian specimens have been found as articulated and disarticulated specimens, but the two elasmobranchs are known only from a single tooth each. Nearly all recovered invertebrates are marine.

4. Conclusions

The three localities Robinson, Hamilton, and Garnett, respectively have a decreasing number of marine forms and an increasing number of ter-


restrial forms if one considers the total assemblage (Fig. 3). The percentage of marine invertebrates is high in all three localities (100-66%) so that an interpretation that the rocks of all three localities were deposited in a marine environment, seems to be justified; a marine depositional environment is compatible with the geological and taphonomical investigations of the three localities. The number of terrestrial invertebrates is highest at Hamilton, even though they comprise a lower percentage than at Garnett. Hamilton is the only locality of the three that has a number of freshwater to brackish-water invertebrates.

The vertebrate assemblage does not show the same picture as the invertebrate assemblage. Aquatic vertebrates decrease from 83% of all vertebrates at Robinson to 38% at Garnett; the terrestrial forms increase. Robinson and Hamilton are similar in the distribution of aquatic and terrestrial vertebrates; only Hamilton has no fish that are accepted unequivocally as marine indicators. In Tables 1-3 and Fig. 3, all vertebrates that have been placed in freshwater or brackish environments at one time or another are counted in the category "freshwater to marine." This uncertainty of the palaeoenvironmental limitations of the fishes opens the door to misinterpretation of palaeoenvironments. Only vertebrate palaeon- tologists (Baird, 1968: Robinson; Zidek, 1976: Hamilton) have interpreted these two localities as freshwater deposits, whereas invertebrate palaeon- tologists accepted the localities (correctly) always as marine despite the fact that Hamilton and Garnett have preserved components of an "upland" flora. In general, invertebrates, being less mobile, are more indicative of their general environment than vertebrates, which move around and can occur in different environments. The three localities are placed either in an intertidal region (Robinson) or estuarine channel (Hamilton, Garnett). Terrestrial forms can, and do, occur in such coastal-marine settings.

Detailed taphonomic investigations (criterion (f) of Gray, 1988) are lhe most reliable basis for palaeoenvironmental interpretations (Robinson: Sawin, 1977; Sawin et al., 1985; Hamilton: Cunningham, 1993b; Garnett: Woodruff, 1984).

Comparisons of fossil animal assemblages (Maples and Schultze, 1989; Schultze and Maples, 1992; criterion (g) of Gray, 1988) support these interpretations. Using only a specific fossil or a group of fossils as an indicator (criterion (c) of Gray, 1988) of a specific environment can lead to misinterpretations (Zidek, 1976; Kaesler, 1989 and others). Occurrences of forms that do not fit within a specific depositional environment must be explained (e.g., Cunningham, 1993a, for Darwinula). Functional morphology can characterize the common habitat of a form, as with terrestrial vertebrates and insects, but it character- izes the habitat to which they are adapted in life, which is not necessarily the habitat of death and burial. They may occur in such uncharacteristic habitats either actively (tetrapods walking onto tidal flats) or passively (insects blown into the area).

Recognition of Palaeozoic freshwater environments is difficult. As long as unequivocal indicators for freshwater are not found, the hypothesis of freshwater environment cannot be tested, only falsified easily by discovery of marine fossils. Comparison of total assemblages between contemporaneous localities is the most powerful indicator if clear marine indicators are missing. But for fishes, this only suggests a connectedness to the marine realm, not necessarily a marine depositional environment.

AcknowLedgments

This paper has benefited from the discussions of the Kansas Lagerst~itten Research Group (A.W. Archer, C.R. Cunningham, H.R. Feldman, C.G. Maples, R.R. West and the author) and discussions with J. Chorn about the Robinson locality. I thank C.R. Cunningham especially for supplying me with most of the references on Spirorbis and C.R. Cunningham, Houston, Texas, and R.R. West, Manhattan, Kansas, for reading critically and improving the manuscript. The manuscript was further improved by critical reviews of A.J. Boucot, J. Gray and R.L. Kaesler. Sharon Hagen of the Division of Biology of the The

270 H.-P. Schultze/Palaeogeography, Palaeoclimatology, Palaeoecology 119 (1995)255 273

University of Kansas prepared the figures. Her work is greatly appreciated.

The National Science Foundation (grants EAR 8903792 and EAR 9018079) and the National Geographic Society (grant no. 4041-89) are grate- fully acknowledged for their support.

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