Life Sciences Contributions Royal Ontario Museum | Bh Naked Species of Gondolella (Conodontophorida): Their Distribution, Taxonomy, and Evolutionary Significance Peter H. von Bitter Glen K. Merrill ROYAL ONTARIO MUSEUM LIFE SCIENCES PUBLICATIONS INSTRUCTIONS TO AUTHORS Authors are to prepare their manuscripts carefully according to the following instructions. Failure to do so will result in the manuscript’s being returned to the author for revision. All manuscripts are considered on the understanding that if accepted they will not be offered for publication elsewhere. ke. GENERAL Papers for publication are accepted from ROM staff members, Research Associates, or from researchers reporting on work done with ROM collections. In exceptional cases,monographic works on the flora and/or fauna of Ontario will be considered for publication by authors not affiliated with the ROM. 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MERRILL LIFE SCIENCES CONTRIBUTIONS ROYAL ONTARIO MUSEUM NUMBER 125 Naked Species of Gondolella (Conodontophorida): Their Distribution, Taxonomy, and Evolutionary Significance ° > ROM C2 ROYAL ONTARIO MUSEUM PUBLICATIONS IN LIFE SCIENCES The Royal Ontario Museum publishes three series in the Life Sciences: LIFE SCIENCES CONTRIBUTIONS, a numbered series of original scientific publications including monographic works. LIFE SCIENCES OCCASIONAL PAPERS, a numbered series of original scientific publications, primarily short and usually of taxonomic significance. LIFE SCIENCES MISCELLANEOUS PUBLICATIONS, an unnumbered series of publications of varied subject matter and format. All manuscripts considered for publication are subject to the scrutiny and editorial policies of the Life Sciences Editorial Board, and to review by persons outside the Museum staff who are authorities in the particular field involved. LIFE SCIENCES EDITORIAL BOARD Senior Editor: J.H. MCANDREWS Editor: R.D. JAMES Editor: C. MCGOWAN PETER H. Von BITTER is Associate Curator-in-charge, Department of Invertebrate Palaeontology, Royal Ontario Museum, and Associate Professor, Department of Geology, University of Toronto. GLEN K. MERRILL is Associate Professor, Department of Geology, College of Charleston, Charleston, South Carolina, and Research Associate, Department of Invertebrate Palaeontology, Royal Ontario Museum. Canadian Cataloguing in Publication Data Von Bitter, Peter H., 1942— Naked species of Gondolella (Conodontophorida) (Life sciences contributions ; no. 125 ISSN 0384-8159) Bibliography: p. ISBN 0-88854-266-6 pa. 1. Gondolella. 2. Conodonts. I. Merrill, Glen K., date II. Royal Ontario Museum. III. Title. IV. Series. QE899.V66 562'.2 C80-094633-2 Publication date: 10 October 1980 ISBN 0-88854-266-6 ISSN 0384-8159 © The Royal Ontario Museum, 1980 100 Queen’s Park, Toronto, Canada MSS 2C6 PRINTED AND BOUND IN CANADA AT THE ALGER PRESS Naked Species of Gondolella (Conodontophorida): Their Distribution, Taxonomy, and Evolutionary Significance Abstract The oldest known species of Gondolella lacking a platform, indeed the oldest known species of Gondolella s.s., is G. gymna, from northwestern Illinois and Japan, of early Desmoinesian and Morrowan-Atokan age, respectively. The concept of ‘‘naked’’ gondolelliform conodonts was based on Gondolella denuda Ellison, a species that occurs in rocks of early Missourian age in Illinois, Missouri, Nebraska and, rarely, in Ohio. A new ‘‘naked’’ species of Gondolella, G. postdenuda sp. nov., occurs in and is characteristic of rocks of Virgilian age. It is common in the Queen Hill Shale of Kansas and Nebraska but also occurs less frequently in the Heebner Shale of Kansas, Oklahoma, and Nebraska, the lower Beil Limestone of Kansas, the Salem School Limestone of northcentral Texas, and the shale overlying the latter member. A second new ‘‘naked’’ species of Gondolella, G. neospathodiformis sp. nov., is known from only a single locality of the Heebner Shale in northern Oklahoma. Gondolella gymna, G. denuda, and G. postdenuda sp. nov. each apparently possessed an apparatus of seven element types, six of which were paired. The ramiform elements belonging to each of these three (as well as those belonging to broad-platformed species) were apparently vicarious, or nearly so, and the ramiform elements of one cannot be distinguished from those of the other without difficulty and considerable uncertainty. Gondolella neospathodiformis sp. nov. is based on a blade-like platform element. The most probable ancestor of ‘‘naked’’ (and other) species of Gondolella appears to be the Spathognathodus bultyncki group of Lower Carboniferous age. The descendants of the former are probably conodonts placed in the genus Neospathodus Mosher. The four ‘“‘naked’’ species of Gondolella may have evolved during the Pennsylvanian in a direct ancestor-descendant relationship. Alterna- tively, the evolution of this group may have involved any of four or more phylogenetically more complex paths—paths that included regressive platform development and iterative mosaic evolution, as well as dimorphism. Introduction The conodont genus Gondolella was defined by Stauffer and Plummer (1932) on the basis of elements that were tongue-, canoe-, or gondola-shaped, that possessed a bar or base (= platform) that was slender to thick and broad, and that had an aboral flaring loop. All of the species included in this genus by its authors possessed broad platforms. In 1941 a narrow blade-like form, Gondolella denuda Ellison, was included in this genus. This was the first of the naked gondolellids known. The term ‘‘naked’’ is used by us much as it was by Ellison (1941), to describe those gondolellids whose Sp (or platform) elements lack a true, broad platform. In this paper we document the occurrence and biostratigraphic, taxonomic, and phylogenetic significance of G. denuda as well as of three additional naked species, two of which are new. Stratigraphic and Geographic Distribution (Fig. 1) The oldest known species of Gondolella lacking a broad platform is Gondolella gymna from the Seville Limestone of northwestern Illinois (Merrill and King, 1971) and from rocks of either similar or older age (Fig. 1) (depending on correlation source) in Japan (Koike, 1967). Slightly younger rocks in northwestern Illinois (‘‘Seville’’ in Merrill, 1975) and western Missouri (Tiawah Member in Vernon Co., Missouri, and the shale in the Scammon Formation below the Tiawah Member in Henry Co., Missouri) contain Gondolella laevis Kosenko and Kozitskaya, a species that has a platform intermediate in width between G. gymna and younger broad- platformed Desmoinesian species. This species was interpreted by Merrill and King (1971) and Merrill (1975) to be an evolutionary intermediate between G. gymna and the fully platformed Desmoinesian gondolellid species such as G. bella. Only two other, somewhat minor, occurrences of naked gondolellids are known from rocks of pre-Missourian age. Gondolella cf. gymna occurs sparingly with the broad-platformed G. magna in the Lonsdale Member of northwestern Illinois (Merrill, 1975: 54) and in the Holdenville Formation of Jackson Co., Missouri. Missourian occurrences of naked species of Gondolella are confined to one species, G. denuda Ellison, from three nearly contemporaneous early Missourian occurrences, the Hushpuckney and Stark Shales of the Kansas City and Omaha areas, the Cramer Member of northern Illinois, and one specimen from the Lower Brush Creek Member of eastern Ohio. This last specimen was formerly interpreted to have been ancestral to (=Prioniodina camerata sensu Ellison, 1941), rather than conspecific with, G. denuda, and was omitted from the occurrences of species of Gondolella (Merrill, 1975:51). This is the first report of a species of Gondolella from the Appalachians. Species of Gondolella are known from many other units of Missourian age in the Midcontinent and Illinois, but those species, both older and younger, have broad platforms as do other species that occur with G. denuda in the Hushpuckney, Stark, and Cramer. No other species of Gondolella are known from 2 G. gymna G. cf.gymna G. denuda G. postdenuda G. neospathodiformis | Merrill&King | Merrill&King | Ellison sp. nov. | | Beil Mbr. | | | Kansas | | | | Shale over Salem School Mbr. Queen Hill Mbr. Texas Kansas Nebraska VIRGILIAN x Sy @ Heebner Mbr. | | N. Oklahoma | Salem School Mbr. Heebner Mbr. | Texas Kansas, Nebraska, Oklahoma Cramer Mbr. | | | | N. Illinois | | | | | | MISSOURIAN Brush Creek Mbr. | | Stark Mbr. . Ohio eswl eo W. Missouri, E. Nebraska el i | | | Hushpuckney Mbr. W. Missouri Lonsdale Mbr. Holdenville Fm. N.W. Illinois W. Missouri PENNSYLVANIAN | DESMOINESIAN eville Mbr. .W. Iilinois ATOKAN q—— Kodani Fm. Japan MORROWAN | | | | | | | | Fig. 1 Stratigraphic distribution of naked species of Gondolella. the Appalachian region other than the single platform (or Sp) element from the Lower Brush Creek. Virgilian naked gondolellids belong to two species, Gondolella postdenuda sp. nov. and Gondolella neospathodiformis sp. nov. The oldest Virgilian occurrence is in the Heebner Shale of Nebraska, Kansas, and Oklahoma, where both species are very rare. The slightly younger Queen Hill Shale of Nebraska and Kansas contains Gondolella postdenuda sp. nov. somewhat more abundantly and predictably, and contains broad-platformed species only very rarely (Ellison, 1941; Mendenhall, 1951). Gondolella postdenuda sp. nov. is also known from the lowest part of the Beil 3 Limestone at one locality in Kansas (as G. denuda in von Bitter, 1972) as well as from the Salem School Limestone and the shale directly overlying the latter in northern Texas (Fig. 1). Gondolella postdenuda sp. nov. and Gondolella neospathodiformis sp. nov. are the youngest species of Gondolella of which we are aware (with the possible exception of Clark’s 1972 report of specimens from the Upper Pennsylvanian and Lower Permian of Nevada that he called G. bella). The stratigraphic and geographic distribution of naked species of Gondolella is shown in Tables 1| and 2. Environmental Distribution and Faunal Associates The ecological requirements and biologic associates of species of Gondolella and the controversy surrounding their environmental setting (deep vs. shallow water) has been considered at length by Merrill and von Bitter (1976). With the possible exception of species of Diplognathodus, species of Gondolella, including the naked forms, are environmentally the most restricted of Pennsylvanian conodonts. Where found they commonly occur in profusion, but are absent from the greater part of the stratigraphic section. The most abundant and well-represented distribution in terms of the number and stratigraphic spacing of units bearing species of Gondolella occurs in the Kansas City Group (Missourian) of Kansas, Missouri, Nebraska, and Iowa. Naked gondolellids, in common with those species bearing broad platforms, generally occur in rocks containing significant numbers of elements of species of Idioprioniodus (Merrill and von Bitter, 1976). Gondolella generally does not occur in samples with substantial numbers of elements of species of Aethotaxis, although exceptions to these generalizations are known. Some of the occurrences of Gondolella gymna in the Seville Limestone are in rocks containing as many or more elements of species of Aethotaxis than of Idioprioniodus (Merrill and King, 1971). Some of the occurrences of G. denuda in the Cramer Member (Table 2) are in conjunction with uncharacteristically large numbers of specimens assignable to species of Aethotaxis, Cavusgnathus, and Ellisonia as well as to those of Idioprioniodus. In the Queen Hil! Shale and the base of the overlying Beil Limestone, G. postdenuda sp. nov. occurs without species of /dioprioniodus. The fact that von Bitter (1972) found only a few doubtful fragments of elements of species of Idioprioniodus as high stratigraphically as the Queen Hill, and Ellison (1941), von Bitter (1972), and Perlmutter (1975) failed to report any younger occurrences may be the result of disappearance (extinction?) rather than environmental exclusion. Although several apparently suitable stratigraphic units for both genera are present higher in the stratigraphic column, Gondolella and Idioprioniodus seem to disappear from the American Midcontinent at almost the same stratigraphic position. Species of the [diognathodus-Streptognathodus plexus are reduced or eliminated above this horizon, although both Gondolella and Idiognathodus occur higher in the western United States (Clark, 1972). 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Exceptions are dark, shaly limestones such as the Seville, light-coloured shales such as Lower Brush Creek, and clean light-coloured limestones such as the Lonsdale and Cramer (Tables 1 and 2). Analysis of these exceptions should provide additional insight into the ecology of Gondolella. Phylogenetic Methods and Philosophy Conodonts are skeletal parts that performed unknown functions within the body of animals whose taxonomic affinities may never be discovered. Despite this, conodont workers have found conodonts to be readily distinguishable from the remains of other groups such as those of fish and worms. Conodonts of Ordovician to Triassic age, taken collectively, appear to belong to a rather uniform, coherent group, a group that did not contain structural and physiological differences of the magnitude found between, for example, mammals and dinosaurs. Although there is evolutionary convergence within this group, this convergence is not between the taxa of major taxonomic categories such as those referred to above but is, as stated, within a taxonomically restricted and uniform group. The observed convergence in conodonts generally expresses itself as the reappearance of a morphologic feature after a long interval of time—of the duration of a geologic period or more. This observation and the fact that we are dealing in this study with a much shorter interval of time allow us to conclude that morphologically closely similar, but distinct, conodont platform elements are not only vaguely related but form a direct or indirect evolutionary ancestor—descendant relationship. This conclusion may also be applied, with some caution, to the nonplatform elements. Phylogenetic trees depend rather heavily on taxonomic decisions and it is necessary to outline what taxonomic characters have been used in the recognition of the various taxa. First and foremost when working with gondolellids is the requirement that the platform element possesses the definitive aboral loop. Naked gondolellids should, by definition, lack a broad platform on their Sp elements. If both these criteria are met and it has been established that we are dealing with naked gondolellids, then other criteria such as platform and denticle length, apparatus composition, fusion or lack of fusion of denticles, distribution of white matter, and the presence or absence of a lateral ridge are used to recognize different species of naked gondolellids. The other aspect of great importance in the construction of phylogenetic trees is where, in time and space, a particular fossil has been found. Any phylogenetic tree must be compatible with the stratigraphic positions, and consequently the ages, of the fossils on which it is based. In outlining the five phylogenetic alternatives (Figs. 2—6) we have carefully considered the stratigraphic occurrences and ranges of the various taxa (Fig. 1). The Phylogeny of Naked Species of Gondolella Two possibly interrelated aspects must be considered when attempting to trace the phylogeny of this group of conodonts. The first is the evolutionary relationship the 10 four naked species of Gondolella had to one another. Secondly, it is necessary to determine the evolutionary relationship of this group to the more common species of Gondolella that have broad platforms in their apparatuses. Both aspects are difficult to evaluate, not only because of the presence of thick sequences lacking any species of Gondolella, but also because of the scarcity of convincing intermediate forms. Considering the first question, the simplest phylogeny is shown in Fig. 2. It is based on the premise that morphological similarity implies or denotes relationship and that because the naked platforms of each of these four species are more similar to each other than they are to broad-platformed species of Gondolella, then they are in fact directly phylogenetically related. The above is the simplest and most attractive phylogeny and may ultimately prove to be the one that comes closest to representing the evolutionary history of naked species of Gondolella during the Pennsylvanian. It does, however, suffer from a number of shortcomings. (a) Sp elements of Gondolella cf. gymna from the Lonsdale Member and the Holdenville Shale are morphologically similar to those of G. gymna. Indeed, if these elements were found in the Seville Member they would be identified as the Sp elements of G. gymna; however, not only is the occurrence of Gondolella cf. gymna Stratigraphically a considerable distance from that of G. gymna, but it also occurs with abundant broad-platformed gondolellids. This fact makes it uncertain to which apparatus the associated ramiform elements (Fig. 8) belong and creates the (to us unlikely) possibility that the Gondolella cf. gymna Sp element is really an Oz element of a broad-platformed species of Gondolella, one that resembles the Sp element of G. gymna owing to possible convergence. There is considerable stratigraphic distance separating the highest occurrence of G. gymna from the lowest occurrence of G. denuda. There are, in this interval, a number of stratigraphic levels such as the Hanover, the Mineral Wells, the Sniabar, and the Hertha in which broad-platformed species of Gondolella occur. Although one would expect evolutionary forms intermediate to G. gymna and G. denuda to occur in environments that are suitable for other species of Gondolella, this is, for unknown reasons, not the case. (c) In considering the phylogenetic relationship between G. denuda and G. postdenuda sp. nov. one is struck by the close morphologic similarity between their platform elements as well as the similarity between their ramiform elements. There is, however, the same difficulty as in (b) in that there is a thick stratigraphic interval containing units such as the Quivira Shale, the Muncie Creek Shale, the Eudora Shale, and the LaSalle and Little Vermillion Limestones that contain abundant specimens of Gondolella with broad platforms but lack the evolutionary intermediate naked gondolellids that one would expect. Gondolella neospathodiformis sp. nov., if directly evolved from G. postdenuda sp. nov., represents a rather drastic shortening of the platform and enlargement of the main cusp. In addition, it is much smaller than are most specimens of G. postdenuda sp. nov. encountered and apparently lacked ramiform elements in its apparatus. (b — (d — A second possible phylogeny is outlined in Fig. 3. It has been established (Merrill, 1975: fig. 12) that Gondolella gymna, a species bearing a naked platform, gave rise to I] PERMO-TRIASSIC Neospathodus bransoni Muller ible tor 6 Ure Lowen ee Gondolella postdenuda sp. nov. VIRGILIAN Gondolella neospathodiformis Sp. NOV. . Broad-platformed & 6 ramiform > 2° Gondolellids? a go elements oe el Gondolella denuda MISSOURIAN Zz =! Z < > axl > o Z Z LL Oo Broad-platformed & 6 ramiform tgp Gondolellid elements Gondolella laevis Gondolella gymna ATOKAN/ DESMOINESIAN MISSISSIPPIAN pace Spathognathodus bultyncki Groessens Fig. 2 Evolution of naked species of Gondolella, the simplest and most parsimonious phylogenetic solution. 4 PERMO-TRIASSIC Broad-platformed Gondolellids I % Gondolella neospathodiformis sp. nov. VIRGILIAN & 6 ramiform Gondolella elegantula —» elements Gondolella postdenuda sp. nov. & 6 ramiform Gondolella symmetrica—> elements MISSOURIAN Gondolella denuda Z = Zz < > al > O Z Z LL oO Gondolella magna —p» & 6 ramiform elements ? Gondolella laevis Gondolella cf. gymna I Gondolella gymna & 6 ramiform elements ATOKAN/DESMOINESIAN MISSISSIPPIAN Fig. 3. Evolution of naked species of Gondolella from a stock of broad-platformed species by regressive platform development. i3 the more broad-platformed G. laevis as well as to younger Desmoinesian gondolellids. Although the phylogeny of younger broad-platformed species remains to be described the phylogeny of these broad-platformed species as at present understood is shown in Fig. 3. In this phylogeny there is no reason known to us why retrogressive (# reverse) evolution (i.e., broad platform to naked platform) could not have taken place a number of times in the Desmoinesian (G. magna to G. cf. gymna), in the Missourian (G. symmetrica to G. denuda), and in the Virgilian (G. elegantula, a species similar to G. sublanceolata, to G. postdenuda sp. nov.). In this phylogeny, the development of G. neospathodiformis sp. nov. was probably from G. postdenuda sp. nov. by direct evolution. Ellison (1942:121, pl. 21, figs. 1-4) illustrated transitional forms between G. denuda and G. symmetrica and interpreted the evolution as having proceeded in that direction. Our second suggested phylogeny is the reverse of that evolutionary pathway. Other transitions such as that illustrated by Ellison are not known, and until they are, this suggestion must be viewed as a weak phylogenetic contender. A third phylogenetic possibility for this group, one first suggested by Ellison (1941), is that a naked gondolellid species gave rise to all broad-platformed species (Fig. 4). Ellison (1941) suggested that G. denuda, the naked Missourian species, was ancestral to all gondolellid species with broad platforms, a suggestion that unfortunately neglected to explain the geologically older species with broad platforms that both Ellison, and Stauffer and Plummer (1932), had found. The transition from G. gymna to G. laevis described by Merrill (1975) however, does represent one such transition. Unfortunately, despite the fact that Merrill (1975) suggested a continuous lineage of naked gondolellids that gave rise to species with broad platforms several times, we at present lack proof of other instances of such iterative mosaic evolution in species of Gondolella with broad platforms. In the preceding we have been concerned with understanding and relating morphologic changes at different stratigraphic levels within the naked platform elements of a number of species of Gondolella. Ellison (1941) and Merrill and King (1971) noted morphologic transitions and intermediates between the conodont elements they called ‘‘camerata’’ and ‘*‘denuda’’ and “‘transitans’’ and *‘gymna’’, respectively. von Bitter (1972) considered that such pairs represent normal symmetry transitions between Sp and Oz elements of G. denuda and G. gymna, respectively, an opinion we follow herein. Nevertheless, it remains a possibility that ‘‘camerata’’ and ‘‘denuda’’ as well as “‘transitans’’ and *‘gymna’’ do represent homologous elements that expanded from the normal Oz to naked (Sp) platforms to broad (Sp) platforms in some parts of the generic range of Gondolella and not during others (Fig. 5). In such a phylogeny the two pairs indicated would form the first two-thirds of such a series. This remains an alternative, one that constitutes a fourth phylogenetic possibility. A fifth, and final, possibility (Fig. 6) explaining the distributions and morphologic intergradations that we observe in Pennsylvanian naked gondolellids is based on observations and ideas already touched upon: first; the morphologic plasticity exhibited by the Sp and Oz elements, i.e. they are morphologically similar; and second, the concept that the Sp and Oz elements may have been homologous elements that were part of a series consisting on one hand of Oz elements, and on the other of an intergrading series ranging from naked to broad platforms. In this postulated phylogeny two morphotypes, possibly sexual dimorphs (Jeppsson, 1972; Merrill and Merrill, 1974), existed concurrently (Fig. 6) throughout most of the ’ 14 PERMO-TRIASSIC VIRGILIAN Gondolella neospathodiformis sp. nov. Gondolella elegantula & 6 ramiform elements ? Gondolella postdenuda sp. nov. Gondolella sublanceolata Gondolella symmetrica IN & 6 ramiform elements ? MISSOURIAN Gondolella denuda Z S Z < > — > O Z Z LL Oo lell Gondolella magna & 6 ramiform elements ? Gondolella laevis ~ Gondolella cf. gymna & 6 ramiform elements ATOKAN/DESMOINESIAN Gondolella gymna MISSISSIPPIAN Fig. 4 Evolution of naked species of Gondolella by iterative mosaic evolution. PERMO-TRIASSIC Gondolella neospathodiformis sp. nov. Gondolella postdenuda sp. nov. eee» “postdenuda Oz” J "O02" ——————— Gondolella sublanceolata es Gondolella denuda VIRGILIAN za < z z l= - a = Slay ~ “ z = z Lu OW “camerata” "Oz" ——» Gondolella magna Ss ESS sirensiians id Gondolella gymna a ATOKAN/DESMOINESIAN MISSISSIPPIAN Fig. 5 Evolution of naked and broad-platformed species of Gondolella from a stock of Oz-like elements. 16 PERMO-TRIASSIC Gondolella neospathodiformis a ) Gondolella postdenuda Gondolella spp. BROAD -—-—-> Gondolella cf. merrilli Gondolella sublanceolata Gondolella denuda Z es Zz < > — > Y) Z Zz LL oO We. condoletta symmetrica MISSOURIAN VIRGILIAN Gondolella elegantula Gondolella magna BROAD = Gondolella bella Gondolella laevis ATOKAN/DESMOINESIAN Gondolella gymna MISSISSIPPIAN Fig. 6 Evolution of naked species of Gondolella. The platform elements are dimorphic and range from being naked to broad-platformed, while the Oz elements remain more or less constant in morphology through time. 17 Pennsylvanian. One morphotype of this pair (Fig. 6) bore a platform surrogate (the element that we refer to as an Oz element) and as many as five ramiform element types. The other morphotype (Fig. 6) contained the intergrading elements we refer to as the naked and broad-platformed Sp elements, again with as many as five ramiform element types. This phylogenetic possibility is based not only on the two observations and concepts already outlined but is also dependent on a number of other considerations. (a) Natural conodont assemblages present on black shale bedding planes from the Stark Shale of Nebraska show the presence of paired Oz elements but lack associated paired platform elements. Some samples, such as one from the Scammon Formation of western Missouri containing abundant well-preserved Sp elements of G. laevis, lack the typical Oz elements that we would have expected in the apparatus of this species. This is despite the fact that Oz elements of another genus, those of species of Idiognathodus, are rather abundant in this sample. (c) The presence of elements having great morphologic similarity in stratigraphic units such as Hushpuckney and Cramer Members of western Missouri and northern Illinois, respectively. More specifically, this refers to naked gondolel- liform elements (G. denuda) and Oz elements called Prioniodina ? camerata by Ellison. This point was already raised in the second phylogenetic alternative. (d) The apparatus of G. sublanceolata as reconstructed by von Bitter (1976) contains an element, the Hi element, that functionally could probably have played the part that the Oz element played in the apparatuses of species of Streptognathodus, Idiognathodus, and Cavusgnathus (von Bitter, 1972). The presence of two elements in the above reconstruction that could have taken on this role is anomalous—apparatuses of species of other genera do not have two morphologi- cally similar Oz-like elements. This, added to the fact that most platform-bearing apparatuses usually have a complement of six rather than seven distinct element types (Klapper and Philip, 1971:431-434) supports the possibility that the ‘‘camerata’’ element may in fact have functioned in a platform-surrogate role. (e) Even though Kozur (1976) makes the claim that Permo-Triassic species of Neospathodus have an apparatus closely similar in kinds of elements to those of species of Gondolella, a number of the species that have been placed in Neospathodus (and which we would consider to be Sp elements) strongly resemble the Oz elements of three of the Pennsylvanian naked gondolellid species. This may be support for the idea that these Pennsylvanian Oz elements were functionally Sp rather that Oz elements. This possibility is, of course, exceedingly difficult to demonstrate owing to the fact that even if such dimorphs existed their remains would normally be mixed together after death. If the existence of such dimorphism could be demonstrated by statistical or other means then this knowledge would have considerable impact on the taxonomy of Pennsylvanian gondolellids. For one thing, a number of broad-platformed and naked species would have to be drawn into synonymy. (b — A consideration of the phylogeny of naked gondolellids has, we believe, important implications for the phylogenies of conodonts both younger and older than those of Pennyslvanian age. Relationships between Gondolella and other genera are not known. By pointing out that this group of gondolellids, as distinct from a group of 18 species that more or less cluster about the type species, has some morphologic characters in common with other genera not only suggests possible phylogenetic relationships, but possible similar environmental adaptations as well. In discussing these faunas, nomenclature can, and does, often have an obscuring effect, and it should be noted that once the names are ignored conodont species having many of the characters of naked gondolellids have been found in rocks of Devonian, Mississippian, Permian, and Triassic age. Thus, some species of I/criodus, Pelekysgnathus, Eotaphrus, and Neospathodus share important morphologic characteristics with the naked gondolellids, particularly in the aboral loop and the possible presence in some species of Gondolella of multiple basal cavity tips. Indeed, Huddle (1934) placed the species Icriodus nodosa in Gondolella and Butler (1973) placed Spathognathodus bultyncki Groessens, a form probably ancestral to the naked gondolellids, in Pelekysgnathus. Spathognathodus bultyncki Groessens (Fig. 131-L) from the Tournaisian of Belgium has an Sp element morphologically closely comparable to the platform or Sp elements of Pennyslvanian naked gondolellids. Initially, Groessens (1971) included two morphotypes in Spathognathodus bultyncki—a platform-like element and a second element bearing a short posterior bar. It was subsequently found (Groessens, 1974) that these two elements have nearly mutually exclusive ranges in Belgium and that in the Lower Carboniferous of some areas, notably in the Canadian Rocky Mountains, the Oz-like element with the posterior bar occurs to the exclusion of the more platform-like element (S. Baxter, pers.comm., 1978). This resulted in two species, S. bultyncki and S. cf. bultyncki, being recognized in Belgium (Groessens, Conil, and Lees, 1973) whereas in the Canadian Mississippian S. Baxter (pers.comm., 1978) is planning to erect a new genus for the Oz-like forms. It is not known what, if any, ramiform elements were present in the apparatus of S. bultyncki. We initially thought that S. bultyncki and S. cf. bultyncki (sensu Groessens) could be part of the same apparatus. On the basis of the disparate ranges in Belgium (Groessen, 1974) and the nonoccurrence of the former in the Canadian Rocky Mountains (S. Baxter, pers.comm., 1978), we now believe that this probably represents the start of an evolutionary alternation in which an evolutionary pliable Oz-like element gave rise to a more Sp-like element. This alternation is nearly identical to that which we suggest might have taken place in Pennsylvanian naked gondolellids (Figs. 5, 6). Spathognathodus bultyncki has not been reported from North America, but S. cf. bultyncki (non sensu Groessens, 1971) (Fig. 13M-N, P-Q) from the Lower Windsor Group of Nova Scotia is very similar and occurs with an Oz element (Fig. 130) at the single locality from which it is known. It seems likely that these two elements represent the Sp and Oz elements of a single species. S. bultyncki Groessens is the best candidate for a Mississippian ancestor for Pennsylvanian naked gondolellids of which we are aware. Species placed in the Permo-Triassic genus Neospathodus Mosher (1968) share many characteristics with the Pennsylvanian naked gondolellids. Not only do they have the typical blade-like form that is not a true platform, but they also have the characteristic loop-like basal cavity. Indeed, there is some question, as is implied by the specific name, of whether Gondolella neospathodiformis sp. nov. could not be conveniently placed in what is at present recognized as a Permo-Triassic genus (Sweet in Ziegler, 1973). Some species of Neospathodus such as N. pakistanensis 19 and N. timorensis possess short posterior processes. We cannot help but wonder if such elements are homologous in their respective apparatuses to the Oz elements of the naked gondolellid species G. gymna, G. denuda, and G. postdenuda. Alternatively, it may be that these Oz-like elements are not platform surrogates and have taken on the function and position of the platform element as in the fourth and fifth phylogenetic possibilities already outlined. This may be a similar to identical expression of the morphologic (and possibly functional) plasticity and interchange- ability postulated between Sp and Oz elements of the naked gondolellids. All of the preceding taken together with Kozur’s (1976) claim that the apparatuses of species of Neospathodus and Gondolella are practically identical may be support for the idea that Pennsylvanian naked gondolellids and Permian and Triassic neospathodids not only are closely related but are part of the same phylogenetic lineage. Apparatus Element Composition von Bitter (1972, 1974, 1976) and Merrill (1975) concluded that species of Gondolella bore an apparatus consisting of not only a pair of platform (Sp) elements, but also paired and unpaired ramiform elements. von Bitter (1972) was able to partially reconstruct the apparatus of Gondolella denuda (G. postdenuda sp. nov. herein) and Merrill (1975) reconstructed most of the apparatus of Gondolella gymna Merrill and King. Since that time it has been possible to recognize the ramiform elements of G. postdenuda sp. nov. as well as those that probably belonged to G. denuda Ellison. The uncertainty about the ramiform elements of G. denuda stems from the fact that it occurs in the same samples with other, broad platform-bearing species of Gondolella and we cannot be sure which ramiform elements belonged with which Sp elements. Some of the ramiform elements almost certainly belonged to G. denuda, especially in samples where Sp elements of that species greatly outnumber the Sp elements of all other gondolellid species. Gondolella gymna, G. denuda, and G. postdenuda sp. nov. each have similar apparatuses, and the six kinds of ramiform elements cannot generally be distinguished from one species to the next. An exception to this is provided by the Oz elements of the three species. Each mimics the Sp element of the same species and can be distinguished in much the same way as can the respective Sp elements (see Systematic Palaeontology). In general, however, no matter which of the five phylogenetic paths outlined was followed, evolutionary change appears to have been restricted to the paired platform and ozarkodinid elements. Gondolella neos- pathodiformis sp. nov. is known only from Sp elements. The element composition of the naked species, exclusive of Gondolella neospathodiformis sp. nov., is directly homologous to that of species of Gondolella with broad platforms, as reconstructed for the representative species, G. sublanceolata, by von Bitter (1976). The six distinct kinds of ramiform elements present in G. sublanceolata (and probably other broad-platformed species of Gondolella as well) cannot at present be distinguished from those in the apparatuses of G. gymna, G. denuda, and G. postdenuda sp. nov. This lack of clearcut differences in ramiform element morphology of naked and broad platform-bearing species of Gondolella poses problems in apparatus reconstruction when dealing with samples containing both kinds. Fortunately, there are a number of 20 units and/or localities where their occurrence is mutually exclusive, or nearly so, and it is this fact that allows apparatus reconstruction to proceed. For example, although nearly 4000 platform and ramiform elements of the broad-platformed G. sublanceolata were recovered from six Pennsylvanian shale samples in western Iowa (von Bitter, 1976; von Bitter and Heckel, 1978) these same samples were totally devoid of the elements of naked gondolellids. The reverse situation has been found to be nearly always true at three localities of the Queen Hill Shale in Kansas and Nebraska (Fig. 1 and Table 2) and to be true in the Seville Member of northwestern Illinois (Fig. 1 and Table 1). The reasons for this mutual exclusiveness are either environmental and/or evolutionary in nature and were discussed in the section dealing with environmental distribution and faunal associates. The preceding discussion purposely avoids a consideration of the ratios of one element to another present in each of Gondolella gymna, G. denuda, and G. postdenuda sp. nov. It would be unwise to base any discussions regarding such ratios on the material available to us (Tables 1 and 2) because more substantial numerical data are required for such studies. Systematic Palaeontology Order Conodontophorida Eichenberg, 1930 Superfamily Gondolellacea Lindstrom, 1970 Family Gondolellidae Lindstrom, 1970 Genus Gondolella Stauffer and Plummer, 1932 Gondolella Stauffer and Plummer, 1932:41 Illinella Rhodes, 1952:898. Type Species Gondolella elegantula Stauffer and Plummer, 1932, by original designation. Gondolella gymna Merrill and King, 1971 Diagnosis A species containing paired Sp, Oz, Lo, Hi, Ne, and Syn elements as well as a probably unpaired bilaterally symmetrical Tr element. Sp element blade-like, narrow, lacking platform, but possessing a conspicuous lateral ridge about two-thirds to three-quarters the height of the blade above the aboral edge and extending the entire length of the anterior blade. A comparison of the Sp element of G. gymna with those of G. denuda and G. postdenuda sp. nov. is provided in Table 3. Sp, Oz, and Hi elements form a symmetry transition as do the Ne and Syn elements. Sp element (Fig. 7A-G, 1) Gondolella ? sp. A—Koike, 1967:302, pl. 1, figs. 29-32. 22 Gondolella gymna—Merrill and King, 1971:655, pl. 75, figs. 10-14. Gondolella gymna—Merrill, 1975:55, fig. 17, only numbers 53 and 55 are Sp elements of G. gymna. Oz element (Figs. 7H, J-L, 103J) Lonchodina sp. A—Koike, 1967:306, pl. 4, figs. 26-29. Gondolella gymna—Merrill and King, 1971:655, pl. 75, figs. 7-9. Lonchodina transitans—Merrill and King, 1971:658, pl. 75, figs. 15-18. Lo element (Figs. 7N, U, 10K) Gondolella gymna ‘‘Lonchodinid’’ element—Merrill, 1975:56, fig. 17, no. 51. Hi element (Fig. 7M, Q) Gondolella gymna ‘‘Lonchodinid-ozarkodinid’’ element—Merrill, 1975:56, fig. 17, no. 52. Ne element (Fig. 70, P) Syn element (Fig. 7S, T) Gondolella gymna ‘‘Synprioniodinid’’ element—Merrill, 1975:56, fig. 17, no. 54. Tr element (Figs. 7R, V, 10L) Fig. 7 A-V Gondolella gymna Merrill and King, Seville Member, Spoon Formation, Kewanee Group, Henry Co., Illinois; locality 14KSSS, sample 14C. A Sp element, lateral view of paratype. From Merrill and King (1971: pl. 75, fig. 10), USNM 165023, x34. Sp element, lateral view, ROM 38061, x68. Sp element, lateral view, ROM 38062, x70. Sp element, lateral view, enlargement of posterior end, ROM 38061, 139. Sp element, lateral view, ROM 38063, x55. Sp element, aboral-lateral view, ROM 38064, x42. Sp element, lateral view, enlargement of posterior end, ROM 38063, x110. Oz element, dextral, outer lateral view, ROM 38065, x75. Sp element, aboral view, enlargement of posterior end, ROM 38066, 125. Oz element, dextral, aboral view, enlargement of basal cavity and basal groove, ROM 38067, 119: Oz element, sinistral, outer lateral view, ROM 38068, 72. Oz element, sinistral, outer lateral view, ROM 38069, x37. Hi element, dextral, outer lateral view, ROM 38070, X87. Lo element, sinistral, outer lateral view, ROM 38071, x 107. Ne element, sinistral, inner lateral view, ROM 38072, «123. Ne element, dextral, inner lateral view, ROM 38073, 121. Hi element, sinistral, outer lateral view, ROM 38074, x75. Tr element, anterior view, ROM 38075, x 106. Syn element, dextral, inner lateral view, ROM 38076, x62. Syn element, sinistral, inner lateral view, ROM 38077, X88. Lo element, aboral, inner lateral view, ROM 38078, x114. Tr element, lateral view, ROM 38079, x62. Se ac yeaa! Sigh Moh). alee) 2a wo wo ZS nn nnn nnn nnn nner nnn nn nner SSS SuLleyy pue popunos Suey pue popunol AjIAeo [eseg suuelj pue popunol AjIAed [eseg AjjestnouuAs jou AWAvo [eseg AWABS [eseq 33Ie'] A}IABO [eseq o31e7] AWARD Jeseq [[ePWIS a3pll [esaje] poounouoid sso] a3pli [e19}e] poounouold ssa] a3pli [e19}e] psounouolg Suryore SWI0S Bulyose sWIOS Sulyoe apy] dsno 0} 10110)sod dsno 0} yuasaid oq Aew sapotuap juasaid Jol1a}sod juasaid 3q Aeu aJoNUSP BUG Jousa}sod ‘om) Ayares ‘ouQ uayjo ssac0id JoLIAa\sod [eay dsno jnojs ‘woYys dsno Zuo] A[qeoon0N dsnd jo y)3u9] ae1pouts9quy JAN [JUOD sso] aseq a[9IUEG JUIN[JUOS saseq s]ONUEG JWaNTJUOd saseq s[oNUNG SQ]DIUSP ISaMI,F Sa[sUSp snoJouINN sajouap JO JoquInu aJeIPsULI|}Uy sgjoUep passarduio0s sajouap possaidwios AT[e193e] saponuep possoidwios Aq[e19}e] ‘WOYS Ajjesoyey Ajqeaonou ‘ayesuolq Apysiys Ajuo ‘ayI]-3og sgjonUap JsauOYs sgjonuep suo] sgpouep wo0US ee a I ee ee “aou ‘ds ppnuapisod ‘5 jo yuauraya ds uosi||[q Bury pure [Lua ppnuap “5 jo juaulaja ds Duds “5 Jo Juauata dg Se ‘aou ‘ds ppnuapjsod ‘5 pue ‘vpnuap *D ‘vuudks vyajopuoy yo syuauiata dg ay) JO SdNSI1a}9e1eYD ay} JO UOSIAedUIOD ¢ JIqGUT 24 DISCUSSION Sp elements (Fig. 7A-G, I) of this species form a symmetry transition with the Oz element. This element, may, like the Oz element, have several denticles posterior to the cusp (Fig. 7F, 1) but is distinguished from the Oz element (Fig. 7H, J-L) by the presence of a closed aboral loop below the cusp (Fig. 71). The Oz element lacks a closed aboral loop and the basal groove extends into both the anterior and posterior blade (Fig. 7J). The posterior blade of the Oz element and the denticles posterior to the cusp of the Sp element are often missing, the result of structural weakness just posterior to the cusp. The resulting break generally is through the expanded basal cavity below the cusp and often makes it difficult, if not impossible, to state definitely with which of these two elements one is dealing. The nonplatform elements of Gondolella gymna (Fig. 7H, J-V) appear to be completely homologous in kinds of element types present with those of G. denuda, G. postdenuda sp. nov., and G. sublanceolata (von Bitter, 1976), the last a broad-platformed species. Although the ramiform elements of the three gondolellid species bearing naked Sp elements cannot be distinguished from one species to another, an exception to this observation is provided by the Oz element (Fig. 7H, J-L), the characteristics of which are tabulated in Table 4. The Oz element of G. gymna is differentiated from that of the younger G. denuda by the fact that it possesses fewer, shorter, and more-discrete denticles and by the presence of a strong lateral ridge along the length of the element. The presence of a strong lateral ridge in both the Sp and Oz elements of G. gymna (Fig. 7A, B, K, L) is another feature in which these two elements intergrade and mimic one another. With the exception of the Oz element, the ramiform elements of G. gymna are apparently indistinguishable from those of G. denuda and G. postdenuda sp. nov. Elements of G. gymna possess a microsculpture of both parallel and anastomosing ridges (Fig. 10J-L) on their cusps and denticles. MATERIAL Figured specimens ROM 38061 to 38079 inclusive; unfigured material ROM 38080 to 38084 inclusive. DISTRIBUTION Late Atokan and/or early Desmoinesian of Illinois; late Morrowan to early Atokan, Japan (Fig. 1 and Table 1). Gondolella cf. gymna Merrill and King, 1971 Sp element (Fig. 8A-C, 0) Gondolella gymna—Merrill, 1975:85, fig. 14, no. 22. ?Gondolella gymna—Merrill, 1975:85, fig. 16, numbers 38, 39 (probably = Oz elements of G. bella). 25 26 Possible ramiform elements of Gondolella cf. gymna. Oz element (Fig. 8D-F) ?Gondolella gymna—Merrill, 1975:85, figs. 14, 23. (Unable to determine from illustration whether complete aboral loop or posterior groove present.) Lo element (Fig. 8G, H, K, Q) Hi element (Fig. 81, L, P) Ne element (Fig. 8M) Syn element (Fig. 8J) Tr element (Fig. 8N) DISCUSSION Rare platform specimens (Fig. 8A-C) from the Lonsdale Member in northwestern Illinois and the Holdenville Formation of Jackson Co., Missouri bear a strong Fig. 8 A-C2O Ow > D-N, P-Q Gondolella cf. gymna Merrill and King, Sp element, Lonsdale Member, Modesto Formation, McLeansboro Group, Peoria Co., Illinois. Lateral view, locality 2AMGL, sample 2C, RoM 38085, x59. Aboral view, locality 7AMGL, sample 7D, ROM 38086, x 106. Aboral view, enlargement of aboral view showing basal cavity, locality 7AMGL, sample 7D, ROM 38086, 209. Detail view of striae on first denticle posterior to the cusp, locality 2AMGL, sample 2C, ROM 38085, x902. ?Ramiform elements of Gondolella cf. gymna Merrill and King, Lonsdale Member, Modesto Formation, McLeansboro Group, Peoria Co., Illinois. Oz element, dextral, outer lateral view, locality 2AMGL, sample 2C, RoM 38087, x55. Oz element, sinistral, outer lateral view, locality 2AMGL, sample 2C, ROM 38088, x99. Oz element, sinistral, outer lateral view, locality 7AMGL, sample 7D, ROM 38089, x 106. Lo element, sinistral, outer lateral view, locality 7AMGL, sample 7D, RoM 38090, «114. Lo element, sinistral, outer lateral view, locality 2AMGL, sample 2C, ROM 38091, tt 0 39 PsP. uct By — Yo stnsmm Pigs © sey 40 ETYMOLOGY With reference to the morphology of this species which is suggestive of species of Neospathodus Mosher. MATERIAL Figured specimens ROM 38150 (holotype), ROM 38151 to 38154 inclusive (paratypes); unfigured paratypes ROM 38155, 38156, 38163, and 38164. DISTRIBUTION Lower Virgilian, Oklahoma. (See Table 2.) Fig. 13. A-H S(O 2 A. = Gondolella neospathodiformis sp. nov., Heebner Shale, Oread Formation, Shawnee Group, Osage Co., Oklahoma, locality He-7. Sp element, enlarged lateral view of cusp showing striae, holotype, sample He-7-1, ROM 38150, x 792. Sp element, lateral view, holotype, sample He-7-1, RoM 38150, x 154. Sp element, aboral-lateral view, paratype, sample He-7-1, ROM 38151, 204. Sp element, aboral-lateral view, paratype, sample He-7-1, Rom 38152, 207. Sp element, aboral view, paratype, sample He-7-Rec., ROM 38153, x264. Sp element, enlarged aboral view of basal cavity, paratype, sample He-7 Rec., ROM 38153, «506. Sp element, aboral view, paratype, sample He-7-1, ROM 38154, x273. Sp element, enlarged aboral view of basal cavity, sample He-7-1, ROM 38154, x 539. Spathognathodus bultyncki Groessens, Tournaisian, Salet, Belgium, sample B(6) 92-95, ROM Acc. No. 74PB24. Sp element, aboral view, ROM 38157, x92. Sp element, enlarged aboral view of basal cavity, ROM 38157, 231. Sp element, lateral view, ROM 38158, x46. Sp element, enlarged lateral view of posterior end, ROM 38158, x92. Spathognathodus cf. bultyncki Groessens, Viséan, Windsor Group, Wentworth Formation of Moore and Ryan (1976), Phillips Limestone (Moore, in Geldsetzer et al..1980), Miller’s Creek Quarry, Hants Co., Nova Scotia, sample Phil-1-2, ROM Acc. No. 75PB29. Sp element, lateral-aboral view, ROM 38160, x79. Sp element, lateral-aboral view, ROM 38161, x44. Sp element, enlarged lateral-aboral view of posterior end, ROM 38161, X114. Sp element, enlarged lateral-aboral view of posterior end, ROM 38160, x 183. Oz element, (?) of Spathognathodus cf. bultyncki Groessens, Viséan, Windsor Group, Wentworth Formation of Moore and Ryan (1976), Phillips Limestone (Moore, in Geldsetzer et al., 1980), Miller’s Creek Quarry, Hants Co., Nova Scotia, sample Phil-1-1, Rom Acc. No. 75PB28, ROM 38162, X87. 4] Acknowledgements We are indebted to the following members of the Department of Invertebrate Palaeontology, Royal Ontario Museum, for their help and co-operation: Mrs. Hilary Geberl and Ms Maureen Wende who assisted in the preparation of the figures; Miss Joan Burke who typed the numerous drafts and did much of the preliminary editing; and Mrs. LaVerne Russell who, some years ago, did much of the preparation and mounting of specimens and who was the first to locate and recognize the new species G. neospathodiformis. Acknowledgement is made for the use of the scanning electron microscope in the University of Toronto, established through a grant from the National Research Council to the Department of Zoology, University of Toronto, for the development of a programme in systematic and evolutionary zoology. We are grateful to Mr. George Gomolka and Mr. Eric Lin of the University of Toronto, for their expertise in the operation of the Cambridge scanning electron microscopes. We thank Messrs. Allan McColl and Brian Boyle of the Photography Department, Royal Ontario Museum, for their assistance in the preparation of figures. Mr. Ted White, Omaha, Nebraska, generously made available natural assemblage specimens of Gondolella denuda Ellison for study and deposit at the Royal Ontario Museum. 42 Literature Cited BRANSON, E.B. 1944 The geology of Missouri. University of Missouri Studies 19: 1-535. BUTLER, M. 1973 Lower Carboniferous conodont faunas from the eastern Mendips, England. Palaeontology 16:477-5S17. CLARK, D.L. 1972 Early Permian crisis and its bearing on Permo-Triassic conodont taxonomy. Geologica et Palaeontologica 1:147-158. CLARK, D.L. and L.C. MOSHER 1966 Stratigraphic, geographic, and evolutionary development of the conodont genus Gondole!- la. Journal of Paleontology 40:376-394. COOLEY, D.R. 1952 Facies change in the Oread Limestone in southern Kansas and northern Oklahoma. M.Sc. Thesis, University of Kansas. 61 pp. ELLISON, S.P., Jr. 1941 Revision of the Pennsylvanian conodonts. Journal of Paleontology 15:107-143. EVANS, J.K. 1966 Depositional environment of a Pennsylvanian black shale (Heebner) in Kansas and adjacent states. Ph.D. Thesis, Rice University. 166 pp. GELDSETZER, H.H.J., P. GILES, R. MOORE, and W. PALMER 1980 Stratigraphy, sedimentology, and mineralization of the Carboniferous Windsor Group, Nova Scotia. Joint Annual Meeting, Geological Association of Canada/Mineralogical Association of Canada, Halifax, 1980, Field Trip 22 Guidebook. 42 pp. GROESSENS, E. 1971 Les conodontes du Tournaisien supérieur de la Belgique; note préliminaire. Service Géologique de Belgique, Professional Paper 4: 1-29. 1974 Preliminary range chart of conodont biozonation in the Belgian Dinantian. International Symposium on Belgian Micropaleontological Limits, Namur, Publication 17:1-193. GROESSENS, E., R. CONIL, and A. LEES 1973 Problemes relatifs a la limite du Tournaisien et du Viséen en Belgique. Société Belge de Géologie, Bulletin 82: 17—SO. GUNNELL, F.H. 1933 Conodonts and fish remains from the Cherokee, Kansas City, and Wabaunsee groups of Missouri and Kansas. Journal of Paleontology 7:261—297. HUDDLE, J.W. 1934 Conodonts from the New Albany Shale of Indiana. Bulletins of American Paleontology 21(72): 1-137. JEPPSSON, L. 1972 Some Silurian conodont apparatuses and possible conodont dimorphism. Geologica et Palaeontologica 6:51-69. KLAPPER, G. and G.M. PHILIP 1971 Devonian conodont apparatuses and their vicarious skeletal elements. Lethaia 4:429-452. 43 KOIKE, T. 1967 A Carboniferous succession of conodont faunas from the Atetsu Limestone in southwest Japan (Studies of Asiatic conodonts, Part VI). Tokyo Kyoiku Daigaku Science Reports, Section C, 93:279-318. KOZUR, H. 1976 Paleoecology of Triassic conodonts and its bearing on multielement taxonomy. Geological Association of Canada, Special Paper 15:312-324. LEE, W. 1938 Stratigraphy of the Cisco Group of the Brazos Basin. University of Texas, Publication 3801: 11-90. MENDENHALL, M.E. 1951 Conodonts and fish remains of the Douglas and Shawnee Groups of the Virgil Series (Pennsylvanian) of Nebraska. M.Sc. Thesis, University of Nebraska. 78 pp. MERRILL, G.K. 1975 Pennsylvanian conodont biostratigraphy and paleoecology of northwestern Illinois. Geolog- ical Society of America, Microform Publication 3: 1-130. MERRILL, G.K. and C.W. KING 1971 Platform conodonts from the lowest Pennsylvanian rocks of northwestern Illinois. Journal of Paleontology 45:645-—664. MERRILL, G.K. and S.M. MERRILL 1974 Pennsylvanian nonplatform conodonts, Ila: The dimorphic apparatus of /dioprioniodus. Geologica et Palaeontologica 8:119-130. MERRILL, G.K. and P.H. von BITTER 1976 Revision of conodont biofacies nomenclature and interpretations of environmental controls in Pennsylvanian rocks of eastern and central North America. Royal Ontario Museum, Life Sciences Contributions 108:1—-46. MOORE, R.G. and R.J. RYAN 1976 Guide to the invertebrate fauna of the Windsor Group in Atlantic Canada. Nova Scotia Dept. of Mines, Paper 76-5: 1-57. MOSHER, L.C. 1968 Triassic conodonts from western North America and Europe and their correlation. Journal of Paleontology 42:895—946. PERLMUTTER, B. 1975 Conodonts from the uppermost Wabaunsee Group (Pennsylvanian) and the Admire and Council Grove Groups (Permian) in Kansas. Geologica et Palaeontologica 9:95-109. STAUFFER, C.F. and H.J. PLUMMER 1932 Texas Pennsylvanian conodonts and their stratigraphic relations. University of Texas, Bulletin 3201: 13—S0. TOOMEY, D.F. 1969 The biota of the Pennsylvanian (Virgilian) Leavenworth Limestone, midcontinent region; part I, stratigraphy, paleogeography, and sediment facies relationships. Journal of Paleontology 43:1001-1018. 44 von BITTER, P.H. 1972 Environmental control of conodont distribution in the Shawnee Group (Upper Pennsylvanian) of eastern Kansas. University of Kansas Paleontological Contributions 59: 1-105. 1974 Element composition and micromorphology of species of Gondolella from the Middle and Upper Pennsylvanian of the central U.S.A. Geological Society of America, Abstracts with Programs 6:551. [Abstract]. 1976 The apparatus of Gondolella sublanceolata Gunnell (Conodontophorida, Upper Pennsylva- nian) and its relationship to //linella typica Rhodes. Royal Ontario Museum, Life Sciences Contributions 109: 1-44. von BITTER, P.H. and P.H. HECKEL 1978 Differentiation of black ‘‘core’’ shales in Missourian and Virgilian cyclothems (Pennsylva- nian) in Iowa and Kansas, using conodonts. Geological Society of America, Abstracts with Programs 10:510. [Abstract]. von BITTER, P.H. and G.K. MERRILL 1977 Neogondolelliform conodonts of Early and Middle Pennsylvanian age. Royal Ontario Museum, Life Science Occasional Paper 29: 1-12. ZIEGLER, W., ed. 1973 Catalogue of conodonts, volume 1. Stuttgart, E. 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