UNh RSITY OF ILLItxUIS LIBRARY AT URBANA-CHAMPAIGN NATURAL HIST. SURVEY FIELDIANA • GEOLOGY Published by CHICAGO NATURAL HISTORY MUSEUM Volume 10 June 12, 1952 No. 13 A CLASSIFICATION OF THE CONULARIDA G. Winston Sinclair Department of Geology, University of Michigan ABSTRACT The seventeen genera of the Conulariidae are grouped into three new subfamilies. Six new genera and two new species are described. INTRODUCTION Although more than two hundred species of Conularia have been described, ranging in age from Tremadocian to Triassic, until 1939 no attempt was made to express their mutual relationships in terms of subfamily groupings. In that year Boucek (1939, p. A120 et seq.) formalized a classification that he had previously proposed (1928) for his Bohemian material. An emendation of this classi- fication has more recently been published by Sugiyama (1942, p. 397). In the course of a monographic study of the group I have become convinced that these attempts at classification, based mainly on surface ornamentation and the structure of the midline of the face of the shell, are inadequate to express our knowledge of the diverse forms that are found. Publication of the morphological detail on which my opinion rests must await the appearance of the monograph, but it is desirable now to set out some of my taxonomic conclusions, so that they may be used in the forthcoming Treatise on Invertebrate Paleontology. In brief, I find the most diagnostic features of the conularid shell to be the corner of the pyramid, and the nature of the structures found there. The character of the midline of the faces is of secondary importance, and the surface ornamentation is of very minor use, although to some extent it is correlated with more important features. The scheme published by Boucek in Schindewolf's Handbuch der Palaozoologie is readily available, and it will be sufficient to summarize the classification given there without comment. The "Gruppen" refer to Boucek' s informal classification of 1928. No. 692 135 136 FIELDIANA: GEOLOGY, VOLUME 10 CLASSIFICATION ACCORDING TO BOUCEK Order? Conularida Miller and Gurley 1896 Family Conularidae (sic) Walcott 1886 Genus Conularia Miller (sic) 1818 Subgenus Archaeoconularia Boucek 1939 (Gruppen der Conularia fecunda und insignis) Subgenus Mesoconularia Boucek 1939 (Gruppe der Conularia fragilis) Subgenus Conularia Miller Subgenus Plectoconularia Boucek 1939 (Gruppe der Conularia proteica) Genus Metaconularia Foerste 1928 (Gruppen der C. exquisita und solitaria) Genus Pseudoconularia Bou6ek 1939 (Gruppe der Conularia grandissima) Family Conulariellidae Kiderlen 1937 Genus Conulariella Boucek 1928 Family Serpulitidae Boucek 1939 Genus Serpulites Murchison 1839 As I pointed out in 1940 (Sinclair, 1940, p. 73), this treatment of Conularia s. str. is based on a misinterpretation of the genotype. Bou5ek's Plectoconularia is Conularia s. str., and his Conularia s. str. is Paraconularia. CLASSIFICATION ACCORDING TO SUGIYAMA Family I. Conularidae (sic) Walcott 1886 Genus 1. Conularia Miller (sic) 1821 Subgenus 1. Archaeoconularia Boucek 1939 Subgenus 2. Mesoconularia Boucek 1939 Subgenus 3. Conularia (s. str.) Miller Subgenus 4. Paraconularia Sinclair 1940 Subgenus 5. Plectoconularia Boucek 1939 Genus 2. Metaconularia Foerste 1928 Genus 3. Pseudoconularia Boucek 1939 Family II. Conulariellidae Kiderlen 1937, emend. Genus 4. Conulariella Bou5ek 1928 Genus 5. Conchopeltis Walcott 1875 Genus 6. Neoconularia Sugiyama 1942 Family III. Conulariopsisdae (sic) Sugiyama 1942 Genus 7. Conulariopsis Sugiyama 1942 Family IV. Serpulitidae Boucek 1939 Genus 8. Serpulites Murchison 1839 Sugiyama's paper is not generally available in the United States, so some comments may be in order. I am greatly indebted to Mr. C. G. Robertson of Tokyo for his kindness in providing me with a photostat of the paper. The inclusion of both Paraconularia and Plectoconularia as subgenera of Conularia must have been based on a misreading of SINCLAIR: CLASSIFICATION OF CONULARIDA 137 my paper (1940a). The type species of Conularia must belong to one or the other of these groups, but not both. Conchopeltis (Knight, 1937, p. 186) may be a conularid, but it has nothing in common with Conulariella, its broad spreading shell contrasting strikingly with the thin lamellar shape of the latter genus. I do not know where to put it, but Conchopeltis does not belong in the Conulariellidae. Neoconularia is a new genus (Sugiyama gives the date as 1941, but this is an error) based on Conularia rectangularis Hayasaka 1920. Unfortunately, this species is known only from steinkerns, and none of the descriptions (Hayasaka, 1920, p. 87; 1924, p. Ill; Sugiyama, 1942, p. 391) give details of the nature of the corner of the shell, or of the facial ornament. One may say from the descriptions that the species has nothing in common with Conulariella, but its correct position cannot be stated with certainty. I place it in the Para- conulariinae mainly because that subfamily is better represented in the Upper Paleozoic than the others. It will not be possible to refer other species to this genus until more information on the type species is available. Sugiyama's Conulariopsis (for which he gives the date 1938, although this refers only to an oral presentation of the description to a meeting of the Palaeontological Society of Japan) is based on a new species, Conulariopsis quadrata, from the Lower Triassic. It is characterized by a lack of corner grooves, the corners of the pyramid being raised; by coarse sinuous longitudinal ridges that apparently corrugate the surface of the shell; and by a very irregular series of widely spaced transverse folds that appear in the photograph as if they could be adventitious. Conulariopsis is possibly a conularid, but it should be placed in a separate family. PROPOSED CLASSIFICATION Order ?Conularida Miller and Gurley 1896 ?Family Conulariopsidae Sugiyama 1942 Family Conulariellidae Boucek 1939 Genus Conulariella Boucek 1928 Family Conulariidae Walcott 1886 Subfamily Conulariinae, subfam. nov. Conularids with the corners of the shell simply inflected, not interrupting the surface ornamentation, and with no strengthening rods or other structures. Genus Conularia Sowerby 1820. Type, Conularia quadrisulcata Sowerby 1820. Genus Mesoconularia Boucek 1939. Type, Conularia fragilis Barrande 1867 (see Boucek, 1928, p. 84). Genus Metaconularia Foerste 1928. Type, Conularia aspersa Lindstrom 1884 (see Sinclair, 1940a, p. 101 et seq.). 138 FIELDIANA: GEOLOGY, VOLUME 10 Genus Pseudoconularia Boucek 1939. Type, Conularia grandissima Barrande 1867 (see Sinclair, 1941, p. 125 et seq.). Genus Archaeoconularia Bou5ek 1939. Type, Conularia insignis Barrande 1867 (see Sinclair, 1944, p. 87 et seq.). Genus Palaenigma Walcott 1886. Type, Tetradium wrangeli Schmidt 1874 (see Schmidt, 1874, p. 42 et seq.; Sinclair, 1944, p. 87). Genus Diconularia gen. nov. Type, Conularia micronema Meek 1871. Genus Exoconularia gen. nov. Type, Conularia exquisita Barrande 1867. Genus Anaconularia gen. nov. Type, Conularia anomala Barrande 1867. Subfamily Paraconulariinae, subfam. nov. Conularids with the corners of the shell sharply inflected; the ornamentation interrupted there, the transverse elements either stopping or alternating. Genus Paraconularia Sinclair 1940. Type, Conularia inaequicostata de Koninck 1883. Genus Calloconularia gen. nov. Type, Calloconularia strimplei, sp. nov. Genus Eoconularia Sinclair 1944. Type, Conularia loculata Wiman 1894 (see Sinclair, 1944, p. 90). Genus ?Neoconularia Sugiyama 1942. Type, Conularia rectangularis Hayasaka 1920. Subfamily Ctenoconulariinae, subfam. nov. Conularids with the corners of the shell strengthened with internal carinae or other types of thickening. Genus Ctenoconularia gen. nov. Type, Ctenoconularia obex, sp. nov. Genus Climacoconus Sinclair 1942. Type, Conularia quadrata Walcott 1879 (see Sinclair, 1942, p. 226). Genus Glyptoconularia gen. nov. Type, Conularia gracilis Hall 1847. Genus Conularina Sinclair 1942. Type, Conularia triangulata Raymond 1905 (see Sinclair, 1942, p. 226). DESCRIPTIONS OF GENERA AND SPECIES Conulariinae, subfam. nov. Diconularia, gen. nov. Conularia auctt. (in part). Mesoconularia Boucek and others (in part). Type species. — Conularia micronema Meek 1871 (see Meek, 1875, p. 316). Diagnosis. — Conulariinae with closely appressed transverse ridges on the faces, set with prominent subcircular pustules; the corners of the shell simple, the midline of the faces unmarked on the sur- face, but with a tendency to be raised on the inner side. Remarks. — In 1940 I followed Boucek in taking Mesoconularia to be an inclusive genus, with rather wide limits. I now feel that this is unwise, and I restrict Mesoconularia to the few forms that closely resemble the type species, M. fragilis. There are, especially in the Upper Paleozoic, a number of species forming a group that has rather obviously developed from Conularia, species of that genus SINCLAIR: CLASSIFICATION OF CONULARIDA 139 often showing as gerontic characters the crowding of the transverse ridges and accentuation of the pustules that are normal in Diconu- laria. In view of this, it seems probable that Diconularia arose several times, and should be considered a form-genus until the relationships of the species can be worked out more thoroughly. The genus ranges from the Middle Ordovician to the PennsyU vanian. Exoconularia, gen. nov. Conularia auctt. (in part). Metaconularia Boucek and others (in part). Type species. — Conularia exquisita Barrande 1867 (see Boucek, 1928, p. 70). Diagnosis. — Conulariinae with accessory longitudinal lines along the faces, varying in position in different species; the surface orna- mented with transverse rows of discrete pustules. Remarks. — This is the group of species that I left as "Metaconu- laria?" in 1940. The relationships are still not as clear as I would like to see them, but these species are clearly separable from Meta- conularia s. str., with its pair of strong internal carinae at the mid- line, and I place them in this new genus. I suspect that Exoconularia itself needs subdivision, but I do not feel that my knowledge of the European species warrants that step at the moment. The genus is exclusively Ordovician, occurring from the Tre- madocian to the Caradocian. Anaconularia, gen. nov. Conularia auctt. (in part). Metaconularia Boucek (in part). Type species. — Conularia anomala Barrande 1867 (see Barrande, 1867, p. 33). Diagnosis. — Conulariinae with no trace of a corner groove, but instead with the corner of the shell elevated into a sharp or rounded ridge. Remarks. — The abundant specimens of this species are all stein- kerns, but the lack of the corner groove is not due to that fact, since in all other conularids the presence of the groove is emphasized, rather than obscured, on the steinkern. Similarly, the apparent lack of any superficial ornamentation cannot be attributed to pres- 140 FIELDIANA: GEOLOGY, VOLUME 10 ervation, since in the same beds occur steinkerns of other species with the fine surface markings well impressed either on the specimen itself or on its external impression. The only species occurs in the Middle Ordovician Drabov quartites (d?) at Drabov and other localities in Bohemia. Barrande had over 1,500 specimens when he drew up his description of the species. Of these, the specimen figured by him (pi. 7, figs. 10 and 11) may be designated as lectotype. It is presumed to be in the National Museum in Prague. Paraconulariinae, subfam. nov. Calloconularia, gen. nov. Conularia Girty (in part). Type species. — Calloconularia strimplei, sp. nov. Diagnosis. — Paraconulariinae with low crowded transverse ridges and, in the interspaces, longitudinal bars. There exists in the Upper Paleozoic of the United States a group of very similar shells that should be separated from Paraconularia. They are always small, and the surface markings are remarkably uniform. At the shoulders of the corner groove the interspaces swell strongly, so that they continue into the groove, while the transverse ridges stop at the shoulders. This feature is also seen in some species now referred to Paraconularia, for example, P. victa (White), so that it is not diagnostic for Calloconularia. I know six species in the Pennsylvanian of Oklahoma and Texas. The only described form is that named Conularia crustula holden- villae by Girty in 1911 (see Girty, 1915, p. 44). Calloconularia strimplei,1 sp. nov. Figure 56, D-F. Horizon and locality. — Pennsylvanian, Ochilate group, an un- named shale 30 feet above the Torpedo sandstone. Two miles north-northeast of Capan, Oklahoma. Types. — C.N.H.M. No. PE 142, holotype; five paratypes, No. PE 143. Harrell L. Strimple, collector. Gift of G. W. Sinclair. Description. — Shell small, but of the usual size for the genus. Length of holotype 15 mm., greatest width of face 4 mm.; shell 1 Named for Mr. Harrell L. Strimple, well known for his work on the Echino- dermata. SINCLAIR: CLASSIFICATION OF CONULARIDA 141 not quite straight, tapering regularly or a little more rapidly apically, apical angle about 15°. Section rhombic, almost square. Faces plane or a little concave, equal; the midline indicated only by the ornamentation or by a very slight ridge apically. Corner with a rather wide, shallow groove, the bottom angulate and filled with the ends of the interspaces, the shoulders broadly but sharply rounded. Apex not known. The apertural lobes have been seen in some speci- mens, but show no real details. The holotype figured is complete at the apertural end, but the lobes are either lacking or strongly infolded, so that they are not seen. Surface with rather low, but prominent, flat-topped transverse ridges, eight in a length of 1 mm. apically, seven and one-half and six in the same length measured successively nearer the aperture. The ridges are smooth. They alternate at the corners, but have no regularity of position at the midline. Their direction can be seen in the figures. The interspaces are from one to one and one-half times as wide as the ridges, and are marked by very irregular longi- tudinal wrinkles. They swell sharply near the corner, so that by the time they reach the shoulder of the groove they are higher than the ridges, and continue past them into the bottom of the groove, where they alternate with those from the adjacent face. Ctenoconulariinae, subfam. nov. Ctenoconularia, gen. nov. Conularia auctt. (in part). Type species. — Ctenoconularia obex, sp. nov. Description. — Ctenoconulariinae with strikingly slender shells. The corner of the shell appears on the surface as a wide shallow groove, on the interior as a strongly carinate ridge. The midline of the face is marked by an interruption of the transverse ridges, but the internal structure is not known except in one species. The transverse ridges that "ornament" the faces are biconvex in section, that is, they extend past the surface both externally and internally, so that the steinkern appears grooved when the test is removed. The ridges stop at the shoulder of the corner grooves. They are pustulose, and the pustules usually continue into the next orad interspace as projections of greater or lesser length. This results at times in an appearance very similar to that of Conularia s. str., but in that genus the longitudinal bars that occur in the interspaces are always continuous with, or opposite, pustules on the next ridge orad, and 142 FIELDIANA: GEOLOGY, VOLUME 10 thus have a very different origin. Neither the apex nor the aperture has been seen in any specimen. No apical schott1 has been seen. Remarks. — The genus ranges from the Black River (Middle Ordovician) to the Hamilton (Middle Devonian). Sixteen species are known to me, of which Conularia crebristria Hall 1876 (Hall, 1879, p. 210) and Conularia trentonensis occidentalis Bradley (1930, p. 242) are previously described American forms. Ctenoconularia obex,2 sp. nov. Figure 56, A-C. Horizon and locality. — Middle Ordovician, Sprechs Ferry member of the Platteville formation. Minneapolis, Minnesota. Types. — University of Minnesota collections, holotype No. 6608; three paratypes Nos. 6609-6611; F. W. Sardeson, collector. Description of holotype. — Shell rather small, 27 mm. long; face 5 mm. wide, straight, tapering regularly and very slowly, apical angle about 6°. Section square. Faces plane, equal, the midline marked only by the course of the ornamentation. Corner with a rather wide shallow groove, the shoulders broadly rounded, the bottom broadly rounded and smooth. Surface with narrow, rather low, rounded, transverse ridges, six in a length of 2 mm. at the smaller end of the specimen, eight in the same length near the middle, and seven near the larger end; the crests marked by well-separated longitudinal elevations that extend very slightly into the next orad interspace but are somewhat longer near the corners; about ten to twelve in a width of 1 mm. The transverse ridges are almost all continuous at the midline and op- posite at the corners. They are slightly curved, well arched across the face, the angle at the midline about 135°. Interspaces two to three times as wide as the ridges, smooth except for the projections mentioned above. Remarks. — A much larger paratype, with facial width at least 11 mm., has seven transverse ridges in a length of 5 mm., set with seven longitudinal elevations in a width of 1 mm. 1 The transverse structure sealing off the apex in many species of Conulariinae (but apparently never in Paraconulariinae or Ctenoconulariinae) has been referred to by authors as a septum, a diaphragm, or in other ways. These terms suggest an undesirable homology with structures in other animals, and I have therefore used the noncommittal German word Schott (plural Schotten) for this peculiarly conularid structure. 2 Derivation of name: Latin obex, a bolt. Fig. 56. A-C. Ctenoconularia obex, sp. nov. Holotype; X%lA and X7. Univ. Minnesota coll. No. 6608. Ordovician, Sprechs Ferry member of Platteville, Minneapolis, Minnesota. D-F. Calloconularia strimplei, sp. nov. Holotype; X5. C.N.H.M. PE 142. Pennsylvanian, Ochilate group, Capan, Oklahoma. 143 144 FIELDIANA: GEOLOGY, VOLUME 10 Glyptoconularia, gen. nov. Conularia auctt. (in part). Genotype. — Conularia gracilis Hall 1847 (see Hall, 1847, p. 224). Description. — Ctenoconulariinae with strong corner strengthen- ings, a pronounced mesial carina on the interior at the midline of the face, and strong longitudinal surface ornamentation. Remarks. — I find it difficult to determine what weight to give the unique surface ornamentation of Conularia gracilis. But for the surface, this species might be referred to Eoconularia. At the moment it seems simplest to erect a monotypic genus. It should be noted that the young "Conularia gracilis" described by Ruedemann (1896, 1898) and copied by so many textbooks, is not a conularid. It belongs to the genus Tubelelloides, which Howell has recently erected (1949, p. 4) for the worm-tubes long known as Serpulites (see Ruedemann, 1916, p. 85). REFERENCES Barrande, Joachim 1867. Systeme silurien du centre de la Boheme — I. 3, Pteropodes, xv+179 pp., 16 pis. Prague. Boucek, Bedrich 1928. Revision des Conulaires paleozoi'ques de la Boheme. Palaeontographica Bohemiae, Nr. XI, 108 pp., 7 pis. 1939. Conularida, in O. H. Schindewolf, Handbuch der Palaozoologie, Bd. 2A, pp. A113-131, figs. Bradley, J. H., Jr. 1930. Fauna of the Kimmswick Limestone of Missouri and Illinois. Walker Mus. (Univ. Chicago), Contr., 2, (6), pp. 219-290, pis. 23-30. Girty, G. H. 1915. Fauna of the Wewoka formation of Oklahoma. U. S. Geol. Surv., Bull. 544, 353 pp., 35 pis. Hall, James 1847. [Descriptions of organic remains of lower division of New York system.] Palaeontology of New York, 1, 338 pp., 99 pis. 1879. [Descriptions of Gasteropoda, Pteropoda and Cephalopoda of Upper Helderberg, Hamilton, Portage and Chemung groups.] Geol. Surv. New York, Palaeontology, 5, pt. 2, 492 pp., 120 pis. Hayasaka, Ichiro 1920. A new species of Conularia from southern Kitakami, Japan. Geol. Soc. Tokyo, Jour., 27, (327), pp. 87-90, figs. 1924. Some Permian fossils from the Kitakami Mountains. Japanese Jour. Geol. and Geogr., 2, (4), pp. 107-116, pi. 15. SINCLAIR: CLASSIFICATION OF CONULARIDA 145 Howell, B. F. 1949. New hydrozoan and brachiopod and new genus of worms from the Ordovician Schenectady formation of New York. Wagner Free Inst. Sci. (Philadelphia), Bull. 24, (1), pp. 1-10, 2 pis. Knight, J. B. 1937. Conchopeltis Walcott, an Ordovician genus of the Conulariida. Jour. Paleontology, 11, (3), pp. 186-188, pi. 29. Meek, F. B. 1875. A report on some of the invertebrate fossils of the Waverly group and coal measures of Ohio. Ohio Geol. Surv., Rept., 2, pt. 2, pp. 269-347, pis. 10-20. RUEDEMANN, RUDOLF 1896. Note on the discovery of a sessile Conularia. Amer. Geol., 17, (3), pp. 158-165; 18, (2), pp. 65-71, pi. 2. Reprinted, with additions, in Rept. of New York State Geol., 15, pp. 701-720, 4 pis., 1898. 1916. Account of some new or little-known species of fossils. New York State Mus., Bull. 189, pp. 7-97, pis. 1-30, figs. 1-33. Schmidt, Friederich 1874. Miscellanea silurica — II. Uber einige neue und wenig bekannte Baltisch- Silurische Petrefacten. Acad. Imp. Sci. St. Petersbourg, Mem., (7), 21, (11), 48 pp., 4 pis. Sinclair, G. W. 1940a. The genotype of Conularia. Canadian Field-Nat., 54, (5), pp. 72-74. 1940b. A discussion of the genus Metaconularia, with descriptions of new species. Roy. Soc. Canada, Trans., (3), 34, sec. IV, pp. 101-121, 3 pis. 1941. Notes on Pseudoconularia and P. magnifica (Spencer). Roy. Soc. Canada, Trans., (3), 35, sec. IV, pp. 125-129, pi. 1942. The Chazy Conularida and their congeners. Ann. Carnegie Mus., 29, (10), pp. 219-240, 3 pis. 1944. Notes on the genera Archaeoconularia and Eoconularia. Roy. Soc. Canada, Trans., (3), 38, sec. IV, pp. 87-95, pi. SUGIYAMA, TOSHIO 1942. Studies on the Japanese Conularida. Geol. Soc. Japan, Jour., 49, pp. 390-399, pi. 15.