COLOR PATTERNS ON PENNSYLVANIAN GASTROPODS FROM OHIO1

R. D. HO ARE, Department of Geology, Bowling Green State Univ., Bowling Green, OH 43403 M. T. STURGEON, Department of Geology, Ohio University, Athens, OH 45701

Abstract. Gastropods are among the more common fossil forms with preserved color patterns. Numerous species of Carboniferous gastropods have been previously re- ported in Europe and North America including 9 species from the Pennsylvania!! of Ohio. Our collections of Pennsylvanian gastropods, made over the past several years, contain specimens representing 11 species with remnants of color patterns, 9 not previously reported from Ohio. The color patterns of 4 of these species, Ian- thinopsis intercalaris? (Meek & Worthen), Palaeostylus (Paleostylus) n. sp., Palaeo- siylus {Pseudozygo pleura) n. sp. 1 and Palaeostylus {Pseudozy go pleura) n. sp. 2, have not been described before. Comparisons were made between Ohio gastropods with color patterns, or closely related forms, and those reported elsewhere. Variation in patterns were noted and illustrated where possible. OHIO J. SCI. 78(1): 3, 1978

Preserved color patterns in fossils have tropod species previously reported from been intermittently reported since Bru- the Pennsylvanian strata of Ohio have guiere (1792) illustrated specimens of the been described by Sturgeon (1964), bivalve Gryphaea, with radiating brown Bellerophon (Pharkidonotus) percarinatus bands, from the Cretaceous of Ireland. (Conrad) and Euconospira turbiniformis Over 200 reports describing or illustrating (Meek & Worthen); Lee (1971), Eucono- color patterns on fossils have been pub- spira turbiniformis (Meek & Worthen); lished since that date, a large number of Morningstar (1922), Naticopsis {Marmo- which pertain to gastropods, one of latella) pulchella n. sp.; Webb (1972), the more common groups of organisms Euconospira riddelli (Shumard), Lepto- in which patterns are preserved. tygma virgatum (Knight), Bellerophon {Pharkidonotus) percarinatus (Conrad), Paleozoic examples of this type of Bellerophon {Bellerophon) graphicus preservation are not as uncommon as Moore, Knightes {Cymatospira) sp., and might be expected. Newton (1907) gives Ianthinopsis paludinaeformis (Hall); and, a summation of European occurrences of Hoare and Sturgeon (1976), Baylea sp. color pattern preservation, noting 13 [ = Callistadia n. sp.]. Paleozoic species of cephalopods, pele- Continued collecting of Pennsylvanian cypods and gastropods, three of which are invertebrates in the Ohio Pennsylvanian gastropods of Carboniferous age. Foerste by the authors and their students has un- (1930), in a summary paper of known oc- covered specimens representing 11 spe- currences of cephalopods, brachiopods, cies with preserved color patterns, 9 of gastropods and bivalves with preserved which have not been previously reported color, lists 44 Paleozoic species of gastro- from Ohio, and 4 species, Ianthinopsis pods, with 33 being of Carboniferous age. intercalaris? (Meek & Worthen), Paleo- Since 1930, specimens of at least 22 stylus {Paleo stylus) n. sp., Paleo stylus Paleozoic species of gastropods, with {Pseudozy go pleura) n. sp. 1 and P. color patterns preserved, have been de- {Pseudozy go pleura) n. sp. 2 that have not scribed in the literature; 11 are from the been previously described with preserved Pennsylvanian of North America. Gas- patterns. In several cases these 11 spe- manuscript received May 25, 1977 (#77-42). cies were represented by single specimens R. D. HOARE AND M. T. STURGEON Vol. 78 but there were several instances where cord ll/i Quadrangle. Specimens col- lected from the shale on the spoil several specimens were present, providing banks. information as to the variability of the Jmi-2. The Vanport Limestone and Shale ex- color pattern within the species. Varia- posed at a small abandoned drift mine tion from previously described patterns on the north side of Buffler Run, S cen- ter sec. 24, Milton Township, Jackson for a species was noted and is listed under County, Ohio. Mulga 7J^' Quadrangle. the color pattern descriptions. Specimens were collected from the 0.5 The figures were produced by drawings foot shale zone just above the Vanport made directly from specimens or from Limestone. LIuc-1. The Lower Mercer Limestone exposed photographs of specimens. Most speci- at an undescribed and uncertain lo- mens were incomplete or distorted. Some cality near Flint Ridge, Ohio. generalized reconstruction was necessary Mp-2. The Vanport Limestone exposed at the to indicate the general form or whorl pro- Carbon Limestone Co. quarries, 0.75 file of specimens but when major portions mi. south of U.S. 224 and just west of the Ohio-Pennsylvania state line near of the shell were missing, they were not Kansas Corners, Poland Township, added. Only the basic shell form and Mahoning County, Ohio. Campbell major features of ornamentation have 73^2' Quadrangle. Specimens collected been indicated so that the color pattern, from weathered limestone blocks. No-4. The Portersville Shale exposed in road- indicated by stippling, is not obscured. cut on the east side of U.S. 21, SEJ4 The pattern represented in the drawings SE^ sec. 10, Olive Township, Noble is as preserved and no attempt was made County, Ohio. Caldwcll South 7^' to reconstruct a total pattern. Descrip- Quadrangle. Specimens collected from shale exposure. tions and illustrations of the new species Pr-3. The Lower Mercer Limestone exposed indicated herein are being published in the B & O RR cut, N\V}| sec. 10, at separately. Repositories for the illus- Somerset, Reading Township, Perry trated specimens are given in the figure County, Ohio. Somerset i^/i Quad- rangle. Specimens collected from legends as: The Ohio State Univer- weathered blocks of limestone. sity, OSU; Ohio University, OU; Bowling Ty-3. The Putnam Hill Limestone and Shale Green State University, BGSU. exposed in a small abandoned borrow pit and strip mine of the Zoar Mining COLLECTING LOCALITIES Co., N\\T14 sec. 17, south of New Cp-3. The Columbiana Limestone at the Philadelphia, York Township, Tusca- rawas County, Ohio. New Phila- abandoned Brookwood Mine (type ] locality for the Columbiana), SE^4 delphia 7}4 "Quadrangle. Specimens NWM sec. 29, Perry Township, Co- collected from a 1-2 foot shale zone lumbiana County, Ohio. Salem 7^' just above the Putnam Hill Limestone. Quadrangle. Specimens collected from the mine dump. DISCUSSION OF COLOR PATTERNS CS1-1. The Lower Mercer Limestone and The preservation of color patterns on Shale exposed in a roadcut on the fossil specimens commonly shows varia- north side of U.S. 36, 1.0 mi. east of tion within a species. In some instances jet. with Ohio 93, NE^ sec. 1, approx. this variation probably is caused by in- 2.0 mi. northeast of West Lafayette, Lafayette Township, Coshocton complete preservation of the pattern on County, Ohio. Fresno 7J^' Quad- a particular specimen. In other cases rangle. Specimens collected from a the variation appears to be biologically 1.0 foot shale zone just above the controlled. It is doubtful that any of Lower Mercer Limestone. these specimens show true coloration, as Ga-1. The Cambridge Limestone and Shale exposed in a small abandoned quarry, present during life, but only remnants of WJ^SE^ sec. 25, approx. 1.0 mi. north- the original color as a result of preserva- east of New Concord, Adams Town- tion vagarities. Preservation may ac- ship, Guernsey County, Ohio. Bloom- field iyPermian of Texas as does Knight (1934) spiralling dark band on the base of the in specimens of this genus from the whorl. The band is centrally located Pennsylvanian of Missouri. and occupies one-third to one-half the Callistadia n. sp. A population of 177 whorl width. In several instances the specimens from the Putnam Hill Shale at band is interrupted to form a series of locality Ty-3 contains 120 specimens patches or dashes as it approaches the showing color patterns. The pattern aperture (figs. 9 and 11). Some speci- consists of transverse sinuous dark gray mens also show a dark narrow band ex- bands of variable width and spacing tending from just below the seienizone to which cross the ramp from the upper the upper margin of the seienizone and suture and extend onto the lateral sur- may have a ramp completely colored, al- face where they are interrupted at a con- though more lightly than the band at the cave zone before extending onto the whorl seienizone, except for the band of nodes base (figs. 12-14). Variability in pattern adjacent to the upper suture (fig. 8). R. D. HOARE AND M. T. STURGEON Vol. 78

14

FIGURES 1 and 2. Bellerophon (Bellerophon) graphicus Moore. Dorsal and lateral views, x2.6, Cambridge Shale, Gwe-1, OU-29498. FIGURES 3-5. Bellerophon (Pharkidonotus) percarinatus (Conrad). 3, 4, dorsal and lateral views, x2.6, Cambridge Shale, Ga-1, BGSU-4163; 5, oblique dorsal view, x3.5, Columbiana Limestone, Cp-3, OSU-19612. Specimen illustrated in figures 3 and 4 illustrates reversed light-dark banding as opposed to specimen in figure 5. FIGURE 6. Euconospira turbiniformis (Meek & Worthen). Lateral view of a partial whorl, xl.3, Vanport Limestone, Mp-2, OSU-19764. FIGURE 7. Straparolus (Amphiscapha) cattiloides (Conrad). Oblique view, x2.6, Cambridge Shale, Ga-1, BGSU-4164. FIGURES 8-11. Glabrocingulum {Glabrocingulum) grayvillense (Norwood & Pratten). Figures 8 and 9, lateral and basal views, x4.0, BGSU-4165; Figures 10 and 11, lateral and basal views, x4.0, BGSU-4166 Cambridge Shale, Ga-1. Views of two specimens showing variation in pat- terns, in part probably due to preservation differences. FIGURES 12-14. Callistadia n. sp. Lateral views of three specimens showing variation in pat- tern, x20.0, Putnam Hill Shale, Ty-3, OSU-30390, 30388, 30382. Ohio I. Sci. FOSSIL GASTROPOD COLOR PATTERNS Other specimens show little or no dif- (fig. 17). The rest of the shell has a very ference in color in the area of the seleni- light color. Knight (1930) describes zone or on the ramp surfaces (fig. 10). numerous color patterns in pseudozygo- Naticopsis (M armolatella) pulchella pleurids from the Pennsylvanian of Mis- (Morningstar). One specimen from the souri. Paleostylus (Pseudozy go pleura) Lower Mercer Limestone at locality Pr-3, tenuivirga Knight has a pattern of one described by Morningstar (1922) and Lee narrow dark band located at the sutural (1971), is still available for study. The line which closely approaches that of this observable pattern consists of a series of species from Ohio. revolving bands with three dark brown Palaeostylus (Pseudozy go pleura) n. sp. bands separated by lighter brown bands. 2. About 200 specimens were collected A light band is present just below the from the Cambridge Shale at locality upper suture followed by a dark band of Ga-1. A few specimens show retention approximately equal width, a narrower of a color pattern which consists of a light band, a second wider dark band, a broad revolving light brown band at second wider light band and then a third midwhorl bordered by narrower light very narrow dark band just above mid- bands between it and the upper and lower whorl. The lower part of the whorl ap- sutures (fig. 16). A broad light brown pears to be light (fig. 15). Most of the revolving band may also be present on body whorl is crushed and extensions of the base of the whorls. Knight (1930) the pattern into this area cannot be does not describe any pattern for species determined. Both Morningstar (1922) of the subgenus Pseudozy go pleura which and Webb (1972) note only the two matches that of the specimens described broad upper dark bands. above. He does indicate that some spe- Knight (1933) reillustrated Morning- cies of the subgenus Paleostylus have a star's specimen without description but similar pattern in the Pennsylvanian of the narrow third (lower) dark band can Missouri. be faintly seen in his illustration (PI. 44, Palaeostylus (Palaeostylus) n. sp. Ap- fig. 3). He also notes revolving bands on proximately 100 specimens were collected neanic specimens of Naticopsis wortheni from the Cambridge Shale at locality Knight and N. virgata Knight from the Ga-1. Many of the specimens show a Pennsylvanian of Missouri but the spac- preserved pattern consisting of a rela- ing and distribution of the revolving tively broad brown revolving band lo- bands is different in both of these species cated at mid-whorl and bordered by as well as being different from N. (M.) narrower light bands which extend to pulchella. Squires (1976) describes and the upper and lower sutures respectively illustrates ephebic specimens of N. (Nati- (fig. 18). Knight (1930) describes four copsis) wortheniana Knight from the species of this subgenus from the Penn- Pennsylvanian Buckhorn asphalt in Okla- sylvanian of Missouri with color patterns. homa which have a pattern of revolving The pattern of P. (P.) wortheni Knight dark bands separated by light-colored seems to be most similar to that of this bands, the pattern, in terms of band species from Ohio. width and spacing, showing some vari- Leptoptygma virgatum (Knight). Webb ability and differ from N. (M.) pulchella (1972) described and illustrated speci- in being wider and extending much lower mens from the Cambridge Shale and on the whorl profile. Limestone from locality Gwe-1 with a Palaeostylus (Pseudozy go pleura) n. sp. preserved pattern which consists of two 1. Several hundred specimens collected dark spiralling bands separated by a cen- from the Cambridge Shale at locality trally located light-colored band. One Ga-1 show a color pattern. The pre- dark band borders the upper suture and served pattern consists of one narrow re- is more prominent, the other extends volving band of grayish-blue to dark above the lower suture and may be par- brown or reddish-brown color located at tially to completely covered by succeed- the sutural line so that it is commonly ing whorls. The bands increase in width exposed above the lower suture but towards the aperture (fig. 19). Knight partially covered by the succeeding whorl (1931) described and illustrated speci- R. D. HOARE AND M. T. STURGEON Vol. 78 mens of this species from the Pennsyl- the aperture before ending well above vanian of Missouri with a dark nucleus mid-whorl. They appear again below (protoconch) and two dark revolving midwhorl as chevron-shaped bands ter- bands. One dark band lies below the minating above the umbilical area (fig. upper suture and is separated from it by 21). Knight (1931) describes specimens a narrower light band and the second of this species from the Pennsylvania!! dark band extends slightly above the of Missouri as having a uniform dark lower suture but is mainly covered by gray to black coloration. succeeding whorls. Ianthinopsis paludinaeformis (Hall). Ianthinopsis intercalaris^ (Meek & Webb (1972) described and illustrated Worthen). One large specimen, col- specimens from the Cambridge Shale at lected from the Cambridge Shale at locality Gwe-1. More than a dozen locality Ga-1, shows transverse bands specimens subsequently collected from of dark brown pigmentation which start the Cambridge Shale at localities Ga-1 at or just below the upper suture and and Gwe-1, and two specimens from the more or less parallel the growth lines on Porterville Shale at locality No-4, show the shell. The bands vary in width and retention of color patterns. A consider- spacing with some apparently interrupted able amount of variation exists in the on the central portion of the body whorl, pattern although the pattern is consist- while others are continuous across the ently composed of revolving bands. A whorl. Some bands extend into the um- relatively narrow light or dark band, bilical area (fig. 20). more commonly the former color, occurs Ianthinopsis primogenius (Conrad). just below the upper suture. Below this One specimen, collected from the Cam- is either a narrow band of the opposite bridge Shale at locality Ga-1, shows a shade or the rest of the whorl surface is color pattern consisting of transverse of the opposite shade. On some speci- dark grayish-blue bands on the body mens the second band may move up to whorl which vary in width and spacing. the upper suture in latter whorls. Oc- The bands begin at the upper suture and casionally three bands are present with a curve away from and then back towards narrow upper dark band, a narrow in-

EXPLANATION OF FIGURES 15-28 FIGURE 15. Naticopsis (Marmolatella) pulchella (Morningstar). Oblique view of holotype, x3.3, Lower Mercer Limestone, Pr-3, OSU-15311. Revolving dark bands cannot be seen on later portion of body whorl because of preservation. FIGURE 16. Palaeostylus (Pseudozy'gopleura) n. sp. 2. Lateral view of specimen without a sec- ond band on the base of the whorl, xlO.O, Cambridge Shale, Ga-1, BGSU-4167. FIGURE 17. Palaeostylus (Pseudozy go pleura) n. sp. 1. Lateral view, x6.7, Cambridge Shale, Ga-1, BGSU-4168. FIGURE 18. Palaeostylus (Paleostylus) n. sp. Lateral view, xlO.O, Cambridge Shale, Ga-1, BGSU-4169. FIGURE 19. Leptoptygma virgatum (Knight). Lateral view showing only a small indication of the lower band which may or may not appear above the lower suture, x4.0, Cambridge Shale, Gwe-1, OU-29435. FIGURE 20. Ianthinopsis intercalaris? (Meek & Worthen). Lateral view, x2.0, Cambridge Shale, Ga-1, BGSU-4170. FIGURE 21. Ianthinopsis primogenius (Conrad). Lateral view, x3.0, Cambridge Shale, Ga-1, BGSU-4171. FIGURES 22-26. Ianthinopsis paludinaeformis (Hall). Lateral views of five specimens showing pattern variation; Figure 22, x.66, Portersville Limestone, No-4, OU-29436; Figure 23, x2.6, Cambridge Shale, Gwe-1, OU-29437; Figure 24, x2.6, Cambridge Shale, Gwe-1, BGSU-4172; Figure 25, x2.6, Cambridge Shale, Gwe-1, BGSU-4173; Figure 26, x2.6, Cambridge Shale, Ga-1, BGSU-4174. FIGURE 27. Meekospira peracuta (Meek & Worthen). Lateral view, x4.7, Cambridge Shale, Ga-1, BGSU-4175. FIGURE 28. Soleniscus typicus (Meek & Worthen). Lateral view, x2.0, Cambridge Shale, Ga-1, BGSU-4176. Ohio J. Sci. FOSSIL GASTROPOD COLOR PATTERNS termediate light band and the entire suture and a second light gray band of lower two-thirds of the whorl being a equal width is just above the lower suture. third band of dark color (figs. 22-26). The intervening band is light brown in Knight (1931) describes specimens of color and is slightly wider than either of this species from the Pennsylvanian of the light bands and fades out on the Missouri as being uniformly dark gray to body whorl (fig. 27). Knight (1932) de- black. scribes specimens of this species from the Meekospira peracuta (Meek & Wor- Pennsylvanian of Missouri as having a then). One specimen, collected from the narrow dark band below the upper suture Cambridge Shale at locality Ga-1, retains followed by an intermediate light band a color pattern. A narrow light gray and a lower dark band which is bisected band is present just below the upper by the lower suture. Apparently the

28 10 R. D. HOARE AND M. T. STURGEON Vol. 78 patterns of the specimens from Ohio and Hoare, R. D. and M. T. Sturgeon 1976 Color pattern variation in a Pennsylvanian gastro- Missouri are the same with the preserved pod. Geol. Soc. Amer. Abs. Progr. 8: 482- dark-light color relationship of the bands 483. being reversed. Kemp, A. H. 1957 Color retention in Steno- Soleniscus typicus (Meek & Worthen). poceras, Euomphalus, and Naticopsis from the Lower Permian of central Texas. J. One specimen, collected from the Cam- Paleont. 31: 974-976. bridge Shale at locality Ga-1, has a pre- Knight, J. B. 1930 The gastropods of the St. served pattern. The pattern, which is Louis, Missouri, Pennsylvanian outlier: The faint, consists of a broad light brown re- Pseudozygopleurinae. J. Paleont. 4(Suppl. volving band extending from the lower 1): 1-88. 1931 The gastropods of the St. Louis suture, or below it, to near the upper Missouri, Pennsylvanian outlier: The Su- suture from which it is separated by a bulitidae. J. Paleont. 5: 177-229. narrow light band (fig. 28). The body 1932 The gastropods of the St. Louis, whorl shows the dark band extending to Missouri, Pennsylvanian outlier: The Pseudo- melaniidae. J. Paleont. 6: 189-202. the abapical end of the shell. Knight 1933 The gastropods of the St. Louis, (1931) describes the color pattern of this Missouri, Pennsylvanian outlier: The Neri- species, from specimens in the Penn- tidae. J. Paleont. 7: 359-392. sylvanian of Missouri, as being a uniform 1934 The gastropods of the St. Louis, Missouri, Pennsylvanian outlier: The Euom- dark gray to almost black. phalidae and Platyceratidae. J. Paleont. Acknowledgments. We wish to acknowledge 8: 139-166. the help of students and friends in making col- Lee, Chun-sun 1971 Pottsville gastropods of lections out of which the specimens described Ohio. Unpubl. M.S. thesis, Ohio Univ., here were obtained. Anthony Zalinski and Athens, OH. 129 pp. David Atwater, while graduate students at Morningstar, Helen 1922 Pottsville fauna of Bowling Green State University, found several Ohio. Geol. Survey of Ohio, 4th ser., Bull. of the specimens during their thesis field work No. 25, 312 pp. and brought them to our attention. The Newton, R. B. 1907 Relics of coloration in Faculty Research Committee of Bowling Green fossil shells. Proc. Malacological Soc. Lon- State University provided a grant to support don 7: 280-292. field work. Squires, Richard L. 1976 Color pattern of Naticopsis {Naticopsis) wortheniana, Buck- LITERATURE CITED horn asphalt deposit, Oklahoma. J. Paleont. Bruguiere, J. C. 1792 Encyclopedie Methodi- 50: 349-350. que; histoire naturelle des vers. Panckouke Sturgeon, Myron T. 1964 Allegheny fossil (Paris) 1:85-132. invertebrates from eastern Ohio—Gastro- Foerste, A. F. 1930 The color patterns of poda. J. Paleont. 38: 189-226. fossil cephalopods and brachiopods, with Webb, Douglas 1972 Conemaugh fossil in- notes on gastropods and pelecypods. Con- vertebrates from eastern Ohio—Gastropoda. trib. Mus. Paleont. Univ. Michigan 3: 109- Unpubl. M.S. thesis, Ohio Univ., Athens, OH. 149. 147 pp.