Col Our Pat Tern of Naticopsis Planispira (Neritimorpha, Gastropoda) Shell from up Per Car Bon If Er Ous of up Per Silesian Coal Ba Sin, South Ern Poland

Annales Societatis Geologorum Poloniae (2013), vol. 83: 87–97.

COL OUR PAT TERN OF NATICOPSIS PLANISPIRA (NERITIMORPHA, GASTROPODA) SHELL FROM UP PER CAR BON IF ER OUS OF UP PER SILESIAN COAL BA SIN, SOUTH ERN POLAND

Wojciech KRAWCZYÑSKI

Depart ment of Palaeon tol ogy and Stratig ra phy, Faculty of Earth Sciences, Univer sity of Silesia, Bêdziñska 60, 41-200 Sosnowiec, Poland; e-mail: [email protected]

Krawczyñski, W., 2013. Col our pat tern of Naticopsis planispira (Neritimorpha, Gastropoda) shell from Upper Car - bon if er ous of Upper Silesian Coal Ba sin, south ern Po land. Annales Societatis Geologorum Poloniae, 83: 87–97.

Ab stract: A zig zag col our pat tern with ad di tional collabral belts and ir reg ular spots was ob served on a shell of Naticopsis (N.) planispira (Phillips, 1836) found in the Go³onóg Sandstone marine fau nal ho ri zon (Namurian A, the Up per Silesian Coal Ba sin, Poland). A dis ruption in the col our pat tern was also ob served to have occurred during ontogenetic shell growth. In this spec i men, it ap peared in the healed shell damage on the outer lip. So far, only nine Car bon if er ous spe cies of Naticopsis have been re ported with the col our pat tern pre served. The most common are the zig zag-type and the spi ral band-type. In di vid ual taxa clearly dif fer in the mor phol ogy of the chev rons (direc tion and an gles of breaks) and the lo ca tion and width of the spiral bands on a whorl. However, the color pat terns may not be di ag nostic fea tures for the Naticopsis species, because of the large intraspecific varia tion and col our pat tern poly morphism on the neritimorph shells. Col our pat terns on Palaeozoic neritimorph shells most likely served as cam ou flage with re spect to the bot tom surface in the photic zone.

Key words: Gastropoda, Neritimorpha, col our pat tern, Car bon if er ous, Po land.

Manu script re ceived 12 May 2013, ac cepted 26 August 2013

IN TRO DUC TION The pres ent pa per de scribes a Carbon if er ous gas tro pod shell of Naticopsis (Naticopsis) planispira (Phillips, 1836) Spec i mens of Palaeozoic ma rine in ver te brates with col - with a preserved colour pat tern, from the Go³onóg Sand - our pat terns pre served on their shells or cara paces are rare. stone marine faunal hori zon. The diverse faunal assem - They were described from the shells of brachio pods (e.g., blage, consist ing of about 50 taxa (in cluding bi valves, gas- Køíž and Lukeš, 1974; Blodgett et al., 1988; Baliñski, 2010), tropods, brachio pods, and nautiloids), has been studied by bivalves (e.g., Hoare et al., 1988; Mapes and Benstock, Cramer (1910), and Weigner (1938) and was revised by 1988), nautiloids (e.g., Manda and Turek, 2009; Turek, Bojkowski (1972). Addi tional faunal research was con- 2009), and also from trilo bite cara paces (e.g., McRoberts et ducted by Schwarzbach (1935), Pøibyl (1951), and Salamon al., 2013). (1997). Among the ten rec og nized gastro pod taxa, Naticop- Colour patterns on Palaeozoic mollusc shells, includ ing sis planispira has not been listed previ ously. gas tro pods, are pre served rel a tively sel dom, ow ing to the frequent recrystallization of the orig inal aragon ite shells. The oldest gastro pods with preserved colour patterns (Stra- GEO LOG I CAL SET TING parollina harpa) have been found in the Middle Ordo vician depos its of the USA (Raymond, 1906; Foerste, 1930). Indi - The Up per Silesian Coal Ba sin is mainly lo cated in the vidual finds have also been recorded from the Silu rian (e.g., southern part of Poland and only its most SW part stretches Køíž and Lukeš, 1974), De vo nian (e.g., Yochelson and into the terri tory of the Czech Re pub lic (Fig. 1B). The Up- Køíž, 1974; Rohr and Smith, 1978; Frýda, 2000; Basse and per Carbon if er ous coal-bearing series rests on the Lower Heidelberger, 2002; Jankovský, 2003) and Permian depos - Car bon if er ous mudstone-sand stone de pos its (Culm fa cies) its (e.g., Yochelson, 1956; Batten, 1958, 1972; Plas, 1972). in the W part and limestones (Carbon ifer ous limestone fa- Their great est di ver sity, how ever, is noted in Car bon if er ous cies) in its east ern part. The Upper Carbon ifer ous depos its forma tions (e.g., Knight, 1932, 1933; Batten, 1966; Hoare of the Up per Silesian Coal Ba sin con sist of the Paralic Se - and Sturgeon, 1978; see also Kobluk and Mapes, 1989). ries (the lower part of the Coal-bear ing Se ries) in its lower 88 W. KRAWCZYÑSKI

Fig. 1. Lo ca tion of out crops and strati graphi cal po sition of the Go³onóg Sand stone ma rine fau nal ho ri zon. A. Stratigraphic sequence of the Up per Car bon if er ous in the NE part of the Up per Silesian Coal Ba sin, showing po si tion of the Go³onóg Sand stone ma rine fau nal ho ri - zon (af ter Kotas, 1995); Z. B. – Zabrze Beds. USSS – the Up per Silesian Sand stone Se ries. B. Lo ca tion and geo log i cal fea tures of the Up - per Silesian Coal Basin (af ter Jureczka et al., 1995). C. Geo log i cal map of the NE part of D¹browa Górnicza with sample lo ca tion of Naticopsis shell (see as ter isk) at the Go³onóg Sand stone ho ri zon out crop (af ter Doktorowicz-Hrebnicki, 1954) COL OUR PAT TERN OF Naticopsis planispira 89

part with nu merous marine faunal hori zons (Namurian A). Above is the Limnic Series with the Up per Silesian Sand- stone Se ries (Zabrze Beds and Ruda Beds; Namurian B and C) and the Mudstone Se ries (the Za³ê¿e Beds and the Orze- sze Beds; Westphalian A and B). The Kraków Sandstone Series (Westphalian C and D), which is present only in the eastern part of the basin, forms the last deposit of the Coal- bearing Series (see Kotas, 1995; Fig. 1A). In the NE part of the basin, the upper Malinowice Beds form the lower part of the Paralic Se ries. The marine faunal hori zon of the the Go³onóg Sandstone occurs locally within the Malinowice Beds. This hori zon corre sponds to the Štúr Faunal Hori zon, present in the Czech part of the basin (Ko- tas, 1972, 1995). Above the Go³onóg Sandstone ho rizon, are the sand stones of the Sarnów Beds and mudstones with nu merous coal seams and marine faunal hori zons (Flora Beds and Grodziec Beds; Fig. 1A). The Go³onóg Sandstone marine faunal hori zon was first described by Roemer (1866, 1870). The Namurian age was deter mined by Doktorowicz-Hrebnicki (1935), Czarniecki (1959), and Kotas (1972). Currently, the ho rizon is exposed in the NE part of the Up per Silesian Coal Ba sin in the area of the D¹browa Górnicza city, in the W rail-cut be tween Laski and Tworzeñ districts (GPS co-ordi nates: 50°19'55.6"N, 19°15'35.3"E; Fig. 1C). The marine fauna belongs to two assem blages (see Weigner, 1938), preserved in the form of im- prints and inter nal moulds in yellow-gray sandstones (bi- valve-brachi o pod as sem blage with cri noids, cor als, and trilobites) and as partially preserved shells in gray mudstones (gastro pod assem blage with nautiloids), from which the shell of Naticopsis planispira co mes.

MA TE RIAL AND METH ODS

In order to better illus trate the col our pat tern on the shell, Fig. 2. Naticopsis (Naticopsis) planispira (Phillips, 1836) from a UV quartz lamp (EMITA VP-60) was used. To enhance the the Up per Silesian Coal Ba sin (Namurian A), Poland, GIUS morpho log i cal details, the shell was powdered with ammo - 5-3605 DG-1. The spec i men was powdered with am monium chlo - nium chlo ride be fore be ing pho to graphed. ride. A. Aper tural view of shell. B. Apical view with slightly The speci men is housed at the col lec tion of the Depart - thickened collabral lines visi ble ment of Palae on tol ogy and Stra tig ra phy of the Uni ver sity of Silesia in Sosnowiec (abbre vi ated GIUS). pl. 4, fig. 4. 1966 Naticopsis (Naticopsis) planispira (Phillips) – Batten: p. 62, pl. 7, figs 4–5. SYS TEM ATIC PALAE ON TOL OGY 1972 Naticopsis planispira (Phillips) – Øehoø and Øehoøová: p. 50, pl. 21, figs 6–12. Order NERITIMORPHA Koken, 1896 1973 Naticopsis planispira (Phillips) – Gromczakiewicz- Family NATICOPSIDAE Waagen, 1880 £omnicka: p. 44, pl. 9, fig. 4. Genus Naticopsis M’Coy, 1844 Ma te rial. One very well-pre served shell (GIUS 5-3605 DG-1). Type spe cies Naticopsis phillipsii M’Coy, 1844 Mea sure ments of the shell: 6.4 mm height and 6.4 mm width (0.9 mm height of spire). Naticopsis (Naticopsis) planispira (Phillips, 1836) De scrip tion. The shell is globu lar and con sists of three, strongly Figs 2A, B, 3A–F in creas ing whorls, cov er ing three quar ters of the shell height. The base is rounded and anomphalous. The spire is low and the suture *1836 Natica planispira sp. nov. – Phillips: p. 244, pl. 14, is rela tively shal low, but distinct. On the pari etal part of aper ture, a fig. 24. thick ened smooth inductura is located (Fig. 2A). The up per part of 1843 Nerita spirata Sowerby – de Koninck: p. 484, pl. 42, the shell sur face is gently flat tened. The whorls are evenly roun- fig. 3d. ded and dis tinctly con vex. Growth lines are strongly prosoclinal. 1881 Naticopsis(?) planispira (Phillips) – de Koninck: p. Or na men ta tion con sists of slightly strength ened collabral lines, 20, pl. 2, figs 23–24, pl. 3, figs 9–10. only on the up per flat tened subsutural part of the whorl (Fig. 2B). non 1930 Naticopsis cf. planispira (Phillips) – Kühne: p. 98, Api cal an gle equals 130°. The protoconch is not pre served. 90 W. KRAWCZYÑSKI

Fig. 3. Zig zag-type col our pat tern of Naticopsis (Naticopsis) planispira (Phillips, 1836) from the Up per Silesian Coal Ba sin (Namurian A), Poland, GIUS 5-3605 DG-1. A, C, E. Photo graphs un der UV light. B, D, F. Hand-made draw ings of shell views, based on pho to - graphs. A, B. Lat eral view of the shell with a damage visi ble. C–D. Apical view of shell. E, F. Basal view of the shell

Col our pat tern. On the sur face of the last whorl and par tially on The colour pattern is rela tively reg ular in the earlier ontogenetic the spire whorls, the col our pat tern is pre served in the form of stage. brown markings on a light background. A zig zag-type pat tern is Gen er ally nar row col oured belts of dif fer ent widths (about ten domi nant. In ad di tion, there are prosoclinal belts and also two on a whorl) run from the su ture in a opisthoclinal di rec tion, and rows of trap e zoidal or ir reg u lar spots in later ontogenetic stage. then break (sharply bend with out loosing its con ti nuity) at an an gle COL OUR PAT TERN OF Naticopsis planispira 91

of 80° in a prosoclinal di rec tion with the chev ron edge di rected tian; Renish Slate Mountains, Germany; see Frýda, 2000). adaperturally (Fig. 3C, D). An other dou ble break at an an gle of Fre quently, col our pat terns are ob served in Carbon if er ous 33° oc curs at a whorl height of seven-eights, fol lowed by ir reg u lar neritimorph shells, es pe cially in Naticopsis M’Coy, 1844 prosoclinal belts. These belts break again adaperturally at an an gle and Trachydomia Meek et Worthen, 1866, which is proba - of 27° on the pe riph ery of the whorl. Some times, the break ing bly a member of the Trachyspiridae Nützel, Frýda, Yancey neigh bour ing belts merge into ir reg u lar spots. Af ter the last break, nar row belts run strongly opisthoclinally and trans versely again to et An der son, 2007. Speci mens of Trachydomia, with a col - ir reg u lar prosoclinal belts just be low the pe riph ery of the whorl. our pat tern, consist ing of either irreg u lar collabral belts or Another break of the col oured belts is formed at a whorl height of dark spots on the tops of numer ous nodes, were found at the one quar ter and at an an gle of 27° with the adapertural chev ron Buckhorn Asphalt Quarry near Sulphur in Oklahoma, USA (Fig. 3A, B). The last vis i ble abapertural chevron is lo cated on the (Bandel et al., 2002). Single finds of shells with the colour base of the shell and then belts bend arcuately in the di rec tion to pattern preserved have also been reported from the Permian. the basal part of the ap er ture (Fig. 3E, F). Kemp (1957) described a taxon of the genus Naticopsis A mechan i cal dam age of the shell labrum run ning ir reg u larly with narrow collabral belts, coming from the lower Permian to prosoclinally is visi ble in later ontogenetic stage of the shell depos its of Texas (USA). Naticopsis minima (Brown) with growth. This damage causes a clear change in a rela tively reg ular col our pat tern (see Fig. 3). Dou ble breaks of belts at the whorl a zig zag-type colour pat tern has been described by Hollin- height of seven-eights be come very in distinct and prosoclinal gworth and Barker (1991) from the Zechstein reef limesto- arched belts of ir reg ular width ap pear in their place. Adapertural nes of Eng land. bent chev rons on the whorl pe riph ery merge into ir reg u lar spots. Differ ent types of colour pattern preserved on neriti- However, the breaks oc cur ring at the whorl height of one quar ter morph shells were also recorded from the Meso zoic (e.g., pass into rel a tively large trap e zoidal spots, above which there are Tichy, 1980; Squires, 1993; Bandel and Kiel, 2003; Bandel, double break of nar row strips. Be low trap e zoidal spots with belts 2007; Kaim et al., 2013) and Ceno zoic (e.g., Bandel, 2001; of ir reg u lar width break to abapertural di rec tion and they are not Wesselingh, 2006; Symonds and Pacaud, 2010). visi ble on the base of the shell (see Fig. 3E, F). The genus Naticopsis has a globu lar or oval shell, Re marks. The shell of Naticopsis (N.) planispira, char ac ter ized which has a rela tively low spire with a strongly expanded by a low spire and a slightly flat tened up per part of the whorl with thick ened collabral lines, has been de scribed by Phillips (1836) last whorl in a di rection oblique to the axis and an anompha- from the Visean of Eng land for the first time. It also oc curs in the lous base. There are two subgen era within the genus Natico- Hotwells Lime stone from Compton Martin (Somerset, Eng land; psis. Naticopsis (Naticopsis) M’Coy, 1844 has a glob u lar Batten, 1966). This taxon, initially classi fied as Nerita spirata shell with a rounded convex profile of the whorl, while Sowerby (see de Koninck, 1843), has been found in the Visean Naticopsis (Jedria) Yochelson, 1953 has a pronounced lime stones in the eastern part of the Namur Syncline in Bel gium, swelling below the suture and a flattened upper surface of from where its concen tric operculum was de scribed as well (de the whorl. The Trias sic genus Marmolatella Kittl, 1894, Koninck, 1881). Gromczakiewicz-£omnicka (1973) also noted N. classified by Knight et al. (1960) as a subge nus of Naticop- planispira in the Visean cri noid-coral limestones from the Holy sis, now is a sepa rate genus within the subfamily Platychi- Cross Mountains, central Po land. This species was also found in the Paralic Series (Namurian A) in Ostrava part of the Upper Sile- lininae Bandel, 2007. Colour patterns are preserved on the sian Coal Ba sin. It ap pears in ma rine fau nal ho ri zons of Naneta, shells of both subgen era Naticopsis and Jedria. Františka, Enna, Koksova and Gaebler (Øehoø and Øehoøová, 1972) and in the older Go³onóg Sand stone ho ri zon, which is an Naticopsis (Naticopsis) subovata Worthen, 1873 equiv a lent of the Štúr Faunal Ho ri zon in the Ostrava area (see Fig. 4C Kotas, 1995). How ever, neritopsid shells, rec ognized by Kühne (1930) and sampled from Ptasia Góra and Jugów near Wa³brzych The zig zag-type col our pat tern con sists of dark, broad belts of (The Sudetes), have a much higher spire and a smaller apical angle ir reg ular width (about eight on the last whorl). At the be gin ning, (105°). be low the su ture, the belts run opisthoclinally and break dou bly in Oc cur rence. Visean: Eng land: Bolland (Yorkshire), Compton the adapertural and abapertural di rec tions at an angle of 100°. The Martin (Somerset); Belgium: Vise and Lives-sur-Meuse near Na- larg est break in the adapertural di rec tion at an an gle of 80° ap pears mur (Namur Syncline); Po land: Ga³êzice (Holy Cross Mountains). just above the pe riph ery of the whorl and the next abapertural one Namurian A: Upper Silesian Coal Basin: Ostrava area (Czech Re- at three-eights of the whorl height, at an an gle of 105°. The col our pub lic); D¹browa Górnicza-Laski (Po land). belts are not pre served on the base. N. subovata was de scribed by Worthen (1873) from the Up- per Penn syl va nian (Mis sourian) of Il li nois, USA. It was also re - corded from the Penn syl va nian of Ar kan sas (as Strophostylus RE VIEW OF CAR BON IF ER OUS subovata; Mather, 1915), the Mid dle Penn sylva nian of Mis souri, NATICOPSIDS WITH PRE SERVED USA (Knight, 1933), and the Up per Penn syl va nian (up per Vir- COL OUR PAT TERNS gilian) of Kan sas, USA (Mudge and Yochelson, 1962). Poorly pre - served spec i mens of this taxon were found in the low er most Perm- ian of New Mex ico, USA (Kues, 1991). Among gastro pods, colour patterns are most often preserved on neritimorph shells, owing to the presence of the outer calcitic pris matic layer, con tain ing the re mains of pig - Naticopsis (Naticopsis) picta Girty, 1912 ments (see Cox, 1960; Nützel et al., 2007; Frýda, 2012). Fig. 4D The old est colour pat tern on neritimorph shells in the The zig zag-type col our pat tern con sists of numer ous (about form of three rows of oval spots was found in Paffrathopsis twelve on last whorl) narrow and rel a tively reg ular belts. At the subcostata (d’Archiac et Verneuil, 1842) at Paffrath (Give- be gin ning, wide, ir reg u lar col our belts run in a prosoclinal di rec - 92 W. KRAWCZYÑSKI

Fig. 4. Re view of col our pat tern types on lat eral views of the Car bonif er ous Naticopsis shells. A. Naticopsis (Jedria) meeki Knight, 1933, Mid dle Penn syl va nian of St. Louis, Missouri, USA (after Knight, 1933, pl. 42, fig. 1c). B. Naticopsis (Jedria) ventrica (Norwood et Pratten, 1855), Middle Penn sylva nian of Pontotoc, Oklahoma, USA (af ter Knight, 1933, pl. 41, fig. 2d). C. Naticopsis (Naticopsis) subovata Worthen, 1873, Up per Penn sylva nian of Il li nois, USA (after Knight, 1933, pl. 43, fig. 2j). D. Naticopsis (Naticopsis) picta Girty, 1912, Up per Mis sissip pian of Perry County, In di ana, USA (af ter Girty, 1912, pl. 1, fig. 9). E. Naticopsis (Naticopsis) wortheniana Knight, 1934, Mid dle Penn sylva nian of the Buckhorn Asphalt Quarry, Oklahoma, USA (af ter Squires, 1976, fig. 1a). F. Naticopsis (Naticopsis) virgata Knight, 1933, Mid dle Penn sylva nian of St. Louis, Missouri, USA (after Knight, 1933, pl. 44, fig. 6g). G. Naticopsis sp., Mid dle Pennsyl va nian of the Buckhorn Asphalt Quarry, Oklahoma, USA (af ter Seuss et al., 2009, fig. 16a). H. Naticopsis (Naticopsis) pulchella Morningstar, 1922, Mid dle Penn syl va nian of Stark County, Ohio, USA (af ter Knight, 1933, pl. 44, fig. 3)

tion be low the su ture and break (lo cally double break) at seven- Naticopsis picta was de scribed only from the Upper Missis - eights of the whorl height at an an gle of 60°, form ing a chev ron in sippian (Chesterian) of In di ana, USA (Girty, 1912; see also Haya- an abapertural di rec tion. Then the belts be come nar row and reg u- saka, 1953). lar with many small sec ond ary chev rons. They run strongly opisthoclinally and break again at an an gle of 23° in an adapertural di - Naticopsis (Naticopsis) wortheniana Knight, 1934 rec tion and con tinue quite far in an abapertural direc tion. Nar row Fig. 4E belts break a few times at a con stant an gle of 23° just above and be low the pe riph ery of the whorl. From there they con tinue in an The col our pat tern con sists of dark spi ral bands, which are opisthoclinal di rec tion and break again at one third of whorl height only visi ble on the last whorl. The adapical band is lo cated just be - in adapertural di rec tion. The di rec tion of the belts is not vis i ble on low the up per su ture. The cen tral and wid est band has con stant the base of the shell. width in the early ontogenetic stages, while in later stages its width COL OUR PAT TERN OF Naticopsis planispira 93

be comes vari able and tri an gu lar spots are pres ent. This belt is lo - above half of the whorl height. Then the belts run strongly opistho- cated on the up per sur face of whorl from a height of three quar ters clinally and break twice at about three-eights of the whorl height, to pe riph ery of the whorl. The third spi ral band has a contin uous at an an gle of 40°. From the last break on ward, the belts run stron- width that is simi lar to the width of the first band. It appears on the gly opisthoclinally to ward the basal part of the labrum. rounded base from one quar ter to one eight of the whorl height. This spe cies is fairly wide spread in Up per Car bon if er ous sed- Naticopsis wortheniana was first de scribed by Knight (1933) i ments. Naticopsis altonensis (McChesney, 1865), N. ventricosus from the Penn syl va nian of St. Louis (Missouri, USA) as Naticop- Meek et Worthen, 1873, N. pricei Shumard, 1858, N. torta (Meek, sis wortheni. This name proved to be a youn ger homonym of 1871) proved to be younger synonyms of this species (see Yochel- Naticopsis wortheni Weller, 1916 (see Knight, 1934). Sev eral son and Saunders, 1967). Naticopsis (Jedria) ventrica was first de - spec i mens of very well-pre served shells with col our pat terns were scribed from the Up per Car bon if er ous of Indi ana, USA (Norwood also found in the Mid dle Penn sylva nian of the Buckhorn Asphalt and Pratten, 1855). The spe cies was also re corded in the Middle Quarry, Oklahoma, USA (Squires, 1976). and Up per Penn sylva nian of St. Louis, Henry County (Missouri, USA; Knight, 1933), Summit, Muskingum, and Vinton Coun ties Naticopsis (Naticopsis) virgata Knight, 1933 (Ohio, USA; Morningstar, 1922; Knight, 1933), San Juan re gion (Col o rado, USA; Girty, 1903), New Mex ico, USA (Kues and Bat - Fig. 4F ten, 2001). The col our pat tern con sists of three spi ral dark bands. The first band oc curs at half of a distance be tween su ture and pe riph ery Naticopsis sp. of the whorl (see Knight, 1933, pl. 44, fig. 6b). The wid est band is Fig. 4G lo cated just be low pe riph ery at five-eights of the whorl height. The slightly nar rower third band is sit u ated in the mid dle of the roun- The col our pat tern is com posed of three dark spiral bands. ded base (from about three-eights to one eight of the whorl height). The up per band ranges from periph ery of the whorl to at three- N. virgata was only re corded in the Middle Penn syl va nian of quar ter of the whorl height. The middle band is the widest one. It is St. Louis, Mis souri, USA (Knight, 1933). sit u ated at one eight to three-eights of the whorl height. The third band is about as wide as the up per band. It is sit uated at the lower Naticopsis (Naticopsis) pulchella Morningstar, 1922 part of the base. This spec i men was found in the Buckhorn As phalt Quarry, in Fig. 4H Oklahoma, USA (Mid dle Penn syl va nian; Seuss et al., 2009), and The shell has a col our pat tern con sisting of spi ral dark bands. clearly dif fers with re spect to both the pro por tion of shell and the There are two wide spi ral bands on the up per sur face of the whorl: col our band ing of the spe cies N. wortheniana Knight, 1934, which a nar rower band is lo cated at three-quar ter of the whorl height and was de scribed ear lier from the same de posits (Squires, 1976). a wider band is sit u ated just above pe riph ery of the whorl. Foerste (1930) men tions two taxa of Car bon if er ous neritids This taxon was found in black shales of the Mid dle Penn syl- with pre served col our pat terns: Naticopsis (Naticopsis) lirata Phi- va nian Potts ville For mation from Ohio, USA (Morningstar, 1922; llips, 1836 and Naticopsis (Jedria) plicistria (Phillips, 1836), but see also Knight, 1933; Yochelson and Saunders, 1967). there are no de scrip tions of these pat terns.

Naticopsis (Jedria) meeki Knight, 1933 Fig. 4A DIS CUS SION The zig zag-type col our pat tern con sists of about eight dark belts on the last whorl. Wide, irreg u lar belts start in the prosoclinal Among nu mer ous Car bon if er ous spe cies of the ge nus di rec tion at the suture and run archwise as chev rons (break an gle is Naticopsis only nine exhibit colour patterns on their shells. 27°) in adapertural di rec tion at a seven-eights of the whorl height. The pre dom inant pattern is the zig zag-type which is rec og- The next belts with nu merous irreg ular sec ondary breaks are di - nized in five taxa. Four taxa have the spiral band-type pat- rected strongly prosoclinally and break in the abapertural di rec tion tern (see Fig. 4; Tab. 1). Zigzag-type colour patterns clearly at an an gle of about 60° on the pe riph ery of the whorl. A fur ther differ from each other with respect to the morphol ogy of adapertural break is sit u ated at a quar ter of the height of the whorl belts, their width, repeat abil ity, break angle and loca tion of with an an gle of 80°, after which the belts run prosoclinally to ward chevrons on the whorl (see Figs 3, 4A–D). Differ ences are the base. Col our belts are not visi ble on the base. also observed among speci mens with spiral band-type pat- This taxon is the type spe cies of the sub ge nus Naticopsis (Jedria) Yochelson, 1953. It oc curs in the Mid dle Penn sylva nian terns, and in partic u lar their rela tive width and posi tion on of St. Louis and Barton Counties, in Mis souri, USA (Knight, 1933; the whorl (see Fig. 4E–G). Hoare, 1961). Poorly pre served spec i mens of Naticopsis (Jedria) Up to now, naticopsid shells with the dotted-type colour meeki was also recorded from the Lower Perm ian lime stones of pat terns, ap pear ing on Tri as sic shells of Naticopsis (see New Mex ico, USA (Kues, 1991). Tichy, 1980, pl. 3, fig. 2; Kaim et al., 2013, fig. 4), were not known from the Palaeozoic. Proba bly, the devel op ment of Naticopsis (Jedria) ventrica (Norwood et Pratten, 1855) new types of colour pattern could be related to the biotic re- Fig. 4B covery of gastro pods after the end-Perm ian mass ex tinction (see Nützel, 2005) and the increas ing im portance of preda - The zig zag-type col our pat tern con sists of eight dark belts on tion in the early stages of the Meso zoic marine revolution the last whorl. Ir reg ular wide belts (about twice as wide as on the base of the shell) run prosoclinally from the suture and then they (see Vermeij, 1977). break at seven-eights of the whorl height at an an gle of 115°. The A colour pattern is formed by tempo ral activ ity of pig - next break is adapertural at two-third of the whorl height, at an an - ment-se cret ing cells in the man tle mar gin at the edge of lab- gle of 70°. From this point on ward the belts are clearly nar rower rum. A specific pattern is formed, depend ing on the loca tion and more reg ular. A fur ther abapertural break ap pears slightly of these cells and their current activ ity (see Cox, 1960). In 94 W. KRAWCZYÑSKI

Ta ble 1 Review of Carbon if er ous taxa of Naticopsis with type of colour pattern, age and local ity of findings

Type of Taxon Age Locality Reference colour pattern Naticopsis (N.) subovata Worthen, Upper Pennsylvanian (Middle Chanute Shale, Turkey Creek, Kansas City, zigzag Knight, 1933 1873 Pennsylvanian–Lower Permian) Missouri, USA Naticopsis (Jedria) ventrica Middle Pennsylvanian (Middle middle Boggy Fm, Pontotoc County, zigzag Knight, 1933 (Norwood et Pratten, 1855) Pennsylvanian–Lower Permian) Oklahoma, USA Naticopsis (Jedria) meeki Knight, zigzag Middle Pennsylvanian St. Louis County, Missouri, USA Knight, 1933 1933 Naticopsis (N.) pulchella Lower Mercer Black Shale, Stark County, Morningstar, 1922; spiral bands Middle Pennsylvanian Morningstar, 1922 Ohio, USA Knight, 1933 Naticopsis (N.) virgata Knight, 1933 spiral bands Middle Pennsylvanian St. Louis County, Missouri, USA Knight, 1933 Naticopsis sp. spiral bands Middle Pennsylvanian Buckhorn Asphalt, Oklahoma, USA Seuss et al., 2009 Naticopsis (N.) wortheniana Knight, St. Louis County, Missouri, USA; Knight, 1933; spiral bands Middle Pennsylvanian 1934 Buckhorn Asphalt, Oklahoma, USA Squires, 1976 Naticopsis (N.) planispira (Phillips, Paralic Series, Upper Silesian Coal Basin, zigzag Namurian A (Visean–Namurian) this paper 1836) Poland Chester Group, Perry County, Indiana, Naticopsis (N.) picta Girty, 1912 zigzag Upper Mississippian Girty, 1912 USA

the case of spi ral bands, the pig ment-se cret ing cells are in a tion of the regu lar zigzag-type pattern into a more irreg u lar fixed posi tion and perma nently produce pigment. The zig- pattern, caused by some dam age of the outer edge of the zag-type pattern forms, when cell ac tiv ity changes over a shell labrum (see Symonds, 2008, fig. 11). specific pe riod of time, whereas the collabral-type pat tern A colour pat tern is an adaptative fea ture, depend ing on arises, when all the cells lo cated at the labrum are tempo - the en viron ment. In the case of gas tropods, it can be im por- rarily active or inactive (see Kobluk and Mapes, 1989). tant as cam ou flage, mak ing the an i mal in dis tin guish able Col our pat terns may change during ontog eny, owing to from the substrate in the photic zone. Zigzag-type and band- exter nal, envi ron men tal changes. This is known as colour type pat terns harmo nize well with a striped net of shadows pat tern poly mor phism and is ob served rel a tively fre quently on the bot tom, created by waves at the sur face of the wa ter. among mod ern and fos sil neritimorphs (see Grüneberg and The zigzag-type pattern can also be a warning sign to poten - Nugaliyade, 1976; Symonds, 2008). There also has been a tial pred a tors, be cause it is simi lar to the zigzag-type pat - large vari ety of col our pat terns in neritimorph shells of the terns on nautiloid shells. As well, it cannot be excluded that same species. For this reason, the colour pattern may not be the colour pat tern could be created to re tain toxic meta bolic a di ag nos tic fea ture for neritimorph spe cies. Ex am ples are products in the shell (see Tichy, 1980; Kelley and Swann, Ceno zoic fresh- or brackish-wa ter neritimorphs with vari - 1988; Kobluk and Mapes, 1989). ous zigzag-type colour patterns: the Eocene Clithon (Picto- neritina) pisiformis (Ferussac) from the Blackheath Beds, Abbey Wood, England (see Symonds, 2008, figs. 10–12), the Plio cene Theodoxus danubialis (Pfeiffer) from Hun gary CON CLU SIONS (see Bandel, 2001, figs. 27–31) and the Re cent Neritina ga- gates Lamarck from South Africa (see Bandel, 2001, figs. 1. Naticopsis (Naticopsis) planispira (Phillips, 1836) 68–71). has a dom inant zigzag-type colour pat tern, com bined with On the shell of Naticopsis planispira from the Upper collabral belts. Silesian Coal Ba sin, there is a clear change in the mor phol- 2. The change of the colour pattern was caused by me- ogy of in di vid ual el e ments from a rel a tively reg u lar zig zag- chani cal dam age to the shell, and was not re lated to colour type pattern to addi tional rows of irreg u lar or trape zoidal pat tern poly mor phism. spots (see Fig. 3A, B, E, F). This change in the colour pat- 3. Zig zag-type and spiral band-type colour pat terns are tern is not a result of col our pat tern poly morphism, because the most common among the Carbon if er ous Naticopsis spe - it abruptly appears along the site of clear mechan ical dam- cies. These colour pat terns were proba bly used for shell ca- age to the shell. The mechan ical damage prob a bly re sulted mouflage on the sea-floor in the photic zone. from en vi ron men tal fac tors (e.g., wa ter tur bu lence) or pre- 4. De spite a sig nif i cant dif fer en ti a tion of col our pat- da tion (see Jeffery et al., 1994). This phenom enon also has terns within the Palaeozoic Naticopsis shells, they cannot be been observed, for exam ple, among the Eocene neritid spe- re garded as di ag nos tic fea tures for the Naticopsis spe cies, cies Clithon (Pictoneritina) pisiformis (Ferussac). One of owing to a high intraspecific variabil ity of colour patterns the spec imens of the latter spe cies clearly shows a dis rup- on their shells. COL OUR PAT TERN OF Naticopsis planispira 95

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