Wojciech Krawczyński

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

De part ment of Palae on tol ogy and Stra tig ra phy, Fac ulty of Earth Sci ences, Uni ver sity of Silesia, Bêdziñska 60,

41-200 Sosnowiec, Po land; e-mail: wojciech.krawczynski@us.edu.pl

Krawczyñski, W., 2013. 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 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 u lar spots was ob served on a shell of Naticopsis (N.) planispira (Phillips, 1836) found in the Go³onóg Sand stone ma rine fau nal ho ri zon (Namurian A, the Up per Silesian Coal Ba sin, Po land). A dis rup tion in the col our pat tern was also ob served to have oc curred dur ing ontogenetic shell growth. In this spec i men, it ap peared in the healed shell dam age 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 com mon 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 (di rec tion and an gles of breaks) and the lo ca tion and width of the spi ral bands on a whorl. How ever, the color pat terns may not be di ag nos tic fea tures for the Naticopsis spe cies, be cause of the large intraspecific vari a tion and col our pat tern poly mor phism 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 sur face 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

Spec i mens of Palaeozoic ma rine in ver te brates with col

-our pat terns pre served on their shells or car a paces are rare.

They were de scribed from the shells of brachi o pods (e.g.,

Køíž and Lukeš, 1974; Blodgett et al., 1988; Baliñski, 2010),

bi

valves (e.g., Hoare et al., 1988; Mapes and Benstock,

1988), nautiloids (e.g., Manda and Turek, 2009; Turek,

2009), and also from tri lo bite car a paces (e.g., McRoberts et

al., 2013).

Col our pat terns on Palaeozoic mol lusc shells, in clud ing

gas tro pods, are pre served rel a tively sel dom, ow ing to the

fre quent recrystallization of the orig i nal ar agon ite shells.

The old est gas tro pods with pre served col our pat terns

(Stra-parollina harpa) have been found in the Mid dle Or do vi cian

de pos its of the USA (Ray mond, 1906; Foerste, 1930). In di

-vid ual finds have also been re corded from the Si lu rian (e.g.,

Køíž and Lukeš, 1974), De

vo

nian (e.g., Yochelson and

Køíž, 1974; Rohr and Smith, 1978; Frýda, 2000; Basse and

Heidelberger, 2002; Jankovský, 2003) and Perm ian de pos

-its (e.g., Yochelson, 1956; Bat ten, 1958, 1972; Plas, 1972).

Their great est di ver sity, how ever, is noted in Car bon if er ous

for ma tions (e.g., Knight, 1932, 1933; Bat ten, 1966; Hoare

and Stur geon, 1978; see also Kobluk and Mapes, 1989).

The pres ent pa per de scribes a Car bon if er ous gas tro pod

shell of Naticopsis (Naticopsis) planispira (Phillips, 1836)

with a pre served col our pat tern, from the Go³onóg Sand

-stone ma

rine fau

nal ho

ri

zon. The di

verse fau

nal as

sem

blage, con sist ing of about 50 taxa (in clud ing bi valves, gas

-tro pods, brachi o pods, and nautiloids), has been stud ied by

Cramer (1910), and Weigner (1938) and was re vised by

Bojkowski (1972). Ad

di

tional fau

nal re

search was con

-ducted by Schwarzbach (1935), Pøibyl (1951), and Salamon

(1997). Among the ten rec og nized gas tro pod taxa,

Naticop-sis planispira has not been listed pre vi ously.

GEO LOG I CAL SET TING

The Up per Silesian Coal Ba sin is mainly lo cated in the

south ern part of Po land and only its most SW part stretches

into the ter ri tory of the Czech Re pub lic (Fig. 1B). The Up

-per Car bon if er ous coal-bear ing se ries rests on the Lower

Car bon if er ous mudstone-sand stone de pos its (Culm fa cies)

in the W part and lime stones (Car bon if er ous lime stone fa

-cies) in its east ern part. The Up per Car bon if er ous de pos its

of the Up per Silesian Coal Ba sin con sist of the Paralic Se

-ries (the lower part of the Coal-bear ing Se -ries) in its lower

88

Fig. 1. Lo ca tion of out crops and strati graphi cal po si tion of the Go³onóg Sand stone ma rine fau nal ho ri zon. A. Strati graphic se quence of the Up per Car bon if er ous in the NE part of the Up per Silesian Coal Ba sin, show ing 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 Ba sin (af ter Jureczka et al., 1995). C. Geo log i cal map of the NE part of D¹browa Górnicza with sam ple 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)

part with nu mer ous ma rine fau nal ho ri zons (Namurian A).

Above is the Limnic Se ries 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 Sand stone

Series (Westphalian C and D), which is pres ent only in the

east ern part of the ba sin, forms the last de posit of the

Coal-bear ing Se ries (see Kotas, 1995; Fig. 1A).

In the NE part of the ba sin, the up per Malinowice Beds

form the lower part of the Paralic Se ries. The ma rine fau nal

ho ri zon of the the Go³onóg Sand stone oc curs lo cally within

the Malinowice Beds. This ho ri zon cor re sponds to the Štúr

Fau nal Ho ri zon, pres ent in the Czech part of the ba sin

(Ko-tas, 1972, 1995). Above the Go³onóg Sand stone ho ri zon,

are the sand stones of the Sarnów Beds and mudstones with

nu mer ous coal seams and ma rine fau nal ho

ri zons (Flora

Beds and Grodziec Beds; Fig. 1A).

The Go³onóg Sand stone ma rine fau nal ho ri zon was first

de scribed by Roemer (1866, 1870). The Namurian age was

de ter mined by Doktorowicz-Hrebnicki (1935), Czarniecki

(1959), and Kotas (1972). Cur rently, the ho ri zon is ex posed

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 dis

tricts (GPS co-or

di

nates: 50°19'55.6"N,

19°15'35.3"E; Fig. 1C). The ma rine fauna be longs to two as

sem blages (see Weigner, 1938), pre served in the form of im

prints and in ter nal moulds in yel lowgray sand stones (bi

-valve-brachi o pod as sem blage with cri noids, cor als, and

tri-lobites) and as par tially pre served shells in gray mudstones

(gas

tro

pod as

sem

blage with nautiloids), from which the

shell of Naticopsis planispira co mes.

MA TE RIAL AND METH ODS

In or der to better il lus trate the col our pat tern on the shell,

a UV quartz lamp (EMITA VP-60) was used. To en hance the

mor pho log i cal de tails, the shell was pow dered with am mo

-nium chlo ride be fore be ing pho to graphed.

The spec i men is housed at the col lec tion of the De part

-ment of Palae on tol ogy and Stra tig ra phy of the Uni ver sity of

Silesia in Sosnowiec (ab bre vi ated GIUS).

SYS TEM ATIC PALAE ON TOL OGY

Or der NERITIMORPHA Koken, 1896

Fam ily NATICOPSIDAE Waagen, 1880

Ge nus Naticopsis M’Coy, 1844

Type spe cies Naticopsis phillipsii M’Coy, 1844

Naticopsis (Naticopsis) planispira (Phillips, 1836)

Figs 2A, B, 3A–F

*1836 Natica planispira sp. nov. – Phillips: p. 244, pl. 14, fig. 24.

1843 Nerita spirata Sowerby – de Koninck: p. 484, pl. 42, fig. 3d.

1881 Naticopsis(?) planispira (Phillips) – de Koninck: p. 20, pl. 2, figs 23–24, pl. 3, figs 9–10.

non 1930 Naticopsis cf. planispira (Phillips) – Kühne: p. 98,

pl. 4, fig. 4.

1966 Naticopsis (Naticopsis) planispira (Phillips) – Bat ten: p. 62, pl. 7, figs 4–5.

1972 Naticopsis planispira (Phillips) – Øehoø and Øehoøová: p. 50, pl. 21, figs 6–12.

1973 Naticopsis planispira (Phillips) – Gromczakiewicz-£omnicka: p. 44, pl. 9, fig. 4.

Ma te rial. One very well-pre served shell (GIUS 5-3605 DG-1).

Mea sure ments of the shell: 6.4 mm height and 6.4 mm width (0.9 mm height of spire).

De scrip tion. The shell is glob u lar and con sists of three, strongly

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 su ture is rel a tively shal low, but dis tinct. On the pa ri etal part of ap er ture, a thick ened smooth inductura is lo cated (Fig. 2A). The up per part of the shell sur face is gently flat tened. The whorls are evenly roun-ded and dis tinctly con vex. Growth lines are strongly prosoclinal. Or na men ta tion con sists of slightly strength ened collabral lines, only on the up per flat tened subsutural part of the whorl (Fig. 2B). Api cal an gle equals 130°. The protoconch is not pre served.

Fig. 2. Naticopsis (Naticopsis) planispira (Phillips, 1836) from the Up per Silesian Coal Ba sin (Namurian A), Po land, GIUS 53605 DG1. The spec i men was pow dered with am mo nium chlo -ride. A. Ap er tural view of shell. B. Apical view with slightly thick ened collabral lines vis i ble

Col our pat tern. On the sur face of the last whorl and par tially on

the spire whorls, the col our pat tern is pre served in the form of brown mark ings on a light back ground. A zig zag-type pat tern is dom i nant. In ad di tion, there are prosoclinal belts and also two rows of trap e zoidal or ir reg u lar spots in later ontogenetic stage.

The col our pat tern is rel a tively reg u lar in the ear lier ontogenetic stage.

Gen er ally nar row col oured belts of dif fer ent widths (about ten on a whorl) run from the su ture in a opisthoclinal di rec tion, and then break (sharply bend with out loosing its con ti nu ity) at an an gle

90

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), Po land, GIUS 53605 DG1. A, C, E. Pho to graphs un der UV light. B, D, F. Handmade draw ings of shell views, based on pho to -graphs. A, B. Lat eral view of the shell with a dam age vis i ble. C–D. Api cal view of shell. E, F. Basal view of the shell

of 80° in a prosoclinal di rec tion with the chev ron edge di rected adaperturally (Fig. 3C, D). An other dou ble break at an an gle of 33° oc curs at a whorl height of seven-eights, fol lowed by ir reg u lar prosoclinal belts. These belts break again adaperturally at an an gle of 27° on the pe riph ery of the whorl. Some times, the break ing 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 ir reg u lar prosoclinal belts just be low the pe riph ery of the whorl. An other break of the col oured belts is formed at a whorl height of one quar ter and at an an gle of 27° with the adapertural chev ron (Fig. 3A, B). The last vis i ble abapertural chev ron is lo cated on the base of the shell and then belts bend arcuately in the di rec tion to the basal part of the ap er ture (Fig. 3E, F).

A me chan i cal dam age of the shell labrum run ning ir reg u larly to prosoclinally is vis i ble in later ontogenetic stage of the shell growth. This dam age causes a clear change in a rel a tively reg u lar col our pat tern (see Fig. 3). Dou ble breaks of belts at the whorl height of seven-eights be come very in dis tinct and prosoclinal arched belts of ir reg u lar width ap pear in their place. Adapertural bent chev rons on the whorl pe riph ery merge into ir reg u lar spots. How ever, the breaks oc cur ring at the whorl height of one quar ter pass into rel a tively large trap e zoidal spots, above which there are dou ble break of nar row strips. Be low trap e zoidal spots with belts of ir reg u lar width break to abapertural di rec tion and they are not vis i ble on the base of the shell (see Fig. 3E, F).

Re marks. The shell of Naticopsis (N.) planispira, char ac ter ized

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) from the Visean of Eng land for the first time. It also oc curs in the Hotwells Lime stone from Compton Mar tin (Somerset, Eng land; Bat ten, 1966). This taxon, ini tially clas si fied as Nerita spirata Sowerby (see de Koninck, 1843), has been found in the Visean lime stones in the east ern part of the Namur Syncline in Bel gium, from where its con cen tric operculum was de scribed as well (de Koninck, 1881). Gromczakiewicz-£omnicka (1973) also noted N. planispira in the Visean cri noid-coral lime stones from the Holy Cross Moun tains, cen tral Po land. This spe cies was also found in the Paralic Se ries (Namurian A) in Ostrava part of the Up per Sile-sian Coal Ba sin. It ap pears in ma rine fau nal ho ri zons of Naneta, Františka, Enna, Koksova and Gaebler (Øehoø and Øehoøová, 1972) and in the older Go³onóg Sand stone ho ri zon, which is an equiv a lent of the Štúr Fau nal Ho ri zon in the Ostrava area (see Kotas, 1995). How ever, neritopsid shells, rec og nized by Kühne (1930) and sam pled from Ptasia Góra and Jugów near Wa³brzych (The Sudetes), have a much higher spire and a smaller api cal an gle (105°).

Oc cur rence. Visean: Eng land: Bolland (York shire), Compton Martin (Somerset); Bel gium: Vise and Lives-sur-Meuse near Na-mur (NaNa-mur Syncline); Po land: Ga³êzice (Holy Cross Moun tains). Namurian A: Up per Silesian Coal Ba sin: Ostrava area (Czech Re -pub lic); D¹browa Górnicza-Laski (Po land).

RE VIEW OF CAR BON IF ER OUS

NATICOPSIDS WITH PRE SERVED

COL OUR PAT TERNS

Among gas tro pods, col our pat terns are most of ten pre

-served on neritimorph shells, ow ing to the pres ence of the

outer calcitic pris matic layer, con tain ing the re mains of pig

-ments (see Cox, 1960; Nützel et al., 2007; Frýda, 2012).

The old est col our pat tern on neritimorph shells in the

form of three rows of oval spots was found in Paffrathopsis

subcostata (d’Archiac et Verneuil, 1842) at Paffrath

(Give-tian; Renish Slate Moun tains, Ger many; see Frýda, 2000).

Fre quently, col our pat terns are ob served in Car bon if er ous

neritimorph shells, es pe cially in Naticopsis M’Coy, 1844

and Trachydomia Meek et Worthen, 1866, which is prob a

-bly a mem ber of the Trachyspiridae Nützel, Frýda, Yancey

et An der son, 2007. Spec i mens of Trachydomia, with a col

-our pat tern, con sist ing of ei ther ir reg u lar collabral belts or

dark spots on the tops of nu mer ous nodes, were found at the

Buckhorn As phalt Quarry near Sul phur in Oklahoma, USA

(Bandel et al., 2002). Sin gle finds of shells with the col our

pat tern pre served have also been re ported from the Perm ian.

Kemp (1957) de scribed a taxon of the ge nus Naticopsis

with nar row collabral belts, com ing from the lower Perm ian

de pos its of Texas (USA). Naticopsis min ima (Brown) with

a zig zag-type col our pat tern has been de scribed by

Hollin-gworth and Barker (1991) from the Zechstein reef

limesto-nes of Eng land.

Dif fer ent types of col our pat tern pre served on

neriti-morph shells were also re corded from the Me so zoic (e.g.,

Tichy, 1980; Squires, 1993; Bandel and Kiel, 2003; Bandel,

2007; Kaim et al., 2013) and Ce no zoic (e.g., Bandel, 2001;

Wesselingh, 2006; Symonds and Pacaud, 2010).

The ge

nus Naticopsis has a glob

u

lar or oval shell,

which has a rel a tively low spire with a strongly ex panded

last whorl in a di rec tion oblique to the axis and an anompha-

lous base. There are two sub gen era within the ge nus Natico-

psis. Naticopsis (Naticopsis) M’Coy, 1844 has a glob u lar

shell with a rounded con vex pro file of the whorl, while

Naticopsis (Jedria) Yochelson, 1953 has a pro

nounced

swelling be low the su ture and a flat tened up per sur face of

the whorl. The Tri as sic ge nus Marmolatella Kittl, 1894,

classified by Knight et al. (1960) as a sub ge nus of

Naticop-sis, now is a sep a rate ge nus within the subfamily

Platychi-lininae Bandel, 2007. Col our pat terns are pre served on the

shells of both sub gen era Naticopsis and Jedria.

Naticopsis (Naticopsis) subovata Worthen, 1873

Fig. 4C

The zig zag-type col our pat tern con sists of dark, broad belts of ir reg u lar width (about eight on the last whorl). At the be gin ning, be low the su ture, the belts run opisthoclinally and break dou bly in the adapertural and abapertural di rec tions at an an gle of 100°. The larg est break in the adapertural di rec tion at an an gle of 80° ap pears just above the pe riph ery of the whorl and the next abapertural one at three-eights of the whorl height, at an an gle of 105°. The col our 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 sou rian) of Il li nois, USA. It was also re -corded from the Penn syl va nian of Ar kan sas (as Strophostylus subovata; Mather, 1915), the Mid dle Penn syl va nian of Mis souri, USA (Knight, 1933), and the Up per Penn syl va nian (up per Virgilian) 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).

Naticopsis (Naticopsis) picta Girty, 1912

Fig. 4D

The zig zag-type col our pat tern con sists of nu mer ous (about twelve on last whorl) nar row and rel a tively reg u lar belts. At the be gin ning, wide, ir reg u lar col our belts run in a prosoclinal di rec

-tion be low the su ture and break (lo cally dou ble break) at seven-eights of the whorl height at an an gle of 60°, form ing a chev ron in an abapertural di rec tion. Then the belts be come nar row and reg u -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 -rec tion and con tinue quite far in an abapertural di -rec tion. Nar row 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 opisthoclinal di rec tion and break again at one third of whorl height in adapertural di rec tion. The di rec tion of the belts is not vis i ble on the base of the shell.

Naticopsis picta was de scribed only from the Up per Mis sis -sip pian (Chesterian) of In di ana, USA (Girty, 1912; see also Haya-saka, 1953).

Naticopsis (Naticopsis) wortheniana Knight, 1934

Fig. 4E

The col our pat tern con sists of dark spi ral bands, which are only vis i ble on the last whorl. The adapical band is lo cated just be -low the up per su ture. The cen tral and wid est band has con stant width in the early ontogenetic stages, while in later stages its width

92

Fig. 4. Re view of col our pat tern types on lat eral views of the Car bon if er ous Naticopsis shells. A. Naticopsis (Jedria) meeki Knight, 1933, Mid dle Penn syl va nian of St. Louis, Mis souri, USA (af ter Knight, 1933, pl. 42, fig. 1c). B. Naticopsis (Jedria) ventrica (Norwood et Pratten, 1855), Mid dle Penn syl va nian of Pontotoc, Oklahoma, USA (af ter Knight, 1933, pl. 41, fig. 2d). C. Naticopsis (Naticopsis) subovata Worthen, 1873, Up per Penn syl va nian of Il li nois, USA (af ter Knight, 1933, pl. 43, fig. 2j). D. Naticopsis (Naticopsis) picta Girty, 1912, Up per Mis sis sip 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 syl va nian of the Buckhorn As phalt Quarry, Oklahoma, USA (af ter Squires, 1976, fig. 1a). F. Naticopsis (Naticopsis) virgata Knight, 1933, Mid dle Penn syl va nian of St. Louis, Mis souri, USA (af ter Knight, 1933, pl. 44, fig. 6g). G. Naticopsis sp., Mid dle Penn syl va nian of the Buckhorn As phalt 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)

be comes vari able and tri an gu lar spots are pres ent. This belt is lo -cated on the up per sur face of whorl from a height of three quar ters to pe riph ery of the whorl. The third spi ral band has a con tin u ous width that is sim i lar to the width of the first band. It ap pears on the rounded base from one quar ter to one eight of the whorl height.

Naticopsis wortheniana was first de scribed by Knight (1933) from the Penn syl va nian of St. Louis (Mis souri, USA) as Naticop-sis wortheni. This name proved to be a youn ger hom onym of Naticopsis wortheni Weller, 1916 (see Knight, 1934). Sev eral spec i mens of very well-pre served shells with col our pat terns were also found in the Mid dle Penn syl va nian of the Buckhorn As phalt Quarry, Oklahoma, USA (Squires, 1976).

Naticopsis (Naticopsis) virgata Knight, 1933

Fig. 4F

The col our pat tern con sists of three spi ral dark bands. The first band oc curs at half of a dis tance be tween su ture and pe riph ery of the whorl (see Knight, 1933, pl. 44, fig. 6b). The wid est band is 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-ded base (from about three-eights to one eight of the whorl height). N. virgata was only re corded in the Mid dle Penn syl va nian of St. Louis, Mis souri, USA (Knight, 1933).

Naticopsis (Naticopsis) pulchella Morningstar, 1922

Fig. 4H

The shell has a col our pat tern con sist ing of spi ral dark bands. There are two wide spi ral bands on the up per sur face of the whorl: a nar rower band is lo cated at three-quar ter of the whorl height and a wider band is sit u ated just above pe riph ery of the whorl.

This taxon was found in black shales of the Mid dle Penn syl -va nian Potts ville For ma tion from Ohio, USA (Morningstar, 1922; see also Knight, 1933; Yochelson and Saunders, 1967).

Naticopsis (Jedria) meeki Knight, 1933

Fig. 4A

The zig zag-type col our pat tern con sists of about eight dark belts on the last whorl. Wide, ir reg u lar belts start in the prosoclinal di rec tion at the su ture and run arch wise as chev rons (break an gle is 27°) in adapertural di rec tion at a seven-eights of the whorl height. The next belts with nu mer ous ir reg u lar sec ond ary breaks are di -rected strongly prosoclinally and break in the abapertural di rec tion at an an gle of about 60° on the pe riph ery of the whorl. A fur ther adapertural break is sit u ated at a quar ter of the height of the whorl with an an gle of 80°, af ter which the belts run prosoclinally to ward the base. Col our belts are not vis i ble on the base.

This taxon is the type spe cies of the sub ge nus Naticopsis (Jedria) Yochelson, 1953. It oc curs in the Mid dle Penn syl va nian of St. Louis and Barton Coun ties, in Mis souri, USA (Knight, 1933; Hoare, 1961). Poorly pre served spec i mens of Naticopsis (Jedria) meeki was also re corded from the Lower Perm ian lime stones of New Mex ico, USA (Kues, 1991).

Naticopsis (Jedria) ventrica (Norwood et Pratten, 1855)

Fig. 4B

The zig zag-type col our pat tern con sists of eight dark belts on the last whorl. Ir reg u lar wide belts (about twice as wide as on the base of the shell) run prosoclinally from the su ture and then they break at seven-eights of the whorl height at an an gle of 115°. The next break is adapertural at twothird of the whorl height, at an an -gle of 70°. From this point on ward the belts are clearly nar rower and more reg u lar. A fur ther abapertural break ap pears slightly

above half of the whorl height. Then the belts run strongly opistho- clinally and break twice at about three-eights of the whorl height, at an an gle of 40°. From the last break on ward, the belts run stron-gly opisthoclinally to ward the basal part of the labrum.

This spe cies is fairly wide spread in Up per Car bon if er ous sed -i ments. Nat-icops-is altonens-is (McChesney, 1865), N. ventr-icosus Meek et Worthen, 1873, N. pricei Shumard, 1858, N. torta (Meek, 1871) proved to be youn ger syn onyms of this spe cies (see Yochelson and Saunders, 1967). Naticopsis (Jedria) ventrica was first de -scribed from the Up per Car bon if er ous of In di ana, USA (Norwood and Pratten, 1855). The spe cies was also re corded in the Mid dle and Up per Penn syl va nian of St. Louis, Henry County (Mis souri, USA; Knight, 1933), Sum mit, Muskingum, and Vinton Coun ties (Ohio, USA; Morningstar, 1922; Knight, 1933), San Juan re gion (Col o rado, USA; Girty, 1903), New Mex ico, USA (Kues and Bat -ten, 2001).

Naticopsis sp.

Fig. 4G

The col our pat tern is com posed of three dark spi ral bands. The up per band ranges from pe riph ery of the whorl to at three-quar ter of the whorl height. The mid dle band is the wid est one. It is 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 u ated at the lower part of the base.

This spec i men was found in the Buckhorn As phalt Quarry, in Oklahoma, USA (Mid dle Penn syl va nian; Seuss et al., 2009), and clearly dif fers with re spect to both the pro por tion of shell and the col our band ing of the spe cies N. wortheniana Knight, 1934, which was de scribed ear lier from the same de pos its (Squires, 1976).

Foerste (1930) men tions two taxa of Car bon if er ous neritids with pre served col our pat terns: Naticopsis (Naticopsis) lirata Phi-llips, 1836 and Naticopsis (Jedria) plicistria (PhiPhi-llips, 1836), but there are no de scrip tions of these pat terns.

DIS CUS SION

Among nu mer ous Car bon if er ous spe cies of the ge nus

Naticopsis only nine ex hibit col our pat terns on their shells.

The pre dom i nant pat tern is the zig zagtype which is rec og

nized in five taxa. Four taxa have the spi ral bandtype pat

-tern (see Fig. 4; Tab. 1). Zig zag-type col our pat -terns clearly

dif fer from each other with re spect to the mor phol ogy of

belts, their width, re peat abil ity, break an gle and lo ca tion of

chev rons on the whorl (see Figs 3, 4A–D). Dif fer ences are

also ob served among spec i mens with spi ral bandtype pat

-terns, and in par tic u lar their rel a tive width and po si tion on

the whorl (see Fig. 4E–G).

Up to now, naticopsid shells with the dot ted-type col our

pat terns, ap pear ing on Tri as sic shells of Naticopsis (see

Tichy, 1980, pl. 3, fig. 2; Kaim et al., 2013, fig. 4), were not

known from the Palaeozoic. Prob a bly, the de vel op ment of

new types of col our pat tern could be re lated to the bi otic re

-cov ery of gas tro pods af ter the end-Perm ian mass ex tinc tion

(see Nützel, 2005) and the in creas ing im por tance of pre da

-tion in the early stages of the Me so zoic ma rine revolu-tion

(see Vermeij, 1977).

A col our pat tern is formed by tem po ral ac tiv ity of pig

-ment-se cret ing cells in the man tle mar gin at the edge of

lab-rum. A spe cific pat tern is formed, de pend ing on the lo ca tion

of these cells and their cur rent ac tiv ity (see Cox, 1960). In

the case of spi ral bands, the pig ment-se cret ing cells are in a

fixed po si tion and per ma nently pro duce pig ment. The zig

-zag-type pat tern forms, when cell ac tiv ity changes over a

specific pe riod of time, whereas the collabral-type pat tern

arises, when all the cells lo cated at the labrum are tem po

-rarily ac tive or inactive (see Kobluk and Mapes, 1989).

Col our pat terns may change dur ing on tog eny, ow ing to

ex ter nal, en vi ron men tal changes. This is known as col our

pat tern poly mor phism and is ob served rel a tively fre quently

among mod ern and fos sil neritimorphs (see Grüneberg and

Nugaliyade, 1976; Symonds, 2008). There also has been a

large va ri ety of col our pat terns in neritimorph shells of the

same spe cies. For this rea son, the col our pat tern may not be

a di ag nos tic fea ture for neritimorph spe cies. Ex am ples are

Ce no zoic fresh or brack ishwa ter neritimorphs with var i

-ous zig zag-type col our pat terns: the Eocene Clithon

(Picto-neritina) pisiformis (Ferussac) from the Black heath Beds,

Ab bey Wood, Eng land (see Symonds, 2008, figs. 10–12),

the Plio cene Theodoxus danubialis (Pfeif fer) from Hun gary

(see Bandel, 2001, figs. 27–31) and the Re cent Neritina

ga-gates Lamarck from South Af rica (see Bandel, 2001, figs.

68–71).

On the shell of Naticopsis planispira from the Up per

Silesian Coal Ba sin, there is a clear change in the mor phol

-ogy of in di vid ual el e ments from a rel a tively reg u lar zig

zag-type pat tern to ad di tional rows of ir reg u lar or trap e zoidal

spots (see Fig. 3A, B, E, F). This change in the col our pat

-tern is not a re sult of col our pat -tern poly mor phism, be cause

it abruptly ap pears along the site of clear me chan i cal dam

-age to the shell. The me chan i cal dam -age prob a bly re sulted

from en vi ron men tal fac tors (e.g., wa ter tur bu lence) or pre

-da tion (see Jeffery et al., 1994). This phe nom e non also has

been ob served, for ex am ple, among the Eocene neritid spe

-cies Clithon (Pictoneritina) pisiformis (Ferussac). One of

the spec i mens of the lat ter spe cies clearly shows a dis rup

-tion of the reg u lar zig zag-type pat tern into a more ir reg u lar

pat tern, caused by some dam age of the outer edge of the

shell labrum (see Symonds, 2008, fig. 11).

A col our pat tern is an adaptative fea ture, de pend ing on

the en vi ron ment. In the case of gas tro pods, it can be im por

-tant as cam ou flage, mak ing the an i mal in dis tin guish able

from the sub strate in the photic zone. Zig zag-type and band-

type pat terns har mo nize well with a striped net of shad ows

on the bot tom, cre ated by waves at the sur face of the wa ter.

The zig zagtype pat tern can also be a warn ing sign to po ten

tial pred a tors, be cause it is sim i lar to the zig zagtype pat

-terns on nautiloid shells. As well, it can not be ex cluded that

the col our pat tern could be cre ated to re tain toxic met a bolic

prod ucts in the shell (see Tichy, 1980; Kelley and Swann,

1988; Kobluk and Mapes, 1989).

CON CLU SIONS

1. Naticopsis (Naticopsis) planispira (Phillips, 1836)

has a dom i nant zig zag-type col our pat tern, com bined with

collabral belts.

2. The change of the col our pat tern was caused by me

-chan i cal dam age to the shell, and was not re lated to col our

pat tern poly mor phism.

3. Zig zag-type and spi ral band-type col our pat terns are

the most com mon among the Car bon if er ous Naticopsis spe

-cies. These col our pat terns were prob a bly used for shell

ca-mouflage on the sea-floor in the photic zone.

4. De spite a sig nif i cant dif fer en ti a tion of col our pat

-terns within the Palaeozoic Naticopsis shells, they can not be

re garded as di ag nos tic fea tures for the Naticopsis spe cies,

ow ing to a high intraspecific vari abil ity of col our pat terns

on their shells.

94

Ta ble 1

Re view of Car bon if er ous taxa of Naticopsis with type of col our pat tern, age and lo cal ity of find ings

Taxon Type of

colour pattern Age Locality Reference

Naticopsis (N.) subovata Worthen,

1873 zigzag

Upper Pennsylvanian (Middle Pennsylvanian–Lower Permian)

Chanute Shale, Turkey Creek, Kansas City,

Missouri, USA Knight, 1933 Naticopsis (Jedria) ventrica

(Norwood et Pratten, 1855) zigzag

Middle Pennsylvanian (Middle Pennsylvanian–Lower Permian)

middle Boggy Fm, Pontotoc County,

Oklahoma, USA Knight, 1933 Naticopsis (Jedria) meeki Knight,

1933 zigzag Middle Pennsylvanian St. Louis County, Missouri, USA Knight, 1933 Naticopsis (N.) pulchella

Morningstar, 1922 spiral bands Middle Pennsylvanian

Lower Mercer Black Shale, Stark County, Ohio, USA

Morningstar, 1922; 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,

1934 spiral bands Middle Pennsylvanian

St. Louis County, Missouri, USA; Buckhorn Asphalt, Oklahoma, USA

Knight, 1933; Squires, 1976 Naticopsis (N.) planispira (Phillips,

1836) zigzag Namurian A (Visean–Namurian)

Paralic Series, Upper Silesian Coal Basin,

Poland this paper Naticopsis (N.) picta Girty, 1912 zigzag Upper Mississippian Chester Group, Perry County, Indiana,

Ac knowl edge ments

I thank Waldemar Kierepka for do nat ing the gas tro pod shell for the cur rent re search. I would like to thank Al ex an der Nützel (München, Ger many) and Andrzej Kaim (War saw, Po land) for their con struc tive re views of the manu script. I also would like to thank Micha³ Zatoñ (Sosnowiec, Po land) for nu mer ous com ments and lin guis tic cor rec tions of the first ver sion of this text.

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