Grzegorz Pacyna, Sławomir Florjan & Robert Borzęcki

NEW MORPHOLOGICAL FEATURES OF ARTHROPLEURA SP.

(MYRIAPODA, DIPLOPODA) BASED ON NEW SPECIMENS FROM

THE UPPER CARBONIFEROUS OF LOWER SILESIA (POLAND)

1

De part ment of Palaeobotany and Palaeoherbarium, In sti tute of Bot any, Jagiellonian Uni ver sity, Lubicz 46, 31-512 Kraków, Po land, e-mail: grzegorz.pacyna@uj.edu.pl

2

De part ment of Palaeobotany and Palaeoherbarium, In sti tute of Bot any, Jagiellonian Uni ver sity, Kopernika 27, 31-501 Kraków, Po land, e-mail: slawomir.florjan@uj.edu.pl

3

Mu seum of Min er als, Rob ert Borzêcki, Czeska 15, 57-303 K³odzko, Po land, e-mail: min er als@redbor.pl Pa cyna, G., Flor jan, S. & Borzêcki, R., 2012. New mor pho logi cal fea tures of Ar thropleura sp. (Myriapoda, Di plo poda) based on new speci mens from the Up per Car bon if er ous of Lower Sile sia (Po land). An nales So cie ta tis Ge olo go rum Po lo niae, 82: 121–126.

Ab stract: New ter gite frag ments of Ar thropleura were found at the Nowa Ruda mine (Lower Sile sia, Po land),

af ter more than sev enty years since any pre vi ous dis cov ery. The small di men sions of the pre served struc tures, in terms of the ge nus Ar thropleura, and the char ac ter is tic fea tures of tu ber cu la tion proba bly in di cate that the re mains do not be long to the type spe cies A. ar mata, but they could rep re sent a new spe cies. The oc cur rence of round pro tru sions in the broken- off tu ber cles is an other fea ture of the new speci mens, which has not been men tioned in the ear lier lit era ture. Un for tu nately, the scanty re mains that we have at our dis posal (only two speci mens were found) are not suf fi cient for the pro posal of a new spe cies. The number of valid Ar thropleura spe cies re quires fur ther in ves ti ga tion, es pe cially re gard ing the fea tures, which are di ag nos tic for spe cies. The ter gites de scribed have very nu mer ous tu ber cles, which may have taxo nomic value in spe cies dis crimi na tion. Be cause the fos sils were found on a dump, their ex act stra tigraphic po si tion is not known. How ever, they oc cur to gether with in dex leaf lets of the seed ferns Pa rip teris gi gan tea and Li nop teris sp., which en abled the age de ter mi na tion of Up per Na murian - Lower West phalian for the fos sils stud ied. This new dis cov ery of Ar thropleura con trib utes to a bet ter un der stand ing of the ge nus and of the Car bon if er ous land fauna of Po land, which is oth er wise poorly known. This is also the first, de tailed de scrip tion of Ar thropleura re mains from the Pol ish Car bon if er ous.

Key words: Ar thropleura, Up per Car bon if er ous, Lower Sile sia, Po land.

Manuscript re ceived 5 January 2012, accepted 4 September 2012

IN TRO DUC TION

-log i cal his tory of the Earth reach ing 2–3 m in lenght (Braddy et al., 2008). This is the rea son for the con tin ual re -search on it (Meyer, 1853; Jor dan and Meyer, 1854; Kliver, 1883, 1884; Pruvost, 1930; Waterlot, 1934; Guthörl, 1936, 1938; Rolfe, 1969, 1985; Don o van, 2002; Kraus and Brauckmann, 2003). Its sys tem atic po si tion among ar thro pods is still un der dis cus sion (Rolfe and Ingham, 1967; Al -mond, 1985; Kraus and Brauckmann, 2003), but at pres ent the over rid ing opin ion is that it should be as signed to the Diplopoda (Kraus and Brauckmann, 2003; Kraus, 2005; Shear and Edge combe, 2010). How ever, there are doubts con cern ing its kin ships among Diplopoda and its ex tinct rel -a tives, Eo-arthropleurid-a -and Microdecemplicid-a (Kr-aus -and Brauckmann, 2003; Kraus, 2005). The best pre served re mains of Arthropleura have been found in Up per Car bon if -er ous and Low-er P-erm ian strata (Guthörl, 1940; Hahn et al.,

1986; Brauckmann et al., 1997; Hannibal, 1997a, b; Schnei -der and Barthel, 1997; Proc tor, 1998; Schnei der and Werneburg, 1998; Kraus and Brauckmann, 2003). Some older, but doubt ful re mains, found in Up per De vo nian strata, have also been re ferred to Arthropleura (Al mond, 1985; Ross and Briggs, 1993). The tracks of this gi gan tic an i mal are well known and have been de scribed fre quently (Briggs et al., 1979, 1984; Ryan, 1986; Pearson, 1992; Don -o van, 2002; Morrissey and Braddy, 2004; Lucas et al., 2005; Hannibal, 2007; Pol lard et al., 2008; Martino and Greb, 2009; Schnei der et al., 2010).

Arthropleura spec i mens have been found in the Lower

Silesian Coal Ba sin since the 19th cen tury, as in other Car -bon if er ous coal bas ins in Eu rope. These finds have contri-buted to a better un der stand ing of the ge nus (Roemer, 1880; Guthörl, 1936, 1938), but they have never been prop erly de -scribed or il lus trated. The spe cies Arthropleura armata, A.

maillieuxi, Arthropleura sp. were de ter mined from Lower

Silesia, and Arthropleura sp. from Up per Silesia (Gothan and Gropp, 1933; Guthörl, 1936; Hoehne, 1948). The ge nus

Arthropleura is also well known from the Czech area of Up

-per Silesia and Lower Silesia (Øehoø and Øehoøová, 1972; Štamberg and Zajic, 2008). Arthropleura armata and A.

maillieuxi were de scribed from the “Nowa Ruda” mine (Guthörl, 1936). Since Hoehne’s work (1948), there have been no new data on Arthropleura from Po land. In the 1950s and 1960s, Sagan col lected nu mer ous, well pre served spec i mens of Arthropleura sp. from Lower Silesia, but he pub lished only a brief note about his find ings (Sagan, 1967). The Sagan col lec tion is cur rently be ing re searched, and the ini tial re sults have been pre sented by Zdebska et al. (2005). Kiepura et al. (2001) col lated the in for ma tion, pub -lished thus far, about Po land’s Car bon if er ous land fauna, but the ge nus Arthropleura was not men tioned. The only spec i men of a Car bon if er ous myria pod, de scribed in de tail from Po land (Hannibal and Krzemiñski, 2005; Zdebska et

al., 2005), be longs to the ge nus Palaeosoma (Diplopoda).

The new spec i mens dis cussed be low were found on a spoil heap at the “Nowa Ruda” mine. Though only two spec i mens were found, they have con trib uted to a better un der stand ing of the ge nus Arthropleura and the Car bon if er -ous land fauna in Po land gen er ally.

MA TE RIAL AND METH ODS

Two frag ments of ter gites were found by R. Borzêcki on a spoil heap at the “Nowa Ruda” mine (Fig. 1) in 2010 (Lower Silesia, Po land) and are stored at the Muzeum Minera³ów, Rob ert Borzêcki, in K³odzko un der the cat a log

num bers MMRB2010.06.16/527/0.00/a-b and MMRB

2010.07.25/564/0.00. They are pre served in black, coaly shale, to gether with some plant re mains. Coal min ing has now ceased in the Pol ish area of the Lower Silesia Coal Ba sin (Bossowski and Ihnatowicz, 2006). There fore, new fos -sils can only be found on spoil heaps, in drill core and at a few nat u ral out crops, many of which are now in ac ces si ble.

As the fos sils were dis cov ered on a spoil heap, their ex act strati graphic po si tion is not known (Fig. 2). For tu nately, some de ter mi na ble plant re mains ac com pany the

Arthro-pleura re mains. In dex leaf lets from the seed ferns Paripteris gigantea (Stern berg, 1823) Gothan, 1941 (Fig. 3) and Linopteris sp. have been rec og nized, and these per mit an age de

-ter mi na tion for the fos sils stud ied as Up per Namurian– Lower Westphalian (Gothan, 1941; Kotasowa and Migier, 1995) (Fig. 2).

The spec i mens were an a lyzed un der a Nikon stereomicroscope SMZ 1000. In the spec i men de scrip tions, we fol -low the an a tom i cal terms, pro posed by Kraus and Brauck-mann (2003, fig. 1).

SYS TEM ATIC PA LE ON TOL OGY

Phy lum ARTHROPODA Latreille, 1829 Super class MYRIAPODA Latreille, 1796 Class DIPLOPODA de Blainville (in Gervais, 1844)

Or der ARTHROPLEURIDA Waterlot, 1934 Fam ily ARTHROPLEURIDAE Zittel, 1885

Ge nus Arthropleura Meyer, 1853 Type spe cies: Arthropleura armata Meyer, 1853

Arthropleura sp.

Fig. 3A–C

Syntergite (spec i men no. MMRB 2010.07.25/564/0.00)

Syntergite frag ment (only pos i tive pre served, vis i ble from the dor sal side of the body), with the di men sions 28 mm by 21 mm (Fig. 3A), and roughly 20 traces of small tu -ber cles, vis i ble on the sur face, with di am e ters at the base from 0.5 mm to 3 mm. These tu ber cles are ir reg u larly dis -trib uted. Only one tu ber cle is clearly larger, and is 6 mm in di am e ter at the base. Some tu ber cles have bro ken-off tips and are trun cated, be cause of this. Plant de tri tus is as so ci -ated with the syntergite, with only leaf lets of the seed fern

Paripteris gigantea de ter mi na ble (Fig. 3D).

Fig. 1. Lo ca tion of spoil heap at Nowa Ruda Fig. 2. Lithostratigraphic scheme of Nowa Ruda area (ac cord ing Nowak, 1995, mod i fied), show ing prob a ble po si tion of ho ri -zon with Arthropleura spec i mens

The spec i men was iden ti fied as a syntergite frag ment, be cause of its pre served, some what rect an gu lar shape (syntergite edges are not clearly vis i ble) and clearly vis i ble tu ber cles, di rected to the rear (Hahn et al., 1986; Brauck-mann et al., 1997; Schnei der and Barthel, 1997; Schnei der and Werneburg, 1998; Kraus and Brauckmann, 2003). The oc cur rence of only one large tu ber cle on the syntergite is an in ter est ing fea ture of the fos sil de scribed here. In spec i mens de scribed in ear lier pa pers, there are more large tu ber cles on syntergites and they form an al most reg u lar line near the rear edge of the syntergite (Waterlot, 1934; Hahn et al., 1986; Brauckmann et al., 1997; Kraus and Brauckmann, 2003). Here, too, the large tu ber cle is near the rear edge of the syntergite.

Paratergite (spec i men no. MMRB 2010.06.16/527/0.00/ a-b)

An in com plete paratergite frag ment (pos i tive and neg a tive pre served) with di men sions of 44 mm by 22 mm, vis i -ble from the dor sal side of the body. The whole paratergite sur face has vis i ble traces of over 70 tu ber cles from 0.6 mm to 3 mm in di am e ter at the base and a line of large tu ber cles, 6 mm in di am e ter at the base (Fig. 3B, C). Small tu ber cles are dis trib uted over the whole paratergite sur face at rel a -tively even dis tances from each other; smaller ones oc cur more densely and larger ones more sparsely. Some tu ber cles have bro ken-off tips and are trun cated, be cause of this. The tu ber cles with bro ken-off tips have round ish struc tures,

located cen trally in side them. The im print of the leaf let

Lino-pteris sp. oc curs with the fos sil, to gether with plant de tri tus.

The spec i men was iden ti fied as a paratergite frag ment, be cause of its shape, which is typ i cal for this struc ture (Hahn et al., 1986; Brauckmann et al., 1997; Schnei der and Barthel, 1997; Schnei der and Werneburg, 1998; Kraus and Brauckmann, 2003). Par tic u larly char ac ter is tic is the para-ter gite’s arched shape. The small di men sions and weak arch ing of the paratergite de scribed al low us to sup pose that it co mes from the front or, more prob a bly, the rear sec tion of the an i mal, be cause paratergites are pro por tion ally larger from the cen tral sec tion, and also strongly arched (Hahn et

al., 1986; Brauckmann et al., 1997). An in ter est ing fea ture

of this paratergite is that the large tu ber cles do not form a reg u lar line, but fol low a stretched let ter S. In the

Arthropleura spec i mens, de scribed from Ger many, the large tu ber

-cles form a clear line (Hahn et al., 1986; Brauckmann et al., 1997; Schnei der and Barthel, 1997; Schnei der and Werne-burg, 1998; Kraus and Brauckmann, 2003). This fea ture is par tic u larly vis i ble on small Arthropleura spec i mens from the for mally un des cribed spe cies from Montceau-les-Mines (Kraus and Brauckmann, 2003; Kraus, 2005). In con trast to those spec i mens, large tu ber cles, fol low ing the shape of a stretched let ter S, are vis i ble on some French Arthropleura

armata spec i mens, which were il lus trated by Waterlot (1934). How ever, this fea ture is more strongly ex pressed in the ma te rial from Silesia. The oc cur rence of round pro tru

-Fig. 3. Ter gite frag ments of Arthropleura sp. and as so ci ated plant fos sil. A. Syntergite frag ment (MMRB 2010.07.25/564/0.00, pos i -tive spec i men). B. Right paratergite frag ment (MMRB 2010.06.16/527/0.00/a, pos i -tive spec i men). C. Right paratergite frag ment (MMRB 2010.06.16/527/0.00/b, neg a tive spec i men). D. Leaf let of Paripteris gigantea (Stern berg, 1823) Gothan, 1941 (MMRB 2010.07.25/ 564/0.00)

sions in the bro ken-off tu ber cles is an other es sen tial fea ture of this spec i men, which has not been men tioned pre vi ously in the lit er a ture. How ever, this fea ture is vis i ble on some spec i mens, pre vi ously il lus trated by Brauckmann et al. (1997, tab. 2, fig. 2). The fea ture is also vis i ble on other spec i mens of Arthropleura from the Lower Silesian Coal Ba sin in the Sagan col lec tion (Wa³brzych Mu seum). The real na ture of this fea ture re quires fur ther in ves ti ga tion. At pres ent, we are un able to de cide whether it is a taphonomic fea ture or an or ganic fea ture of the tu ber cles.

Arthropleura fos sils with nu mer ous and size di ver si fied

tu ber cles, as in our spec i mens, were de scribed and il lus -trated by Rößler and Schnei der (1997), and they de ter mined them to be Arthropleura armata Meyer, 1853. The spec i -mens, de scribed and il lus trated by Hahn et al. (1986) and Brauckmann et al. (1997) as Arthropleura armata, have less nu mer ous tu ber cles. If the num ber, size and ar range ment of tu ber cles are tax o nom i cally im por tant for dis tin guish ing spe cies of Arthropleura, as Kraus and Brauckmann (2003) sug gest, the spec i mens de scribed by Rößler and Schnei der (1997) may not be long to the spe cies Arthropleura armata. The fos sils, de scribed by Rößler and Schnei der (1997), have large tu ber cles, ar ranged dif fer ently on the paratergite, by com par i son with these spec i mens from the Lower Silesian Coal Ba sin. In their spec i mens, tu ber cles are lo cated prob a bly very near the rear edge of the paratergite, al -though this is not cer tain, be cause those spec i mens are not well pre served.

The fos sils from the Lower Silesian Coal Ba sin have tu -ber cles, dis posed the nearer back end of paratergite, but their ar range ment in the shape of the let ter S means that they ex tend to as much as half of the width. The spe cies A.

maillieuxi Pruvost, 1930, A. mammata (Salter, 1863) and A. bri tan nica Andrée, 1910 have a prom i nent, straight sim ple

line of large tu ber cles on the paratergites, which dis tin -guishes them from the ma te rial de scribed (Salter, 1863; Wood ward, 1872; Guthörl, 1936). In A. bri tan nica, the shape of the paratergite is sim i lar to the shape in the fos sil de scribed here. How ever, it is con sid er ably larger in this spe cies (Guthörl, 1936). The spec i mens from Spain, de ter -mined as A. armata, have tu ber cles on the paratergite, which are small and of di men sions be tween small and large and lack tu ber cles that are clearly large (Cas tro, 1997). In the Arthropleura cristata Rich ard son, 1959 from West-phalian D of Mazon Creek, small tu ber cles were only found in the front sec tion of the paratergite; while very large tu ber -cles cover the rest of the paratergite. The paratergite of this spe cies is also con sid er ably larger than the spec i men de -scribed (Rich ard son, 1959; Hannibal, 1997a, b).

DIS CUS SION AND CON CLU SIONS

This new dis cov ery of Arthropleura con trib utes to a better un der stand ing of the Car bon if er ous land fauna of Po land, oth er wise poorly known. Only one, frag men tary spec i men of Arthropleura has been found, pre served within a si -der it ic nod ule from Sosnowiec (Up per Silesian Coal Ba sin), but it was mis iden ti fied as a eurypterid (Krawczyñski et al., 1997; Filipiak per sonal com mu ni ca tion 2012). It is ap pro

pri ate to note that these nod ules con tain the rich est Car bon -if er ous land and fresh-wa ter fauna, dis cov ered in Po land so far (Filipiak and Krawczyñski, 1996; Pacyna et al., 2004; Stworzewicz at al., 2009). Be cause the Arthropleura spec i -mens, noted ear lier from the Pol ish Car bon if er ous (Roemer, 1880; Gothan and Gropp, 1933; Guthörl, 1936, 1938; Hoehne, 1948), were not prop erly de scribed and il lus trated, com par i son with them is al most im pos si ble. Two paratergi-tes from Nowa Ruda, de ter mined as be long ing to

Arthro-pleura armata spe cies, were il lus trated by Andrée (1910).

They have larger di men sions than our spec i men. The tu ber -cles of those paratergites are on av er age big ger and the large tu ber cles form straight line.

The num ber of valid Arthropleura spe cies is a mat ter of de bate. It re quires fur ther in ves ti ga tion, es pe cially with re gard to the fea tures, which are di ag nos tic for spe cies. Pre vi -ously de scribed spe cies (all in the 19th and the be gin ning of the 20th cen tury) do not have clear, di ag nos tic cri te ria (Cal-man, 1914; Guthörl, 1936; Rich ard son, 1956, 1959; Rolfe, 1969). Hence, in the case of the ma te rial be ing de scribed, which is scanty (only two frag ments of ter gites), the spe cies was not de ter mined. The ter gites de scribed have very nu -mer ous tu ber cles, which may have tax o nomic value (Kraus and Brauckmann, 2003). The small di men sions of the pre -served struc tures, which are smaller than those of most spe-cimens, re ferred to Arthropleura armata (but see Waterlot (1934) for small spec i mens, re ferred to Arthropleura

armata). The char ac ter is tic fea tures of tuberculation prob a

-bly in di cate that the re mains do not be long to the spe cies A.

armata, but could rep re sent a new spe cies (Meyer, 1853;

Jor dan and Meyer, 1854; Kliver, 1883, 1884; Pruvost, 1930; Waterlot, 1934). The scanty re mains we have at our dis posal are not suf fi cient to sup port such a de ter mi na tion. The spec i mens may rep re sent a spe cies of Arthropleura with a com par a tively small size of in di vid u als, but it is worth not ing that even smaller spec i mens have been de -scribed that were as signed to the ge nus Arthropleura (Waterlot, 1934; Kraus and Brauckmann, 2003). The small spec i men of Arthropleura from MontceaulesMines (Mas -sif Cen tral, France) has long been in ter preted as a ju ve nile (Briggs and Al mond, 1984). How ever, Kraus and Brauck-mann (2003) and Kraus (2005) showed, with re gard to myria pod de vel op ment, that this is not a ju ve nile spec i men, but a dif fer ent spe cies, in which in di vid u als were par tic u -larly small. The small spec i men, de scribed from Eng land as a new spe cies, Arthropleura moyseyi Calman, 1914, is now rec og nized gen er ally as a ju ve nile spec i men of A. armata (Rolfe, 1969). It is worth not ing, how ever, that groups al lied to Arthropleura, the ex tinct Eoarthropleurida and Mi cro-decemplicida and the ex tant Pselaphognatha, are rep re -sented by small forms, sev eral mil li me ters or so in length (Shear and Selden, 1995; Kraus and Brauckmann, 2003).

We are un able to de cide whether the re mains de scribed rep re sent frag ments of exuviae or the an i mals them selves. The iso la tion of these rem nants may show that they are frag -ments of exuviae, but the lack of the fold ing, char ac ter is tic for exuviae (Cas tro, 1997; Kraus and Brauckmann, 2003), and the very clear im print of an un flat tened paratergite, pre -served (some what) in three di men sions, may show that these are parts of a once liv ing an i mal, and not exuviae.

The match ing rel a tive pro por tions and di men sions of the rem nants and also the sim i lar ity in their tuberculation may show that they are the re mains of in di vid u als of one spe cies. The close ness of their oc cur rence and very sim i lar man ner of pres er va tion might in di cate that they even are the rem nants of one an i mal.

Sin gle leaf lets from seed ferns oc cur with the

Arthropleura rem nants (Fig. 3D). These leaf lets are the dis in te gra

tion prod ucts of large, com pound leaves, which might in di cate that all the re mains were trans ported some dis tance be fore fos sil iza tion. The dam age to the syntergite might con -firm this.

This new dis cov ery of Arthropleura con trib utes to a better un der stand ing of that ge nus and of the Car bon if er ous land fauna of Po land, which is oth er wise poorly known. This is also the first de tailed de scrip tion of Arthropleura re -mains from the Pol ish Car bon if er ous.

Ac knowl edge ments

We of fer sin cere thanks to Pro fes sor D. Zdebska (Jagiellonian Uni ver sity, Cra cow) for dis cus sion and crit i cal com ment, which helped im prove this ar ti cle. C. Brauckmann (TU Clausthal), J. Hannibal (Cleve land Mu seum of Nat u ral His tory), R. Rö¢ler (Mu -seum für Naturkunde, Chemnitz) and P. Selden (Uni ver sity of Kan sas) sent re prints of their pa pers, pre vi ously un avail able to the au thors. Re views by J. Hannibal, W. Krzemiñski and P. Filipiak helped to im prove this pa per. We thank J. Madeja for help in the prep a ra tion of the draw ings.

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