Acta Palaeobotanica 52(2): 247–269, 2012

Carboniferous plants preserved within sideritic nodules – a remarkable state of preservation providing a wealth of information

GRZEGORZ PACYNA and DANUTA ZDEBSKA

Department of Palaeobotany and Palaeoherbarium, Institute of Botany, Jagiellonian University, Lubicz 46, 31-512 Kraków, Poland; e-mail: [email protected], [email protected]

Received 20 November 2012; accepted for publication 10 December 2012

ABSTRACT. Fossil plants preserved within sidertitic nodules have been known from Europaean and North American Carboniferous coal measures since the early 19th Century. However, only a few of them have been described thoroughly palaeobotanically, mainly in the 19th and early 20th centuries; thus their taxonomy often requires revision. Animal fossils preserved within sidertitic nodules beside plant fossils became a base of the description of many rare new taxa of animals undiscovered in other modes of preservation. Published hith- erto works about such preserved fl ora indicate that plants preserved within concretions represent rare taxa, not known in other modes of preservation. The dissimilarities in composition of fl oras preserved in ironstone concretions when compared to those from surrounding sediments likely results from the process of concretion formation which selectively infl uences the preservation of small, delicate plant organs. The most famous fl oras preserved within concretions come from Mazon Creek in the USA and Coseley in Great Britain. These locali- ties were the source of many previously unknown taxa with important evolutionary signifi cance. The new fl ora preserved within concretions has been discovered recently in Poland in Sosnowiec (Upper Silesia). It contains new, hitherto unknown taxa particularly plant reproductive organs. Comparison of the taxonomy and tapho- nomy of the ironstone concretion fl ora from Sosnowiec with other similar assemblages from the Carboniferous of Europe and North America has reveals many similarities steming from a common mode of preservation. Due to the exceptional three-dimensional preservation of the plant fossils, in particular reproductive organs key to the understanding of evolutionary relationships, the locality at Sosnowiec can be regarded as an important new Lagerstätte, and the fi rst such site recognized in Poland. The use of pioneering techniques in high-resolution X-ray microtomography promises to yield yet further information on the biota of this new locality.

KEYWORDS: plant fossils preserved within sideritic nodules, Upper Carboniferous, Sosnowiec, Coseley, Mazon Creek, Fossil- Lagerstätten, Konservat-Lagerstätten

INTRODUCTION

The occurrence of exceptionally well pre- animals, which otherwise are seldom found in served fossils within ironstone concretions a good state of preservation (Garwood & Sut- accompanying Carboniferous coal seams of ton 2010, Garwood et al. 2009, Legg et al. coal-fi elds of Europe and North America has 2012). As well as marine animals, especially been known and described since the 19th Cen- these with a very low fossilization potential, tury (Brongniart 1828–1837). Few of these such as jellyfi shes, hydroids, polychetes etc. specimens were subjected to detailed palaeo- (Nitecki 1979, Shabica & Hay 1997). Some ani- botanical analysis by modern standards and mal fossils have also been described from the techniques. Greater attention was paid to Sosnowiec nodules (Filipiak & Krawczyński the fauna appearing in these concretions and 1995, 1996, Krawczyński et al. 1997, Pacyna there are more papers on this theme (Nitecki et al. 2004, Stworzewicz et al. 2009, Prokop 1979, Shabica & Hay 1997). Sideritic nod- et al. 2012). Carboniferous ironstone concre- ules often contain splendidly preserved land tions create a taphonomic window permitting 248 the recognition of the fauna and the fl ora to volumes of organic material (e.g. plant mate- a degree comparable only with the best palae- rial), the basin water formed reducing condi- ontological sites from other geological epochs, tions above a consolidated sediment base. The e.g. Burgess Shale, Chengjiang, Hunsrück bacterial decay of organic debris quickly laid Slate, Solnhofen Lithographic Limestones, etc. down in the loamy sediment locally changed (Baird 1990). For these reasons, Carboniferous the reaction of the environment around them, localities with fossil preserved within sideritic in special cases enabling the precipitation of nodules are universally recognized as Fossil- the ironstone (Chodyniecka 1973, Woodland Lagerstätten (Baird 1990, Baird et al. 1985a, & Stenstrom 1979, Sellés-Martínez 1996). b, 1986, Schellenberg 2002, Nudds & Selden Then the ironstone concretion formed quickly 2008). The use of new research techniques and protected the organic remains, terminating such as high-resolution X-ray microtomog- decomposition and protecting the resultant fos- raphy reveals new, previously undiscovered sil from compaction through sediment loading. details in such fossils (Garwood & Sutton 2010, Ironstone concretions form syngenetically Garwood et al. 2009, Legg et al. 2012, Spencer with the sediments in which they appear. In et al. 2013). sphaerosiderites the early diagenesis preced- In the 1990s, on the dump of the “Porąbka- ing the compaction provides the opportunity Klimontów” coal mine in Sosnowiec-Zagórze for the three-dimensional preservation of the (Upper Silesian Coal Basin) sideritic concre- fossil (Baird 1997). The mode of preservation tions have been found containing plant and of plants in concretions belongs to the type of animal fossils (Filipiak & Krawczyński 1995, preservation of fossil called authigenic cemen- 1996, Krawczyński et al. 1997, Pacyna 2002a, tation (Schopf 1975, 1979). A very early cemen- b, 2003, 2004, 2007a, b, c, 2008a, b, Pacyna tation in soft sediment by iron and carbonate & Wojciechowski 2004, Pacyna et al. 2004, compounds preserves the surface confi guration Pacyna & Zdebska 2001, 2002, 2005, 2010, of organic parts, even very delicate structures. Stworzewicz et al. 2009, Prokop et al. 2012). Thanks to this the three-dimensional shapes Though ironstone concretions are frequent in of plant morphological-anatomical details are rocks accompanying coal seams in Upper Silesia recorded in exquisite detail. Most soft tissues, (Porzycki 1972) and they were exploited even even if orginally present, are postdiageneti- as source iron ores (Tryjefaczka 1982), regret- cally degraded, and the spaces which remain tably the fossils preserved within them were after them could be secondarily fi lled, e.g. with mentioned earlier only sporadically (Feistman- pyrite or calcite. Of the original organ of the tel 1874, Roemer 1866, Bocheński 1939), and plant only the coal fi lm on walls of the three- a few described fossils (only animals) are now dimensional gap most often remains. This gap lost. This new locality in Sosnowiec provides is often secondarily fi lled with kaolinite which an opportunity to describe completely such covers many details and makes the investiga- preserved fossil assemblage for the fi rst time tion of specimens very diffi cult. The genesis of in Poland. this mineral in concretions has not as yet been The purposes of this article are the discus- explained (Baird et al. 1985b). sion of taxonomical composition and taphonom- ical aspects of fossil fl oras preserved within sid- eritic nodules and comparison between these REVIEW OF THE UPPER fl oras and the new fl ora uncovered in Sosnow- CARBONIFEROUS FLORAS PRESERVED iec alongside discussion on future directions WITHIN IRONSTONE CONCRETIONS in the study of such exceptionally preserved OF EUROPE AND NORTH AMERICA fl oras. Carboniferous fl oras preserved within sider- itic nodules are rare from a global perspec- THE GENESIS OF IRONSTONE tive. Though at least several fl oras preserved CONCRETIONS within ironstone concretions are known from the Upper Carboniferous strata of Europe and The formation of ironstone concretions is North America, their modern complex taxo- tied inseparably with the water environment. nomic and taphonomic descriptions and inter- As a consequence of the accumulation of large pretations are lacking. Every locality of such 249 exceptional preservation invokes broad inter- Mazon Creek est from palaeobotanists, because the plants preserved in them provide a great deal of The Mazon Creek locality is situated in the valuable information. The most famous, rich- north-east part of Illinois. It belongs among est and best recognized locality with Carbonif- the most famous palaeontological localities in erous plant fossils preserved within ironstone the world, fi rst of all because of its unusually concretions is Mazon Creek in Illinois, USA. preserved softbodied fauna (Richardson 1966, In spite of the fact that this locality has been Schram & Nitecki 1975, Nitecki 1979, Baird known since the second half of 19th Century, et al. 1885a, b, Baird et al. 1986, Baird 1990, and has been described in many palaeobo- Shabica & Hay 1997, Schellenberg 2002, Wil- tanical papers (Lesquereux 1866, 1870, 1879, son 2006, Bethoux 2009). Though the fl ora 1880, 1884, Noe 1925, Schopf 1938, 1941, from this locality is as equally well known as Janssen 1940, Langford 1958, 1963, Darrach the fauna, it still awaits a full modern descrip- 1970, Horowitz 1979, Pfefferkorn 1979), to this tion (Lesquereux 1866, 1870, 1879, 1880, 1884, day new taxa of plants from Mazon Creek are Noe 1925, Darrah 1936, 1937, 1938, Janssen still being described, often later recognized 1940, Langford 1958, 1963, Darrach 1970, to be unique to this locality (Pigg & Taylor Peppers & Pfefferkorn 1970, Horowitz 1979, 1985, Drinnan & Crane 1994). Several simi- Pfefferkorn 1979, Pigg & Taylor 1985, Drinnan lar, though signifi cantly taxonomically poorer & Crane 1994). Ironstone concretions are found localities from North America still await full in the Francis Creek Shale Member, overlying elaboration (Baird et al. 1985b). A review of the Colchester (No. 2) Coal seam. According to these localities primarily focusing on the fauna North American Carboniferous stratigraphy, was made by Baird et al. (1985b). A compre- the age of the concretions is Middle Pennsylva- hensive review of European localities is cur- nian – Desmoinesian Series, which correlates rently lacking. In terms of European localities, with Westphalian D according to the West- ironstone concretion lagerstätte have been European division. primarily scrutinised for their faunal content The fossil assemblage at Mazon Creek pre- (Moysey 1910, Anderson et al. 1997, 1999). serves over three hundred species of animal The mode of preservation in ironstone con- taxa and as many plants are also known. The cretions presents a range of problems when fossil remains represent two kinds of environ- trying to extract information and interpret fos- ments: swampy, forested delta plains referred sils. Recently however, advances in the use of to as ‘Braidwood’ after the typical locality of high resolution X-ray tomography in the study occurence and shallow waters of sea surround- of faunal and some fl oral remains preserved in ing the delta referred to as ‘Essex’ type envi- such concretions has allowed the extraction of ronments and biotas. The terrestrial and fresh- much more information from these exceptional water deposits include plants, and also insects, fossils (Garwood & Sutton 2010, Garwood et al. myriapods, spiders, scorpions, fi sh, amphibians 2009, Legg et al. 2012, Spencer et al. 2013). and reptiles. The marine assemblage contains Floras preserved within sideritic nodules remains of jellyfi shes, polychaete worms, mol- from North America and European Pennsyl- lusca, sea cucumbers and crustacea, as well as vanian localities are discussed and compared pelagic cephalopods and fi sh (Schram & Nitecki below. 1975, Nitecki 1979, Shabica & Hay 1997). The fl ora preserved within ironstone con- NORTH AMERICA (USA) cretions from the Mazon Creek area has been known since the beginning of coal mining activ- Illinois ity there in 1855 (Horowitz 1979). The fi rst Numerous well preserved fl oras from shales collection gathered there was sent for investi- and coal balls are known from the Middle Penn- gation and study to Leo Lesquereux who pub- sylvanian Carbondale Formation, Kewanee lished the fi ndings in several extensive works Group, Illinois. Horizons containing ironstone (Lesquereux 1866, 1870, 1879, 1880, 1884), concretions are particularly frequent in this which were classical texts in North American formation and several such preserved fl oras palaeobotany. Since Lesquereux’s work has not have been described from them, particularly appeared in full, professional revisory elabo- the famous Mazon Creek fl ora. ration of this fl ora, though it is known that 250 nomenclature used by Lesquereux is currently leaves for ferns, the foliage for seed ferns and outdated; numerous new specimens have been ovules for cordaites. Pfefferkorn concluded that found, and many of Lesquereux’s taxonomic the minimum-number of recognized natural determinations are incorrect and require veri- taxa in Mazon Creek equals 96 species belong- fi cation. ing to 41 genera. In the 20th century, the Mazon Creek fl ora Pfefferkorn (1979) mentioned only the was illustrated and very briefl y described in most frequently appearing genera and con- several popularized scientifi c works for ama- centrated on the evaluation of the age of the teur collectors (Noe 1925, Janssen 1940, Lang- fl ora, variously given earlier. Based on recog- ford 1958, 1963). These works are now known nition of index species of the macrofl ora (the to contain some errors and do not bring new occurrence Neuropteris ovata together with data to the knowledge of this fl ora. Only Dar- more than ten species of Pecopteris, and also rah (1970) in his revision of the Upper Penn- the occurrence of Stephanian aged species sylvanian fl ora of the USA devoted more space higher in the profi le) he found that this fl ora to the Mazon Creek fl ora, but concentrated parallelled fl oras of Upper Westphalian D in predominantly on common species and strati- Europe. graphically useful taxa. In the second half of the 20th Century there The latest summary of this fl ora is the work appeared several short works revising or of Pfefferkorn (1979). According to Pfefferkorn describing single new taxa from Mazon Creek. the Mazon Creek fl ora is characterized by the These materials were also taken into account large number of species, with a large variabil- in descriptions of the fl ora at large or within ity in size of the most frequent species and descriptions of particular systematic groups: excellent preservation. The large diversity of Abbott 1958 (Asterophyllites, Annularia and taxa is related to the variety of subenviron- Shenophyllum from Mazon Creek), Pfefferkorn ments in the deltaic setting. Plants were pre- 1976 (he described and illustrated stem speci- served as fossils as result of quick submersion mens of arborescent ferns belonging to genera during storms, when the sea fl owed over the Caulopteris, Megaphyton and the new genus area of the delta. This also caused consider- Artisophyton), Pigg and Taylor 1985 (descrip- able mixing of disarticulated plant debris. The tion of a new stem base of the lycopsid – Cor- Mazon Creek fl ora belongs to the taxonomically mophyton mazonensis). richest Pennsylvannian fl oras in North Amer- The separate chapter in research of the ica (Peppers & Pfefferkorn 1970, Pfefferkorn Mazon Creek fl ora constitutes monographic 1979). This results not only from intensive col- elaborations of diversifi ed plant reproduc- lecting of specimens, but also from favourable tive organs of different, numerous and well conditions for plants of during their life and preserved taxa. These monographs appeared taphonomic processes favoring fossil preserva- from the beginning of the 20th century (e.g. tion. It is interesting to note the occurrence Sellards 1903) and were based on single speci- of the gigantism of some plant forms whose mens from the huge (counting above 16 thou- fossils outside Mazon Creek are signifi cantly sand concretions with plant fossils) collection smaller. deposited in the Illinois State Museum and The number of described species of plants the Field Museum of Natural History (Photos from the Mazon Creek locality is over three of specimens from the Illinois State Museum hundred; however, this fi gure infl ates the collection are available on the website http:// diversity artifi cially since many of these spe- www.museum.state.il.us/databases/geology/ cies represent isolated organ morphotaxa (Pep- mazoncreek, unfortunately most often still pers & Pfefferkorn 1970, Pfeffernkorn et al. with Lesquereux’s designations). 1975, Pfeffernkorn 1979). Pfeffernkorn made Based primarily on materials from Mazon an attempt to evaluate the real diversity of the Creek (Lesquereux specimens), Chaloner (1958) Mazon Creek fl ora. For every group of plants carried out revisions of the genus Polysporia. he chose the organ which showed the greatest Kosanke (1955) described Mazostachys pen- diversity, and also appeared often in concre- dulata, a fragment of stem and new genus tions and provided a relatively large amount of calamite cone known from a concretion. of taxonomical information. These are stems Good (1975) again described this specimen for lycopods, the foliage for sphenopsids, fertile and recognized it as a basis for a new genus 251

– Mazostachys and referred to this genus as Carterville yet another species coming from Francis Creek Shale – M. noei. The second famous fl ora from the Carbon- Taylor (1967) described the new species of dale Formation preserved within the ironstone the fertile leaf of fern – Radstockia kidstonii. concretions was discovered in the town of Pfefferkorn et al. (1971) described extremely Carterville in the south of the state of Illinois. rich and splendidly preserved specimens of fer- The concretions appeared in the Energy Shale tile fern leaves, isolated spores in situ from the and Anna Shale Members, over the coal seam sporangia, illustrated and compared them with Herrin (No. 6) Coal and are dated from the dispersed species. Pfefferkorn (1973) described Middle Pennsylvanian – Desmoinesian Series, buds serving the vegetative reproduction of the they appear to be higher in the profi le of the fern as the new genus and species – Kanka- Carbondale Formation rather than the con- keea grundyi, he also revised similar speci- cretions from the Mazon Creek and are some- mens of Lesquereux, which should be referred what younger than them as well, furthermore to this new taxon. Jennigs and Millay (1979) according to the West-European division also described rich fertile material of Pecopteris belong to the Westphalian D. unita. This species is common in the Mazon The fl ora preserved within Carterville con- Creek fl ora. Owing to the use of the combina- cretions was already known about since 1875. tion of different techniques e.g. sectioning of One of the specimens from these concretions the concretion in suitable planes, followed by recognized as Dolerotheca sp. was described polishing and subsequently acetate peeling the by Schopf (1948), while another specimen, of sectioned blocks providing information on the calamite cones, was described in 1965 by Hib- anatomical structure of synangia, sporangia bert and Eggert and discovered to be a new and spores. On the basis of the spore morphol- species – Paracalamostachys cartervillei (Hib- ogy that was isolated, they found that this bert & Eggert 1965). The plant specimen pre- morphological species represented numerous served within this concretion consisted of an different biological species. axis onto which a dozen or so perfectly pre- Delevoryas (1964) described the new genus served cones were cyclicly arranged while still and species of the pollen organ of Medullosales preserving the spores within their sporangia. – Schopfi theca boulayoides on the basis on one Hibbert and Eggert (1965) isolated the spores specimen from the concretion, isolated from it from the sporangia in order to trace their vari- prepollen grains. Drinnan and Crane (1994) ability within the cones, which they recognized published a valuable work discussing pollen belonging to a dispersed species of Calam- organs of Medullosales from Mazon Creek fl ora ospora breviradiata. from the genera Codonotheca, Schopfi theca, In 1977 Gastaldo described a large collec- Dolerotheca and one not generically determi- tion of plants preserved within the ironstone nable pollen organ. In particular they isolated concretions from Carterville, which originated from pollen organs prepollen grains and pro- from the roof shales of Herrin (Nr. 6) Coal duced detailed illustrations of them. Drinnan (Gastaldo 1977). The collection that Gastaldo et al. (1990) described ovules Stephanosper- had at his disposal consisted of 2500 specimens mum konopeonus containing prepollen grains of concretions containing remains of plants: in the micropyle (such specimens enable the from which 1475 concretions were chosen for connection of pollen organs with their corre- study purposes, from these 24 genera and 52 sponding ovules). species of plants were determined. Schopf published two works (1975, 1979) Within the fl ora, the ferns from the orders devoted to the mode of preservation of plants in Filicales and Marattiales dominated numeri- ironstone concretions on the example of speci- cally (44% specimens were recognized as differ- mens from Mazon Creek, this special mode of ent species of the genus Pecopteris), medullo- preservation he named the type authigenic salean seed ferns constituted almost 25% of cementation. the fl ora (the foliage: Neuropteris, Alethopteris, A recapitulation of history of research of the Odontopteris, Callipteridium; the ovules Trigo- Mazon Creek fl ora and the full list of publi- nocarpus schutzianus), while the lyginoterid- cations until the year 1979 was published by alean seed ferns, which were very rare within Horowitz (1979). the assemblage (1%), were represented by only 252 one species called Renaultia (Sphenopteris) followed by its polishing and the application chaerophylloides. Amongst the relatively high of the acetate fi lm method, one could obtain number of sphenopsids, representatives from comparable level of information to that derived the order of Calamitales were mainly found from the study of coal balls. What is more, (the foliage Annularia, Asterophyllites; stems within sideritic concretions single large speci- of Calamites; 1 cone), while Shenophyllales mens are preserved, as opposed to a mixture of were proved to be rare in number (in other many different plants as observed within coal- words sphenopsids – 14% of the assemblage). balls. The ironstone concretions also delivered Isolated sporophylls dominated numerically valuable morphological information which amongst the lycopsids (10% of the assemblage), was diffi cult to obtain from coal-balls, such as and were determined as: Lepidostrobopsis cf. three-dimensional gross morphology of plant missouriensis, Lepidostrobophyllum lanceo- organs. latum, L. brevifolium, L. hastatum, L. fal- lax, L. cf. tumidum, and Lepidocarpon major Gorgetown (this species was particularly numerous and Ironstone concretions in the Carbondale constituted 5% of the entire assemblage). The Formation can also be found in yet another remaining 6% of the assemblage constituted location, this time in the central-eastern state a multitude of taxonomically differentiated of Illinois, near a town called Georgetown (Ver- thin leafl ess branches of lycopsids, determined milion Country). These concretions were found variously as several species of the genus Lepi- within the Energy Shale Member, above the dodendron, stems of Lepidophloios and Asola- Herrin (No. 6) Coal Member like those found nus, the leaves Lepidophylloides, the isolated in Carterville and likewise were dated from the sporangia Lepidocystis, the cones Lepidostro- Middle Pennsylvanian – Desmoinesian Series bus ornatus, the rootstocks Stigmaria fi coides (Westphalian D). The numerous specimens and the herbaceous lycopsids Lycopodites. were preserved in large concretions and are of Cordaites in turn were represented by one a taxonomically similar composition to those of specimen of Artisia. the Mazon Creek fl ora (Baird et al. 1985b). Three taxa of plant reproductive organs with exceptionally well preserved details as Indiana found within the fl ora, was the next subject of Terre Haute and other locations Gastaldo’s detailed works. In 1978 in collabo- ration with Matten (Gastaldo & Matten 1978) Concretions containing fossil assemblages they described the largest seed ever found (10 (fl ora and fauna) similar to those of the Braid- cms in length) in the Carboniferous strata of wood in Mazon Creek were discovered in the North America, which they recognized as the open cast coal mine near Terre Haute (Vigo new species Trigonocarpus leeanus. In 1981 Country) in Indiana (Baird et al. 1985b). They Gastaldo (Gastaldo 1981a) described the new appeared in the Busseron Sandstone Member species from the genus Palaeostachya – P. dir- of the Shelburn Formation, in shales above the cei. Gastaldo performed many cross-sections to coal seam Coal (No. 7) and were dated from the the concretion in the purpose of establishing Middle Pennsylvanian, which correlates with a detailed diagnosis of the new species. Subse- the Westphalian D. The fl ora found within this quently, Gastaldo (Gastaldo 1981b) described location and described by Boneham (1975) has the isolated sporangia of the lycopsid Lepido- much in common with the famous Mazon Creek cystis sp. containing in situ megaspores Valvis- Flora, especially abundant in Annularia stel- porites auritus, the specimen came from the lata, Ptychocarpus unitus, pecopterids with an Anna Shale Member. absence of sphenopterids. Aproximatelly 25% Jennigs and Millay (1978) described a new of the concretions from this location contained species – Scolecopteris macrospora, which were well preserved specimens. Within the fl oral found in the ironstone concretions of Anna assemblage, sterile organs predominate the Shale above the coal seam Herrin near Carte- area with lycopods stems and leaves, sphenop- ville. They demonstrated for the fi rst time that sid stems and leaves, fern and seed fern leaves. the use of suitable techniques in the study of Plant reproductive organs such as the cones ironstone concretions e.g. the sectioning of the of lycopsids and sphenopsids, fertile leaves of concretion in cross-sections of suitable planes, ferns, pteridosperm pollen organs were rare in 253 number, and belonged to the same taxa as in Baird et al. 1985b). Concretions were found in Mazon Creek. the shales above the coal seams of the Crowe- Ironstone concretions were also discovered burg Coal and New Castle Coal within the in numerous coal mines that exploited Pen- lowest part of the Verdegris Formation, and nylvanian coals in the state of Indiana. The were dated back to the Middle Pennsylvanian plant remains that were found in these con- – Desmoinesian Series, which correlated with cretions prevailed in numbers over animal the Westphalian D, they were however slightly fossils (Baird et al. 1985b). Canwright (1959) older than the Mazon Creek concretions (Bode described fl oras preserved within these concre- 1958). Bode (1958) was the fi rst to study this tions and also plant fossils from shales hasting fl ora, but the study was limited to listing taxa concretions. The fl oras are both taxonomically of the stratigraphically important seed fern rich and well preserved. leaves and some of the sphenopsid taxa. Crid- Wood (1963) described a fl ora contained pri- land and Darrah (1968) described only one marily within the ironstone concretions of the new species of the pollen organ – Crossotheca Stanley Cementery in Greene County, Indina. urbani preserved within the concretion, which This was a well preserved and taxonomically they found in a dump, their entire collection of rich fl ora dating back to the Early Pennsyl- nodules contained 1500 specimens. The fossil vanian – Westphalian B, with many plant assemblage (animal fossils were also found in reproductive organs, all beautifully preserved. this location) was very similar to the Braid- Wood (1963) noted the similarities between wood type community in the Mazon Creek this fl ora and the Mazon Creek fl ora, but these (Baird et al. 1985b). similarities likely resulted from the mode of preservation in sideritic nodules. Taxonomi- Knob Noster cally speaking the fl ora was similar to the Konob Noster (Johnson Country, Missouri) Westphalian A and B fl oras of Great Britain is relatively new Pennsylvanian site uncov- and Poland. This was especially visible in the ered in 1990 containing a fl ora and fauna pre- very similar morphology of the Lepidostrobus served within ironstone concretions (Hannibal cones, the lycopsid leafl ess twigs and stems, et al. 2003). Its fossil assemblage was found to and the sphenopsid cones. The species of seed be similar in composition to the Windsor and fern leaves were however typical for the West- Braidwood of Mazon Creek sites. The concre- phalian B fl oral assemblages. tions were found to originate from either the delta area or the estuary area while the plants Iowa were represented by the foliage of ferns, stems, The fl ora which was preserved within the seeds, cones and the trunks of arborescent ironstone concretions of the Pennsylvanian plants. Investigations of this site are in prog- strata of south-eastern Iowa (near Dunerath) ress and could potentially yield new fi ndings. was described by Spencer (1894) and Condit and Miller (1951). Only a few leaves of seed Oklahoma ferns and sphenopsid leaves from this site were Henryetta and Morris illustrated (Condit & Miller 1951). The fossil assemblage was found to be similar in com- From the mudstones and siltstones above the position to that found at Mazon Creek (Baird coal seam of Croweburg Coal, within the lower et al. 1985b). The fl ora included the same taxa part of Senora Formation (Middle Pennsylva- as in Mazon Creek and are of stratigraphically nian – Westphalian D) in the town of Henryetta equivalent ages (Westphalian D). and Morris (Okumlgee Country), eastern Okla- homa State, another site yielding concretions Missouri has been unearthed but is yet to be described. Preliminary recconaisance indicates that the Windsor fl ora is similar to that of the Braidwood biota This taxonomically rich and fossil abun- of Mazon Creek (Baird et al. 1985b). dant fl ora comes mainly from the ironstone concretions and also grey shales of the region Sallisaw of Windsor (Henry Country) in central-western Numerous plant and animal fossils from Missouri (Bode 1958, Cridland & Darrah 1968, another concretion assemblage have been 254 discovered in dark grey mudstones above the from the Coseley nodules were described by coal seam of Stigler Coal, the McAlester For- Kidston in earlier monographic papers about mation, the Cabaniss Group (the base of the selected taxa, as well as taxa from other sites Desmoinesian series, Middle Pennsylvanian, (Kidston 1903, 1905a, b, 1906a, b). In these Westphalian C) within the eastern Oklahoma papers the importance of this locality was State town of Sallisaw (Sequoyah Country). already recognized by Kidston’s description of This fl ora, which was different from the fl ora new species of seed fern reproductive organs. that was found at the Francis Creek Shale, is Specimens of this fl ora were gathered by Kid- yet to be described. Its assemblage of fauna is ston as well as other workers of the geological similar to Braidwood in Mazon Creek (Baird survey at the end of 19th century, most of the et al. 1985b). speciemens were splendidly preserved three- dimensionally within ironstone concretions, NORTH AMERICA (CANADA) especially differentiated reproductive organs, this became the basis for Kidston (1914) from Nova Scotia which he described a dozen or so new taxa. Within the fl ora preserved in ironstones, among Stellarton Basin the taxonomically differentiated representa- Sideritic concretions and atypical siderite- tives of ferns belonging to genera, the follow- dolomite coal balls with preserved anatomi- ing specimens were discovered: Sphenopteris, cally plants have been found in the Foord Boweria, Sphyropteris, Adiantites, Pecopteris, Seam (Stellarton Basin, Nova Scotia) in coal Dactylotheca, the fertile leaves of the Zeilleria and shale splits (Lyons et al. 1995, Zodrow avoldensis have also been recognized. et al. 1996, Zodrow & Cleal 1999). The plant The richest of taxonomical diversity can be tissue preservation was rather poor in sideritic found in the reproductive organs of seed ferns. nodules with only Stigmaria rootlets recogniz- Pollen organs were found in the numbers of able (Zodrow & Cleal 1999). The age of Foord several dozen specimens: 2 species of Crosso- Seam was estimated at Bolsovian (Middle theca, Telangium asteroides, a new genus and Pennsylvanian, Upper Carboniferous) based species of pollen organ Coseleya glomerata on palynomorphs. (description based on large material). New spe- cies of pollen organ – Neuropteris Carpentieri EUROPE (currently Potoniea carpenteri) was described based on specimen in which pollen organs were Great Britain connected with axes. Kidston also isolated pre- pollen grains from this same organ. Smaller Coseley and more simply built pollen organs that were Ironstone concretions accompany many not connected with axes were described as coal seams in Great Britain from the roofs of Whittleseya (?) fertilis (currently Boulayatheca which many are known to hold well preserved fertilis). Kidston illustrated the organs’ vari- plant fossils within sideritic nodules. There abilites and isolated in situ prepollen grains is only one fl ora however which was found for analysis. Ovules found in connection with to be preserved this way and it is yet to be the Neuropteris heterophylla leaves were described in details. Kidston (1914) described referred to this species. Other, isolated ovules the fl ora from the coal basin of Staffordshire, were described as the new species: Hexagono- which was located in the town of Coseley near carpus hookeri, Lagenostoma oblonga, L. (?) Dudley. Most of the ironstone concretions that urceolaris (fully preserved three-dimensionally were discovered came from the level Ten Foot specimen with all internal details), Polyptero- Ironstone above coal seam Thick Coal, while spermum orantum, Rhabdocarpus elongatus, the rest of the described specimens came R. renaulti, R. olivieri, R. wildi, Trigonocarpus from shales accompanying the coal seams. noeggerathi, Trigonocarpus sp., Tripterosper- The age of the fl ora was estimated at West- mum ellipticum, T. johnsonii. Within many phalian B (Lower/Middle Pennsylvanian). It concretions seed fern leaves were found, the was the fi rst time in Europe that a complete following genera were recognized: Mariopteris, description of a fl ora preserved within iron- Alethopteris, Aphlebia, Lonchopteris, Odontop- stone concretions was given. Some specimens teris, Neuropteris, Spiropteris. 255

Lycopsid cones were found: Lepidostrobus Bickershaw variabilis, L. triangularis, Sigillariostrobus In 1997 Anderson and co-authors reported sp., as well as cones connected with branches the discovery of a new site rich in both fl ora of Lepidodendron ophiurus. One of the most and fauna preserved in ironstone concretions remarkable specimens in the entire collection (Anderson et al. 1997). Nodules were found was a three-dimensional fully preserved small in the shales above the coal seam Haigh Yard fragment of lycopsid cone described by Kidston Coal in Bickershaw (Lancashire) and numerous as the new species Lepidocarpon westphali- plant remains were found within the ironstone cum. This specimen was preserved so perfectly concretions. Anderson et al. (1997) identifi ed that Kidston noticed even the integumented on the basis the Cleal and Thomas (1994) key sporangia (which were also well illustrated on the following genera and species: Alethopteris his photographs). In no material referable to decurrens, Annularia radiata, Sphenophyllum this genus, with the exception of specimens cuneifolium, Maripteris muricata, Stigmaria preserved within coal-balls, had anybody to fi coides, Cyperites, Lepidodendron, Lepidostro- date recognized integuments with such cer- bus, Trigonocarpus, Calamites, Neuropteris, tainty. The lycopsid stems that were found in Pecopteris and Sphenopteris. This assemblage the Coseley nodules were referred by Kidston of plants was typical for late Westphalian (1914) as belonging to the genera Sigillaria A (Lower Pennsylvanian). Plants within the and Bothrodendron. ironstone concretions were preserved mainly The sphenopsid remains contained within as coalifi ed remains. Only the Lepidostrobus concretions predominantly consisted of ster- cones were occasionaly piritized, and are also ile organs: Equisetites, Calamites, Annularia, sometimes fi lled with a calcite and a kaolinite. Asterophyllites, Sphenophyllum, Pinnularia, The concretions in Bickershaw were found on few cones belong to Calamostachys and Palae- a dump similar to the one from Sosnowiec. ostachya elongata. The cordaites were represented by the dif- Westhoughton ferent species of Cordaites leaves, Artisia piths, cones Cordaianthus volkmanni and Cordaian- In 1999 Anderson et al. (Anderson et al. thus sp., and ovules Samaropsis gutbieri. 1999) reported a discovery in the region of The specimens originating from the Cose- Westhoughton (Lancashire) of another well ley preserved in the ironstone concretions preserved site of plants and animals within were eventualy described and illustrated ironstone concretions, dating from the Upper- in different papers (Kidston 1923–1925, most Westphalian A (Lower Pennsylvanian). Crookall 1955–1976, Halle 1933, Stockmans A lower number of plant fossils however were & Williere 1961, Kurmann & Taylor 1984, found in these concretions in comparison to Taylor & Kurmann 1985), most of which those in Bickershaw, seed fern leaves were dis- contained Kidston’s specimens, along with covered to be very rare, with only the genus other specimens found at later times. Cleal Mariopteris being recognized. Surprisingly one and Thomas (1994) also illustrated the Cose- of the more common plants that were found ley concretions containing: Asterophyllites within the nodules was the Lepidostrobus cones. longifolius, Aulacotheca hallei, Boulayatheca Genera Trigonocarpus and Cyperites were also fertilis, Potoniea carpenteri. Cleal and Shute recognized. The location of this site was dis- (2003) in the monograph of the genus Lavein- covered in a working opencast coal-pit from eopteris illustrated a well preserved yet to be which the concretions were collected in situ recognized by Kidston, specimens of Cyclop- from shales above the coal seam Wigan Four teris orbicularis. Foot. Within these shales stems of Calamites, Currently animal fossils preserved within Stigmaria fi coides root systems with preserved sideritic nodules from Coseley are being exam- anatomical structure and standing trunks of ined intensively with the use of the newest Sigillaria were found. Paleobotanical analyses techniques, e.g. scanning using high-resolution of this site have yet to be undertaken. X-ray microtomography (μCT) (Dunlop 1999, Other British localities Wilson & Almond 2001, Wilson 2005, Garwood et al. 2009, Garwood & Sutton 2010, Garwood Besides the described specimens men- & Dunlop 2011, Legg et al. 2012). tioned above, a variety of Carboniferous plants 256 preserved within sideritic nodules have also found, from which 51 taxa were determined. been known to originate from British coal Within the plant fossils assemblage the spe- measures and were discovered as early as the cies Annularia stellata (59%, 4041 specimens) 19th century. Many of them were described dominated in number, with the remainder along with compression and impression fossils sphenopsids: Calamites, Calamostachys tuber- with added comments that they were preserved culata and Sphenophyllum constituted 10% within sideritic nodules (mostly exclusively in of assemblage. Ferns constituted 22% of the plate captions). assemblage, particularly numerous were the Special attention should be given to the magnifi cently preserved fertile specimens of holotype of the typical species of genus Lepi- different Pecopteris species. Seed ferns con- dostrobus – L. ornatus, preserved within an stituted somewhat less than 6% of the assem- ironstone concretion from the Upper Carbonif- blage, and were represented by only their foli- erous of Derbyshire. This specimen was illus- age. Some specimens were found to be fertile trated for the fi rst time by Parkinson in 1804, (Dicksonites pluckenetii). Lepidodendralean and then described by Brongniart (1828–1838), lycopsids constituted 3% of the assemblage, while furthermore revised by Brack-Hanes and represented mainly by Cyperites leaves, poorely Thomas (1983) who isolated Lycospora granu- preserved trunks of Asolanus and Syringoden- lata spores from the specimen. dron and one tree-dimensionally preserved Cleal and Thomas (1994) illustrated the specimen of Sigillariostrobus. Cycadales Cordaitanthus sp. found preserved within the (Taeniopteris jejunata) and Cordaitales consti- concretion from above the coal seam Crow tuted each 1% of the assemblage. Particularly Coal in Crawcrook, the Rhyton, County Dur- interesting are the as yet indeterminate small ham (Westphalian B, Lower/Middle Pennsyl- specimens of Cordaitanthus cones. An attempt vanian). to isolate the spores from fertile organs has yet Cleal and Shute (2003) illustrated within to be undertaken. the monograph of the genus Laveineopteris, the well preserved in concretions speci- POLAND mens of Cyclopteris orbicularis from West- phalian B (Lower/Middle Pennsylvanian) of The fossils preserved within the ironstone Coalbrookdale, Shropshire. concretions that were found in Poland have been barely recognized, eventhough the site in Upper Silesia was already know to Roemer FRANCE (1866). From the concretions found in the area of Katowice, Roemer (1866) described a new Montceau-les-Mines genus and species of one of oldest spiders Pro- This well preserved fl ora from the Upper tolycosa anthracophila. In the same work he Stephanian (Upper Pennsylvanian) shales in also mentioned the discovery of plant remains the area of Montceau-les-Mines (Massif Cen- preserved within ironstone concretions. Accord- tral) in France has been known since the 19th ing to Roemer, the fl ora that was discovered in century (Grand’Eury 1877), and the fl ora and the Katowice nodules consisted of Sigillaria, fauna preserved within the ironstone concre- calamites and ferns, unfortunately however, tions of the area had been known since the none of these were either described by him or early 1980’s (Rolfe et al. 1982, Heyler & Pop- illustrated. lin 1988). The perfectly preserved land and Feistmantel (1874) was also familiar with freshwater fauna that were found within the fl oras preserved within the sideritic nod- these concretions instigated the greatest of ules of the Upper Silesia, as he determined interests (e.g. Racheboeuf et al. 2002, Vannier numerous preserved taxa within concretions et al. 2003, Racheboeuf et al. 2008, Racheboeuf found in Mysłowice, Katowice, Załęże, Orzesze, et al. 2009), although the fl ora itself has been Pszczyna and Zagórze in Sosnowiec. The speci- described in detail only comparatively recently mens described by Feistmantel were deposited (Charbonnier 2004, Charbonnier et al. 2008). in the Mineralogical Museum in Wrocław. The material worked upon was collected in Feistmantel however did not know the exact situ in three opencast coal-pits, where 6812 stratigraphic position of these specimens, but concretions containing remains of plants were only their place of origin, and in spite of his 257 plans to properly describe and illustrate them, The so far assembled collection of concre- he only published short notes about them. tions from Sosnowiec counts about 1500 speci- These specimens, as described by both Roe- mens with plant remains, from which 500 of mer and Feismantel, are believed to be by now them were found to have taxonomically diver- probably lost. sifi ed reproductive organs (Pacyna 2002a, Some specimens of plants found preserved 2007c). within sideritic nodules from the Westphalian The size of the concretions which hold the strata (Lower/Middle Pennsylvanian) in the plant fossils range from two centimeters in Jaworzno mine were collected by B. Rydzewski diameter, to more than twenty centimetres in early 20th century but unfortunately they in diameter, while their colour scheme ranges were never published. from darkgray, to beige, to brick-red. Bocheński (1939) illustrated two cone frag- Most of the plants within the concretions ments of Sigillariostrobus as preserved within were found to be well preserved in their three- ironstone concretion, but he did not described dimensional form, either coalifi ed or mineral- them in his text, even though they were the ized. When remains are coalifi ed then a fi lm only three-dimensionally preserved specimens of carbon covers the walls of the empty space in his collection. that remains from the biological degradation Outside the area from the Upper Silesia to of the soft tissues of the three-dimensional the Upper Carboniferous strata of Poland only organs. Mineralized plant fossils on the other Zimmerman (1962) mentioned in his unpub- hand are fi lled with a calcite, pyrite or with lished work of several species of sphenopsid the kaolinite. Mixed mineralisation of fossils and seed fern leaves preserved within sider- are also frequent e.g. calcitically – kaolinitic, itic nodules. These concertions were found in whereby such preserved fossils could also have the neighbourhood of Okrzeszyn in the Lower some of their parts coalifi ed (Pacyna & Zdeb- Silesia, however, Zimmerman (1962) did not ska 2001, 2002). describe nor illustrate any of these speci- Concretions with remains of plants and mens. animals as found in Sosnowiec originate from the Mudstone Series, Załęże Beds. The par- Sosnowiec ticularly frequent occurrence of ironstone con- Concretions with plant remains preserved cretions in the Mudstone Series, unparalleled within the Sosnowiec site were collected from in the remaining Carboniferous beds of the a waste dump of an already closed coal mine Upper Silesia, has been a known fact for a long “Porąbka-Klimontów” in Sosnowiec (Zagórze time (Porzycki 1972). Within the fi rst half of district) between 1998–2006. The area of 20th century they have been even exploited research was located within the Upper Silesia as a source of iron ores (Porzycki 1972, Tryje- Coal Basin, in its north-east part, called the faczka 1982). The ironstone concretions often Dąbrowa region (Pacyna 2022a, b, 2003, 2004, appeared irregularly within strata and were Pacyna & Zdebska 2001, 2002, 2005, 2010). not connected with any specifi c stratigraphical Numerous concretions with preserved levels. They could be found in all types of sedi- plants within them that were found in the mentary rocks, frequently in coal seams, but Ignacy mine in Zagórze (later called the Ignacy most often they accompanied mudstones. They shaft in Mortimer mine, the Mortimer-Porąbka appeared commonly in the form of single nod- mine, and the Porąbka-Klimontów mine) were ules, most often regular in shape with a diam- already known to Feistmantel (Feistmantel eter ranging about from 3 to 20 cm, however 1874). Unfortunately Feistmantel only gave some specimens have been found reaching in a list of taxa, without any detailed descrip- sizes up to 1 meter in diameter (especially in tions or illustrations. Furthermore A. Kota- the Dąbrowa region where such nodules have sowa – the author responsible for the descrip- been described). The Mudstone Series, dated tion of the fl ora which were found within the at Westphalian A-B, is divided into the Załęże mentioned above coal mine (Kotasowa 1968), and the Orzesze Beds. It is a monotonous series also made her own discoveries of concretions in respect of its lithology, mudstones and clay- with preserved plants within them (A. Kota- stones dominate its thickness, with some local sowa per. comm.), but however never did she intercalatations of sandstones, while intra-for- describe them. mational conglomerates are found to be rare. 258

Within the Mudstone Series a large number numerous. Sphenopsid diversifi ed stems, of coal seams appear. Many well preserved whorls of leaves and cones were also found. coalifi ed plant remains were found within this Ferns and seed ferns and their reproductive series (Kotasowa 1968). organs were differentiated taxonomically, but The fl ora preserved within the concretions are less numerous than the other groups men- from Sosnowiec was mainly allochthonous in tioned earlier. The presence of cordaites was origin. Plant debris must have been transported marginal, represented mainly by cones and through fl uvial action into the sedimentary rare leave fragments (Pacyna 2002a, b). Par- basin, where concretions then formed. Some ticularly interesting and of exceptional value to plant fragments were also probably carried by this location, and not unlike that which could the wind e.g. some fragments of leaves. These be found in Coseley (Kidston 1914), was the changes of location must have been probably large presence of various reproductive struc- very short however, since many of the very tures and small plant forms that have seldom delicate structures (e.g. microsynangia of seed been described at all (Pacyna & Zdebska 2001, ferns, lycopsid cones) were found perfectly pre- Pacyna 2003). served and intact. Only calamites, which could With the last salt water ingressions into be found at the borders of channels and lake the Upper Silesia having been dated at Namu- banks and constituted a large part of recog- rian A, the described fossil assemblage having nized plant fossils, could grow there in situ. been represented by both land and freshwa- Large nodules with three-dimensional internal ter environments therefore, and Sosnowiec casts of calamite stems and the stem bases that could be perceived as a parallel counterpart were preserved within concretions, give evi- to the fl ora and fauna type from Braidwood in dence to this interpretation. The dimensions Mazon Creek. Its faunistic assemblage hith- of the plant remains from the other systematic erto described consisted of horseshoe crabs, groups were usually signifi cantly smaller. eurypterids, insects and bivalves (Filipiak The fl oristic assemblage from the ironstone & Krawczyński 1995, 1996, Krawczyński et al. concretions as found in the Sosnowiec was gen- 1997), with subsequent searches bringing forth erally similar in nature to the fl ora described the discovery of various animal groups (Pacyna in the mudstones from which the concretions et al. 2004, Stworzewicz et al. 2009, Prokop originated (Kotasowa 1968), the organs which et al. 2012). prevailed in these concretions however were The age of the fossil assemblage was deter- seldom found in mudstones. Most interest- mined using the same method as that used at ing is the particularly frequent occurrence of Mazon Creek. Unfortunately, earlier palyno- reproductive organs and small plants that were logical attempts at an age evaluation that were generally seldom found within these strata undertaken did not yield conclusive results (Pacyna 2002a, b, 2003, 2007a, b, c, Pacyna (Krawczyński et al. 1997). On the basis of mac- & Zdebska 2001, 2010). Fossils from the concre- roscopic plant remains the age of the concre- tions were also preserved three-dimensionally, tions could be evaluated explicitly towards the as opposed to the fl ora from the shales where late Langsettian (late Westphalian A), Lower as a result of the strong diagenetic mudstone Pennsylvanian, with subzone Laveineopteris compaction, fossils were fl attened. This three- loshii, which resulted in the fl oristics assem- dimentionality was particularly visible in the blage VIII within the Carboniferous fl oral case of lycopod cones that are preserved within succession of Upper Silesia, as in accordance the concretions (Pacyna 2007c). with the Kotasowa and Migier (1995) division. Palaeoecological reconstructions place lyco- Within the fl ora from Sosnowiec concretions pods as the dominant component of the land- were found following index species for the scape of the fl ora. Most numerous within the Upper Silesia Carboniferous strata: Lepido- concretions were their leafl ess twigs which dendron aculeatum, L. obovatum, Sigillaria represented a new taxon (Pacyna & Zdebska mamillaris, S. rugosa, Mesocalamites ramifer, 2005). Cones and isolated sporophylls from Stylocalamites cisti, S. suckowii, S. undulatus, the new species of Lepidocarpon genus could Sphenophyllum cuneifolium, Annularia radi- be very frequently found, possibly belonging ata, Asterophyllites charaeformis, A. equiseti- to the same plant as the leafl ess twigs. Stems formis?, Sphenopteris (Renaultia) schwerini, and cones of other lycopsid species were less Pecopteris (Senftenbergia) pulmosa-dentata, 259

P. (Asterotheca) miltoni, Lyginopteris hoe- the following species were also determined as ninghausii, Neuropteris heterophylla, Neu- belonging to the Sosnowiec concretions: Sig- ralethopteris schlehanii, N. rectinervis, Pari- illaria mamillaris, S. rugosa, Sphenopteris pteris gigantea, P. pseudogigantea, Mariopteris (Renaultia) schwerini, Pecopteris (Senftenber- muricata, Karinopteris acuta, Margarito pteris gia) pulmosa-dentata, P. (Asterotheca) miltoni, pseudocoemansi, Alethopteris decurrens Neuropteris heterophylla, Paripteris gigantea, (Pacyna 2002a, b). These species were typical Paripteris pseudogigantea, Mariopteris muri- for the assemblage VIII in the Carboniferous cata, Margaritopteris pseudocoemansi (Pacyna fl oral succession of Upper Silesia, in accord- 2002a, b). ance with the Kotasowa and Migier (1995) Three of the recognized species in the Sos- division. nowiec nodules appeared in the Westphalian A In the case of the age estimation of the con- stage, each successively higher within their cretions from Sosnowiec the most essential ele- profi le. These were: Margaritopteris pseudo- ments of study were species extinct from the coemansi, Neuropteris heterophylla and Neu- end of Westphalian A stage or such species ralethopteris rectinervis (Kotasowa & Migier whose stratigraphical occurrence were limits 1983, 1995), these are index species for the exclusively to that of the Westphalian A stage. upper part of Westphalian A stage, subzone In examining the available material four such Laveineopteris loshii, enabling therefore to species were recognized: Lyginopteris hoening- pinpoint the specifi c age of the concretions at hausii, Neuralethopteris schlehanii, N. rectin- late Westphalian A (late Langsettian), subzone ervis, Karinopteris acuta and K. beneckei. From Laveineopteris loshii. These results also con- these, the most important was the species fi rm the numerous occurrences in the concre- Lyginopteris hoeninghausii, which was exclu- tions of the species Neuropteris heterophylla sively limited to the stratigraphical range of which species fi rst appeared in this subzone and Westphalian A. This became the main index the relatively few occurrences of Lygino pteris species for Westphalian A, as found not only hoeninghausii and the genus Neuralethopteris in Poland, but also for entire European Car- what is characteristic for this subzone (Cleal boniferous (Kotasowa 1975). The species Neu- & Thomas 1994). ralethopteris schlehanii and Karinopteris acuta Presumably as it was the case in Montceau- fi rst appeared in fossil records at the Namu- les-Mines, the lake in Sosnowiec was the point rian, and eventually achieved full quantitative of origine for the concretions as discussed culmination at the occurrence of Westphal- above (Doktor & Gradziński 1985, Vannier ian A towards the end of which they became et al. 2003). While allochtonous plant remains extinct (Kotasowa & Migier 1995). The species drifted down channels of crevasses or during Karinopteris beneckei fi rst appeared in Namu- greater fl oods, the calamites in turn, which rian B in which it was very frequently found, constituted a larger fraction among the vegeta- as well as in Westphalian A towards the end of ble remains, related to the borders of channels which they became eventually extinct. The spe- and lakes banks, they were the only species to cies Neuralethopteris schlehanii had the same grow as such in situ. Large nodules with three- stratigraphical range in Upper Silesia as it did dimensional internal casts of calamite stems, in Western Europe and was well index species as well as the very frequent occurrence of the (Goubet et al. 2000). N. rectinervis appeared whorls of Asterophyllites leaves, both not unlike exclusively in the upper part of Westphalian those found in Montceau-les-Mines, gave evi- A. Presence of those species indicating that dence to such an interpretation (Charbonnier the concretions originated from Westphalian 2004). The dimensions of the plant remains A stage, the Lyginopteris hoeninghausii zone from the other plant groups remained usually (the fossil plants assemblage VIII in the Car- signifi cantly smaller. As a result of such a pal- boniferous fl oral succession of Upper Silesia aeoenvironmental interpretation it is interest- according to the Kotasowa and Migier (1995) ing to note such a large presence, of otherwise division, the fl oristic assemblage IV according rare in fossil record, isolated megasporophylls to the local Kotasowa division (Kotasowa 1968) of genus Lepidocarpon. Lepidocarpon sporo- for the Dąbrowa region). In addition to the spe- phylls were able to travel to the lake, where cies that were typically part of this fl oral assem- concretions were formed during the greater blage, including the ones mentioned above, fl ood. Numerous occurrences of this type of 260 sporophylls were recognized in the Mazon were preserved within the sideritic nodules in Creek and Carterville nodules (Schopf 1938, Sosnowiec are common as fossils, as is the case 1941, Gastaldo 1977, Pfefferkorn 1979). with the Bickershaw and Westhoughton sites A very early cementation of soft sediment (Anderson et al. 1997, 1999). Interestingly con- by iron and carbonate compounds can preserve cretions often formed around these spherical the surface confi guration of organic parts, but reproductive organs. nonetheless most tissues, even those initially The number of recognized plant organ spe- preserved have been degraded and the voids cies found in the Mazon Creek fl ora have now which remain after could be secondarilly fi lled reached up to three hundred (Pfeffernkorn in. The structure of the epidermis is generally 1979). An attempt has been made to compare not well preserved, this phenomenon was well the diversity of the fl ora from the Mazon Creek observed in the Mazon Creek fl ora (Pfefferkorn and from the Sosnowiec, and this to this day 1979). However, with this particular mode of this operation is still in progress, as not all the fossil preservation the delicate structures can taxa have yet been determined in the Sosnow- preserve itself, e.g. such as it was the case iec fl ora. In Sosnowiec the amount the pteri- with the microsynangia of seed ferns which dophyte reproductive organs have been discov- were described from Mazon Creek and Coseley ered to be greater in number, particularly the (Kidston 1914, Drinnan & Crane 1994), and lycopsid cones, while the amount of leaves of also found in Sosnowiec and described as the seed ferns signifi cantly smaller. To this day new species Codonotheca silesiaca, Boulayath- only one fertile fern leaf have been discovered eca ciliata and Dolerotheca migierii (Pacyna in Sosnowiec, while in Mazon Creek the fertile 2007a, b, c, Pacyna & Zdebska 2010). Since fern leaves were diversifi ed and abundant (Pfef- such similar morphological structures were ferkorn et al. 1971, Pacyna & Zdebska 2002). preserved often in ironstone concretions such Pfeffernkorn (1979) postulated that on the as its been discovered in the different sites (in basis of fossils found in Mazon Creek the mode Mazon Creek, in Sosnowiec), and since these of preservation of plants in the concretions did types of plant structures were rarely, if ever, not have an infl uence on the specifi c composi- found within rocks accompanying coal seams, tion contained within them fl ora, while on the therefore it could be deduced that some types other hand the occurrence of rare forms of speci- of morphological structures were predisposed mens within the concretions intensifi ed the col- more than others to become the nucleus of the lections of specimens, hence within a suitably concretion. An example of such type of plant large collection one would have the chance of organ could be the thin lycopod twig, which fi nding increasingly more and more rare forms has an aggregation of leaves on its apex. Such of specimens. Unfortunately the composition types of fossils have been found in the Sosnow- of the fl ora from Sosnowiec does not confi rm iec concretions and could probably be referred such an interpretation. The collection of plants to a new species, very similar in nature, but from Sosnowiec nodules, though quite large in different taxonomically from those described number with about 1500 specimens, is dispa- in the Mazon Creek and Carterville fl oras (Noe rately smaller than the one from Mazon Creek. 1925, Gastaldo 1977, Pfefferkorn 1979, Pacyna In the Museum of Illinois can one fi nd above & Zdebska 2005). 16 thousands specimens of concretions con- The Mazon Creek fl ora was characterized by taining plants. Among the plants within Sos- a large number of species with large intraspe- nowiec nodules almost half constituted rare or cifi c size ranges. The occurrence of gigantism unknown forms of species as found within the in some of the forms is interesting, such as it shales (various reproductive organs, lycopsid was the case with some of the species found thin twigs). It is worth noticing that similar outside of the Mazon Creek nodules, which and delicate morphological structures were tend to be signifi cantly smaller in sizes (Pfef- often preserved within sphaerosiderites, but ferkorn 1979). In Sosnowiec on the contrary, rarely were they ever found in the shales. One the large presence of small forms was strik- can thus perhaps conclude that some types of ing, the size of the lycopsid and the sphenopsid organs are predisposed to become the nucleus cones being there signifi cantly smaller in com- of the concretion. An example of such types parison to such similar cones as those found of delicate organs are the seed fern microsy- in the coeval shales. The lycopsid cones, which nangia, which appeared in great numbers 261 in the Mazon Creek as well as in Sosnowiec and mixed plants presumably from different (Pacyna 2007a, b, c, Pacyna & Zdebska 2010), places of origin. For this reason the palaeon- another example could be the thin lycopsid vironmental reconstruction for this fl ora was twigs with leaves aggregated onto their apices, deemed to be very diffi cult (Baird et al. 1986, as discovered in Mazon Creek (Noe 1925), as Pacyna 2004). However the frequency of the well as recently the concretions in Sosnowiec occurrences of the plant remains from the dif- (Pacyna & Zdebska 2005). ferent taxa, the differences in sizes of the pre- Since such delicate organs decayed the quick- served organs, the degree of the disintegration est, one could suppose that their preservation of the seed fern complex leaves, can give us in the concretions was a result of a surplus in some clues as to the kind of environments from feedback, whereby the delicate organ quickly which these plants came from. One can there- decomposed, and changed the local conditions fore try to relate some of the found taxa to the of the environment so as to favour the neces- otherwise described taxa as found in the litera- sary formation of a concretion. As result of this ture of the palaeoenvironment coal forests. process the concretion thus arises, the more Generally known reconstructions of the such plant remains the more easily concretions Carboniferous coal forests have little in com- will therefore form (Baird 1997). Of course mon with the truth. They present coal-form- organisms of this type, which were frequently ing plants growing side by side on a peatbog, preserved in the shales (e.g. leaves, fragments which we know today is false, since they did of twigs), also appeared frequently within the not appear growing together at all. The envi- concretions themselves, with the proportions ronments of Carboniferous lowlands were also of the organisms being either often present or much more differentiated, which can be well sporadically present within the shales, the lat- observed on the basis of the sedimentologi- ter of which was of much greater important to cal research (e.g. Doktor & Gradziński 1985). the concretions. Althought some have tried to reconstruct the The fossils assemblage from the Sosnow- Carboniferous “forests” from the very beginning iec concretions was identical in respect to the of palaeobotany development, these reliable taxonomic composition within the assemblage environmental reconstructions of the Carbon- described by Anderson et al. (1997) in Bick- iferous lowlands based on plant macroremains ershaw. Such taxonomical compositions were have had a relatively short history (DiMichele typical for the Europaean late Westphalian & Phillips 1985). Only the pioneering works A fl oras, and permited the evaluation of the by Gastaldo (1987, 1991, 1996) contributed to age of the fossils to be made not only from the modern reconstruction of such Carbonifer- Bickershaw but also from Sosnowiec, confi rm- ous lowlands, as well as the observations made ing that both concretions had indeed the same by Scheihing and Pfefferkorn (1984) during correct age. Plants preserved within the iron- their expeditions to the Orinoco delta, which stone concretions from Bickershaw were disco- permitted a better understanding of the tapho- everd as being mainly coalifi ed, only the cones nomic aspects of the Carboniferous lowlands of Lepidostrobus were sometimes pirytized, as environment. they could also be fi lled with calcite and kao- Palaeogeographical reconstructions place linite. Identical modes of preservation were the Upper Silesia in the Carboniferous period found in Sosnowiec. Similarly, the concretions only several degrees north of the equator in from Bickershaw were found on the spoil tip, the area of the tropical climate. The environ- as were the ones from Sosnowiec. ment for the deposition of the Mudstone Series Within the shales that were found in the was an extensive, fl at river plain, subjected to spoil tip in Sosnowiec from which the concre- constant subsidences, formed by meandering tions were likely derived, stems of Calamites, and anastomosing rivers. Fine-grained sedi- Stigmaria fi coides root systems and fragments ments were deposited mostly on the forested of the bark from trunks of Sigillaria were areas outside river channels and only in the found. These taxa were also recognized in the small part in shallow, relatively short-termed shales from which came the concretions in lakes. Sands were deposited mainly in the Westhoughton (Anderson et al. 1999). river channels and in the areas of crevasse The fl ora from Sosnowiec was alloch- splays. Peatbogs were developed on badly thonous in origin, meaning it contained drifted drained swamp areas of the external fl ood 262 plane, with signifi cantly limited deliveries of well as herbaceous ferns (e.g. Renaultia schw- clastic materials being made to the peatbogs erini). Seed ferns recognized in Sosnowiec were (Doktor & Gradziński 1985). mainly representatives of the Medullosales Plants that were preserved within the con- group. Their leaves were found to be: Neu- cretions represent different environments of ropteris obliqua, Neuralethopteris schlehani, the river-plain as occupied through the mean- N. rectinervis, Paripteris gigantea, P. pseudo- dering rivers. Most often in the concretions gigantea, and Alethopteris decurrens, as well, thin leafl ess lycopsid twigs were found which it so appears, their reproductive organs, e.g. belong probably to a new pseudoherbaceous microsynangia Codonotheca silesiaca, Bou- rizomorphic lycopsid taxon (Bateman & DiMi- layatheca ciliata, Dolerotheca migieri. Less chele 1991, Bateman 1992, Bateman et al. numerous were the seed ferns from the fam- 1992, DiMichele & Bateman 1996, Pacyna ily of Mariopteridaceae (Karinopteris acuta, & Zdebska 2005). Presumably this was a plant K. beneckei, Mariopteris muricata). There rep- which formed compact thicket, as it had a very resentatives of Lyginopteridales could also be short, lignifi ed trunk growing out of its repeat- found: the leaves Lyginopteris hoeninghausi edly divided stigmaria, while from its trunk and pollen organs Silesiatheca formosa. The branches grew out accross the surface of the Seed ferns were climbing plants, their slender ground. This lycopsid grew probably within the stems holding on to the trunks of Sigillaria and area of a fl ood plane. Flood plains were most arboresent ferns (Gradziński & Doktor 1995). often covered and grew by congeneric commu- They could also form thick thickets in which nities of lycopsids resistant to stagnant water their thin trunks covered the tangle of roots (Phillips 1979, DiMichele & Phillips 1994). and produced 4-metre huge leaves supported Probably most of the cones and isolated sporo- by each other. In the undergrowth there surely phylls that were found within the concretions grew thickly herbaceous horsetails from the from Sosnowiec belonged to this new species genus Sphenophyllum. of lycopsid, mature megaspore cones disinte- As they quickly grew during the fl oods cre- grated onto single sporophylls, which were vasse splays delivered sediments to the fl ood adapted to the distribution of water (Bateman plain area. Under these circumstances pioneer et al. 1992). At least during part of the veg- species were calamites from the group Styloca- etative season these plants were suberged in lamites e.g. S. cistii, S. suckowii, and S. undu- water, be it during a fl ood or a rainy season. latus (Gradziński & Doktor 1995). Calamites The most characteristic feature of this plant created also “bulrush” communities over banks was its quick sheding of leaves, some leaves of lakes. Within the concretions taxa were remainded only on the very ends of twigs, and found which could have created these commu- displayed not unlike that of brush. Sheded lyc- nities: Diplocalamites paleaceus, Mesocalam- opsid leaves (Cyperites bicarinatus) were fre- ites cistiformis, M. ramifer, Annularia radiata, quent in concretions. A. sphenophylloides, Asterophyllites charae- Within the area which is know today as formis – particularly often found together Sosnowiec, in sediments formed in river bars, with the cones Calamostachys charaeformis, a taxonomically distinct fl ora once grew, com- Asterophyllites equisetiformis? (Pacyna 2002a, posed of several species of Lepidodendrales: b, Pacyna & Zdebska 2001, 2002). Lepidodendron aculeatum, L. dichotomum, and Cordaitales were still rare element of coal- L. obovatum, from which concretions contain- forming forests in Westphalian A (DiMichele ing imprints of bark scales, twigs and leaves. & Phillips 1994), however by then they already The environment of river bars – taxonomi- had occupied different ecological niches from cally rich among environments of Carbonif- the thicket of the mangroves type, to the erous lowlands (DiMichele & Phillips 1994) mountainous forests (Falcon-Lang & Scott was represented in the concretions by frag- 2000). Their participation in the concretions ments of the bark of lycopsids and the leaf- from Sosnowiec was small. The specimens lets of complex leaves of ferns and seed ferns. were represented by only a few leaves from Lycopsids from the genus Sigillaria (S. mamil- the genus Cordaites and differentiated them- laris, S. rugosa) lived there, among them grew selves, but only ever so slightly in comparison arborescent ferns (Sphenopteris sp., Pecopteris the described in literature cones, by a complete plumosa-dentata, P. miltoni, P. avoldensis) as lack of seeds. In the landscape of Sosnowiec 263 they could appear far from the fl ood plain (Flo- Sites with fossils preserved within the rin 1938–1945, Meyen 1987, 1997, Gastaldo ironstone concretions, particularly the Mazon 1991, 1996, Šimůnek 2000, Šimůnek & Lib- Creek site, are recognized universally as ertín 2006). examples of fossil assemblages carrying a lot of information about extinct organisms, the sites characterizing themselves by their huge CONCLUSIONS wealth and completeness. They are examples of Konservat–Lagerstätten (Nitecki 1979, The preservation of an organism within Baird et al. 1885a, b, Baird et al. 1986, an ironstone concretion was due to its quick Baird 1990, Shabica & Hay 1997, Shields formation, and enabled the preservation of 1998, Schellenberg 2002, Nudds & Selden the original three-dimensional shapes of the 2008). However, none of these, as of yet organisms as well the preservation of its deli- described Carboniferous palaeontological cate morphological elements. sites in Poland have gained the recognition The fl oras preserved within the ironstone of being named a Lagerstätte. The locality concretions united in a very interesting way: in Sosnowiec is signifi cant as the fi rst exam- its allochtonic origin with unusual taphonomi- ple of a Carboniferous Lagerstätte in Poland cal aspects resulting from syngenetic with (Pacyna 2008b). The exceptionally well pre- the surrounding sediment formation of iron- served three-dimensional fossils are impor- stone concretions (Baird et al. 1986, Schopf tant specimens which elucidate new informa- 1975, 1979). tion on the morphology of plant reproductive Most of the fl oras preserved within ironstone organs. The Sosnowiec site is a signifi cant concretions from Europe and North America Konservat – Lagerstätte from a region of originated from the Westphalian stage – the middle Europe that is at present relatively period with the greatest diversity of tropi- understudied (Pacyna & Zdebska 2002, Zatoń cal Carboniferous fl oras. Similarities in their & Marynowski 2004). taxonomic composition are likely the result of New research techniques like high-resolu- a taphonomical fi lter during the formation of tion X-ray microtomography which has been the concretions, which favoured the comport- used to extract detailed anatomical informa- ment of delicate and rare taxa, particularly the tion in plant fossils from deposits elsewhere reproductive organs. promise future discoveries from this locality The assemblage of plant remains from the (Spencer et al. 2013). Sosnowiec ironstone concretions was very dif- ferentiated and contained forms that are yet to ACKNOWLEDGMENTS be described. The assemblage was exceptional This paper benefi ted from the discussion with because among the remains of the plants, R. Gastaldo, W. DiMichele, H. Pfefferkorn, A. Kotas, reproductive organs that were seldom found A. Kotasowa, T. Migier, Z. Šimůnek, E. Purkyňová in other fossil plant sites dominated the area, during workshop “Namurian Floral Change Project” as well as many other small forms of well pre- (Sosnowiec August 4–8, 2003). We submit hearty served plants. Such small plant forms as found thanks to I. Liparski, S. Charbonnier and S. Florjan for discussions and crtical remarks. J. Hilton and B.J. in the Carboniferous shales and mudstones Slater, the journal reviewers are thanked for helpful were commonly poorly preserved and therefore comments which helped improved this paper, they also treated in the literature as detritus and very kindly corrected English. We especially thank jour- rarely described, eventhough they often car- nal’s Editor-in-Chief L. Stuchlik for help and encour- ried much important informations about the agement during manuscript preparation. This review is based on a part of the senior author PhD thesis. The morphology and the anatomy of small taxa. research was supported in part by State Committee for Within nodules from Sosnowiec an intriguing Scientifi c Research KBN grant No. 2 P04C 103 26. fauna of land and freshwater animals have been also found, such animals were described up to now from only several locations in the REFERENCES world (Filipiak & Krawczyński 1995, 1996, Krawczyński et al. 1997, Pacyna & Zdebska ABBOT M.L. 1958. The American species of Astero- 2001, Pacyna et al. 2004, Stworzewicz et al. phyllites, Annularia, and Sphenophyllum. Bull. 2009, Prokop et al. 2012). Am. Paleont., 38(174): 285–390. 264

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