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Annales Societatis Geologorum Poloniae (2021), vol. 91: 137–148 doi: https://doi.org/10.14241/asgp.2021.12

MATONIACEOUS PRESERVED IN GROWTH POSITION IN LOWER DUNE SANDSTONES OF THE HOLY CROSS MOUNTAINS (POLAND)

Grzegorz PACYNA

Jagiellonian University in Kraków, Faculty of Biology, Institute of Botany, Department of , Phytogeography and Palaeobotany, Gronostajowa 3, 30-387 Kraków, Poland; e-mail: [email protected]

Pacyna, G., 2021. Matoniaceous ferns preserved in growth position in Lower Jurassic dune sandstones of the Holy Cross Mountains (Poland). Annales Societatis Geologorum Poloniae, 91: 137–148.

Abstract: The occurrence of the braunii (Goeppert) Harris, preserved in the growth position in sandstones of dune origin, is documented from the late Pliensbachian Szydłowiec Sandstone of the Drzewica Formation in the Holy Cross Mountains. This formed monotypic stands on coastal dunes during a marine highstand. The presence of this common Early Jurassic fern in such an unfavourable environment attests to the huge ecological plasticity of this species.

Key words: Pliensbachian, foreshore, aeolian dune, ferns in life position, , Matonia.

Manuscript received 20 January 2021, accepted 10 May 2021

INTRODUCTION

Ferns were an important element of vegeta- Phlebopteris was proposed in 1829 by Brongniart tion, with numerous taxa and a diversified ecology (e.g., (Brongniart, 1828–1837) for several European Jurassic Van Konijnenburg-van Cittert, 2002; Barbacka et al., ferns. In 1830, Brown (in Wallich, 1830) proposed the 2014b). Families that today are of restricted occurrence as genus Matonia for extant fern specimens, collected from relicts (e.g., , , , the Malayan Peninsula. The family Matoniaceae was pro- , Osmundaceae, Schizaeaceae) were then abun- posed by Presl (1848). Schenk (1871), Zeiller (1885) and dant (Taylor et al., 2009). They lived in a variety of environ- Raciborski (1894) were among the first researchers to rec- ments, also colonising open areas and forming a Mesozoic ognise that Phlebopteris Brongniart (= Laccopteris Presl in analogue of today’s meadows. This is in contrast to modern Sternberg) belongs to the Matoniaceae and referred other fern families (e.g., Pteridaceae, Athyriaceae, Blechnaceae, fossil species to this family. The Matoniaceae originated in Aspleniaceae), which diversified in the in the the during the second fern radiation and had their shade of early arborescent angiosperms and today are main- widest range during the Mesozoic (Rothwell and Stockey, ly undergrowth (Schneider et al., 2004). As in some 2008; Pott et al., 2018). About twenty fossil genera have tropical areas today, Mesozoic ferns were early succession- been described on the basis of foliage, petrified and al-stage colonisers of disturbed and open areas (Mehltreter petioles, which were distributed on all continents (Schenk, et al., 2010; Barbacka, 2011). Ferns also constituted a signif- 1865–1867; Seward, 1899; Harris, 1931, 1961; Hirmer and icant part of the diet of dinosaurs, especially low browsers Hoerhammer,1936; Van Konijnenburg-van Cittert, 1993; (e.g., Gierliński and Pieńkowski, 1999; Brown et al., 2020). Tidwell and Ash, 1994; Popa, 1997; Givulescu and Popa, Among the typical Mesozoic fern families, Matoniacaeae 1998; Taylor et al., 2009; Barbacka et al., 2016). Today, this are especially frequent and well known (Harris, 1931, family is represented by two genera, Matonia (known since 1961; Van Konijnenburg-van Cittert, 1993; Popa, 1997; the Early Jurassic) and Phanerosorus Copeland, each rep- Klavins et al., 2004; Schmidt and Dörfelt, 2007; Kvaček resented by two species and confined to Indonesia, Borneo and Dašková, 2010; Naugolnykh and Pronin, 2015; Pott et and New Guinea (Seward, 1899; Kramer, 1990; Taylor et al., 2018; Badihagh and Uhl, 2019; Van Konijnenburg-van al., 2009). The family is characterised by the presence of Cittert et al., 2020). This family was first known from the pedate fronds with pinnate or more rarely bipinnate pinnae fossil record and secondarily from modern floras. The fossil and sporangia, arranged in sori, with a peltate indusium 138 g. pACYNA

(Kramer, 1990; Lindsay et al., 2003; Taylor et al., 2009; supraregional sea-level changes. Pieńkowski (2004) de- Van Konijnenburg-van Cittert et al., 2020). fined the deposits as the Kamienna Group and subdivided Matoniaceae are well represented in Polish Mesozoic flo- them into lithologic formations, showing regional facies ras, especially the Early Jurassic of Holy Cross Mountains, differentiation, in a study using high-resolution sequence with genera such as Matonia Brown in Wallich, Matonidium stratigraphic analysis, describing the lithofacies, identifying Schenk, Phlebopteris Brongniart (= Laccopteris Presl in the depositional systems and subsystems, and determin- Sternberg) and Stiehler, numerous in species and ing their fluctuation in space and time. The Holy Cross specimens (Raciborski, 1891, 1892, 1894; Makarewiczówna, Mountains are a marginal south-eastern part of the basin, 1928; Reymanówna, 1963, 1965; Barbacka et al., 2010, but the strata of seven formations and ten discerned se- 2014a, 2016; Pacyna, 2013). quences are present (Fig. 2) and accessible for research in This paper describes matoniaceous ferns, preserved in many outcrops and drill cores. the growth position in Lower Jurassic coastal (foreshore) The plant fossils described here originate from the late dune sandstones in the abandoned Podkowiński Quarry Pliensbachian sequence VI, which belongs to the Drzewica at Szydłowiec, in the Holy Cross Mountains (Fig. 1). Formation. The depositional sequence VI (Pieńkowski, Although in situ plants in the growth position are rather fre- 2004, 2006) lies on a prominent erosion surface; the sedi- quent in some strata, for example in the Coal ments overlying the erosional sequence boundary (parase- Measures (Falcon-Lang, 2015), plants preserved in aeolian quence VIa) are coarse alluvial deposits. The depositional dune strata are exceptional and are rarely described in the system is interpreted as a meandering river. The ensuing late palaeobotanical literature; more frequent are reports of fos- Pliensbachian transgression was associated with the devel- sil plants from interdune deposits (Gradziński and Uchman, opment of a foreshore-shoreface-barrier lagoon deposition- 1994; Parrish and Falcon-Lang, 2007; Taylor et al., 2009; al system. Rapid inundation of the palaeorelief with alluvial Florjan and Worobiec, 2016). Only very seldom are such valleys and coastal plains resulted in an embayed coastline preserved plant remains as taxonomically determinable as with detached beach-barrier ridges and a nearshore deposi- was possible for the material presented here. tional system with extensive shoreface deposits. The tran- sitional deposits between the transgressive systems tract GEOLOGICAL SETTING and the highstand systems tract (maximum flooding stage) are represented by very well sorted sandstone lithofacies, During the Early Jurassic, terrigenous, continental, mar- up to 20 m thick, with hummocky cross-stratification, hori- ginal-marine and marine deposits up to 1,400 m thick ac- zontal bedding and tabular cross-bedding. They are known cumulated in a large epeiric basin, extending across Poland as the Szydłowiec Sandstone, exploited for construc- as a part of the Central European Basin (Pieńkowski, tion use since medieval times (Karaszewski, 1960, 1962; 1991, 2004, 2006; Pieńkowski et al., 2008, 2020; Barth Karaszewski and Kopik, 1970). These quartz sandstones et al., 2018). Sedimentation was mainly controlled by occur in the Szydłowiec area, specifically in the lower parts

Fig. 1. Map showing the Podkowiński Quarry at Szydłowiec (red arrow), other localities discussed in the text, and the location of the study area in Poland MATONIACEOUS FERNS PRESERVED IN GROWTH POSITION 139 of the quarries at Szydłowiec (the Antecki and Podkowiński to Karaszewski (1962), Karaszewski and Kopik (1970) and quarries) and the exposures at Szydłówek and Śmiłów Pieńkowski (2004) the plants in the growth position were (Pieńkowski, 2004). Higher in the sequence, a regressive present at the bases of aeolian sandstone bodies, up to 2 m trend, associated with a highstand systems tract, is devel- thick, which wedge out laterally and are covered by argil- oped in a foreshore-dune-backdune/lagoon environment. laceous strata of coastal lagoon origin with drifted plant The Podkowiński Quarry is now abandoned and flooded. remains. Plant macroremains are present at many levels However, a detailed profile of this interval is known from of the Drzewica Formation, but in general are poorly pre- the nearby Śmiłów Quarry (Pieńkowski, 2004). According served and undeterminable. Karaszewski (1962) observed

Fig. 2. Lithologic profile of Lower Jurassic deposits of the Holy Cross Mountains (Profile courtesy of G. Pieńkowski, based on Pieńkowski, 2004, fig. 67, modified), with source strata of material investigated and other fossil floras. 140 g. pACYNA plant remains and rhizoids along the entire profile of this Selected synonyms: formation. Skurczyńska-Garwolińska and Woroncowa- 1841 Laccopteris brauni sp. nov. – Goeppert, Marcinowska (2015) illustrated several specimens, includ- pp. 7–8, pl. 5, figs 1–7. ing those collected by Karaszewski, Ploch and Gierliński, with plant remains from the Szydłowiec Sandstones at 1841 Laccopteris germinans sp. nov. – Goeppert, Szydłowiec, Śmiłów and Mirzec, near Starachowice, from pp. 9–10, pl. 6, figs 1–12. the collection of the Polish Geological Institute Museum; 1865–1867 Laccopteris göpperti sp. nov. – Schenk, none of these, probably with one exception, are taxonom- pp. 94–97, pl. 23, figs 1–12, pl. 24, figs 2–5. ically determinable. Plants buried in the growth position 1865–1867 Laccopteris münsteri sp. nov. – Schenk, pp. (although difficult to determine) were found in the same dune 97–98, pl. 24, figs 6–10, pl. 25, figs 1, 2a, b. interval in the Śmiłów Quarry (Pieńkowski, 2004; Gierliński 1891 Laccopteris elegans Presl in Sternberg et al., 2006). Specimens from Szydłowiec (Skurczyńska- – Raciborski, pp. 305–306, pl. 4, figs 1, 4. Garwolińska and Woroncowa-Marcinowska, 2015, p. 224, fig. 2) and from Mirzec (p. 224, fig. 1) probably contain 1899 Laccopteris elegans Presl in Sternberg plant shoots preserved in the growth position. The rest rep- – Seward, pp. 193–196, text-fig. 8. resent medium-sized detritus, preserved as moulds and casts 1928 Laccopteris elegans Presl in Sternberg of undeterminable shoots or twigs. The remains described – Makarewiczówna, p. 3. here are unique in being taxonomically recognisable. 1928 Laccopteris Münsteri Schenk 1867 Strata of the Drzewica Formation are well exposed to- – Makarewiczówna, p. 4, p. 1(10), figs 2, 2a. day in exposures at Szydłówek and Śmiłów (Pieńkowski, 1936 Phlebopteris brauni (Goeppert) Hirmer and 2004), and also are known from boreholes, such as Hoerhammer – Hirmer and Hoerhammer, the Gutwin (Rogalska, 1976) and Brody-Lubienia bore- pp. 7–17, text-figs 3, 4, 5, 1A–D, pl. 1, holes (Pieńkowski et al., 2016). Depositional sequence VI figs 1–12, pl. 2, figs 1–4, pl. 4, fig. 7. deposits belong to the late Sinemurian–Pliensbachian mega- Horstisporites planatus Zone (Marcinkiewicz, 1971; 1936 Phlebopteris muensteri (Schenk) Hirmer et Marcinkiewicz et al., 2014). Hoerhammer – Hirmer et Hoerhammer, pp. 17–26, text-figs B, 5, 2A–B, pl. 3, figs 1–7, pl. 4, figs 1–6, pl. 5, figs 1–6. MATERIAL AND METHODS 1980 Matonia braunii (Goeppert) Harris – Harris, The specimens were collected by the late Władysław pp. 295–310, figs 1–20. Karaszewski in the 1950s, when the Szydłowiec Sandstone 2016 Matonia braunii (Goeppert) Harris was exploited intensively in the Podkowiński Quarry. They – Barbacka et al., pp. 860–865, figs 2–5. are stored in the Institute of Botany of the Jagiellonian 2019 Matonia braunii (Goeppert) Harris University, collection number Palaeobotanical Collection – Barbacka et al., pp. 167–169, figs 6h, 7a, b. IB UJ 109. The plant fossils are preserved in orange-grey, 2020 Phlebopteris muensteri (Schenk) Hirmer medium-grained sandstone three-dimensionally, as voids et Hoerhammer – Van Konijnenburg-van after the decay of stipes and as the impressions of leaf Cittert et al., pp. 9–11, pl. 5, figs 1–6. blades. The suprageneric classification follows Cronquist et al. (1966), Smith et al. (2006) and PPG I (2016). The Material: PC IB UJ 109/1 (isolated stipes, one frond, isolat- plant remains were observed with a Technival 2 stereoscop- ed pinnules), 109/2 (groups of stipes, one pinna). ic microscope and photographed with a Nikon NIKKOR Description: Frond fragments preserved in growth position AF-s DX Micro NIKKOR 85 mm f/3.5G ED VR camera. (Figs 3–5). Frond palmately dissected with pinnatifid pin- nae. More than twenty stipes preserved singly or in groups SYSTEMATIC PALAEONTOLOGY of ~6 (Fig. 5A, B). Stipes 1.5–2.0 mm in diameter, up to 20 cm long, incomplete, not attached to rhizomes (rhizomes not preserved). Only one frond reasonably close to complete, POLYPODIOPSIDA Cronquist, with at least 12 pinnae still attached to the stipe (Fig. 3A). Takhtajan et Zimmermann, 1966 Isolated pinnae also present (Fig. 5C, D). Preserved pin- Schimper, 1869 nae incomplete, up to 15 cm long, 15–18 mm wide. Pinnae Family MATONIACEAE Presl, 1848 rachis ~1 mm wide. Pinnules linear, opposite to subopposite, densely arranged, attached at 70–80° angles, slightly sick- le-bent toward upper end of the pinnae. Pinnules 7–10 mm Genus Matonia Brown in Wallich, 1830 long, 2–4 mm wide, separated from each other by convex Type species: Matonia pectinata Brown in Wallich, 1830; ridges of sandstone 1–1.5 mm wide (Fig. 3A–D). Pinnules Mount Ophir near Malacca, Malay Peninsula; extant of approximately uniform size along pinnae, smaller in basal and apical parts of pinnae, up to 3 mm wide and up to Matonia braunii (Goeppert, 1841) Harris, 1980 5 mm long (Fig. 3C). Pinnule apices mostly poorly pre- Figs 3–5 served, rather rounded. Pinnule venation indistinct, only central vein sometimes visible (Fig. 3B, C). Probably all pinnae sterile, no traces of sori detectable. MATONIACEOUS FERNS PRESERVED IN GROWTH POSITION 141

Fig. 3. Matonia braunii from Podkowiński Quarry. PC IB UJ 109/1. A. Palmately dissected frond with pinnatifid pinnae three-dimensionally preserved on several rock surfaces. B. Upper part of specimen from 3A, seen from above, void after stipe decay and several pinnae visible. C. Enlargement of pinnae fragments (detail from 3B), pinnules separated from each other by convex ridges of sandstone. D. Lower part of specimen from 3A, pinnae with pinnules separated from each other by prominent convex ridges of sandstone.

Remarks: The material described here is preserved well the pinnae are still attached to the stipe. There are also nu- enough to be taxonomically determined. This is rather ex- merous isolated stipes (Fig. 4) and some isolated pinnae in ceptional for plants growing in such an unfavourable envi- both described specimens (Fig. 5C, D). The isolated stipes ronment (a dunefield) and found in a rock type that generally have the same shape and dimensions as stipes still connected does not preserve fossils well (medium-grained sandstone). to the pinnae, so most probably they also belong to this fern. Palmately dissected fronds with pinnatifid pinnae and No other plant remains are preserved in the two rock specimens. the characteristic shape of the pinnules allow this fossil to Morphologically similar were described and illus- be assigned to the Matoniaceae family. In one specimen trated by Goeppert (1841), Schenk (1865–1867), Seward (PC IB UJ 109/1, Fig. 3A) the leaf is almost complete and (1899), Hirmer and Hoerhammer (1936), Barbacka et al. 142 g. pACYNA

Fig. 4. Matonia braunii from Podkowiński Quarry. PC IB UJ 109/1. A. Specimen from 3A, lower side, flat surface, visible numerous voids after stipes decay. B. Side of specimen from 4A, visible moulds of stipes split lengthwise; note low-angle cross bedding of sandstone perpendicular to stipes. C. Opposite side of specimen from 4A, visible moulds of stipes split lengthwise.

(2016, 2019) and Van Konijnenburg-van Cittert et al. (2020) in Sternberg) Hirmer and Hoerhammer, different in many as belonging to Phlebopteris braunii (Goeppert) Hirmer and aspects from Matonia braunii (Raciborski, 1891, 1892; Hoerhammer, Phlebopteris muensteri (Schenk) Hirmer et Makarewiczówna, 1928; Pacyna, 2013). Members of oth- Hoerhammer, Matonia braunii (Goeppert) Harris and some er fern families present, such as the Osmundaceae and other clearly synonymous species (see list of selected syn- Dipteridaceae, are morphologically different enough not to onyms). Harris (1980) considered the specimens classified be confused with the Matoniaceae. as Phlebopteris muensteri as sun fronds and Phlebopteris Distribution: Matonia braunii, as understood here braunii as shade fronds of the same species. He also demon- (Phlebopteris braunii and P. muensteri as its synonyms), is strated the presence of indusiate sori and on this basis re- known from the Late Triassic to , main- ferred both species to the extant genus Matonia and merged ly from Europe, but most frequent and characteristic for them under the name Matonia braunii (Goeppert) Harris. the Early Jurassic (Goeppert, 1836; Schenk, 1865–1867; Worth noting is the usual co-occurrence of both of them Harris, 1980; Van Konijnenburg-van Cittert, 1993; Van at most localities (Schenk, 1865–1867). Barbacka et al. Konijnenburg-van Cittert et al., 2020). In the Early Jurassic (2016, 2019) followed Harris’ (1980) interpretation, when floras of Poland, this species is a characteristic but not describing specimens from the Lower Jurassic of Poland very numerous element (Barbacka et al., 2014; Pacyna, and Hungary. However, some authors still question these 2021). Specimens similar to those described here were de- species synonymy (see e.g., Van Konijnenburg-van Cittert, scribed and illustrated from the Lower Jurassic of the Holy 1993; Schweitzer et al., 2009; Van Konijnenburg-van Cittert Cross Mountains area (Chlewiska, Chmielów, Dźwiertnia, et al., 2020), based on subtle differences in leaf and sori Gromadzice, Jędrzejowice, Szewna) by Raciborski morphology in forms from the two extremes of variability. (1891) and Makarewiczówna (1928), and referred main- Interestingly, the leaf morphology (especially the relative- ly to Laccopteris elegans Presl in Sternberg (summarized ly short sterile pinnules) of the presently described plants, in Pacyna, 2013). The species Laccopteris elegans was which most probably grew on dunes and were exposed to proposed by Presl in Sternberg (1838), on the basis of direct sunlight, rather resembles that of P. braunii, interpret- poorly preserved, rather indeterminate isolated pinnules ed by Harris as shade leaves. (Jung and Knobloch, 1972; Kvaček and Straková, 1997). The only other species from the Matoniaceae present The species name Laccopteris elegans is a nomenclatu- in the Lower Jurassic strata of the Holy Cross Mountains ral synonym of Asterocarpus lanceolatus Goeppert, 1836, is the easily identifiable Phlebopteris angustiloba (Presl and is an illegitimate name, as superfluous, including the MATONIACEOUS FERNS PRESERVED IN GROWTH POSITION 143

Fig. 5. Matonia braunii from Podkowiński Quarry. PC IB UJ 109/2 A. Two groups of stipes bent in one direction. B. Upper side of specimen from 5A, two groups of stipes bent in one direction. C. Opposite side of specimen from 5A, one pinna im- pression above several moulds of stipes. D. Detail from 5C, one pinnula visible; note low-angle cross bedding of sandstone. type of the formerly validly published name (Goeppert, material described here and were assigned by him as at least 1836; Turland et al., 2018). The species affiliation of the in part to the ferns, but without a detailed description and Polish material, historically referred to this species, needs taxonomical determination. One specimen (Karaszewski, revision. Material probably referable to Matonia braun- 1962, pl. 8, fig. 1), refigured by Skurczyńska-Garwolińska ii was described by Barbacka et al. (2014a) from strata of and Woroncowa-Marcinowska (2015, p. 223, fig. 2, invert- Toarcian–Aalenian age from the Ciechocinek IG 1 borehole ed 180° in relation to Karaszewski’s figure), may represent as Phlebopteris muensteri. New well-preserved material a frond of the same fern species as that identified here, but of Matonia braunii from the Niekłań borehole in the Holy the photos are of poor quality and at very low magnification. Cross Mountains was described by Barbacka et al. (2016). The bend of the groups of stipes in one direction in the material described here (Fig. 5A) may indicate wind blow- DISCUSSION ing sand from one direction. Low-angle cross bedding, per- pendicular to the vertically preserved stipes, indicates that Preservation of plant remains in growth position the plants were still alive and in the growth position during in dune sandstones the deposition of the sand. Two preserved groups of stipes (Fig. 5A, B) are not attached to rhizomes. The lack of rhi- Plant remains are known from several horizons in the zome preservation is interesting and needs explanation, as Lower Jurassic successions of the Holy Cross Mountains the plants are preserved in the growth position. Karaszewski (Fig. 2); they are especially numerous in and (1962) did not mention the presence of rhizomes in his rarer in Sinemurian, Pliensbachian and Toarcian strata published material; no traces of them are present in the (for the most recent summary, see Pacyna, 2013). Only in specimens described here. In extant Matonia pectinata the the Drzewica Formation, they are frequently preserved in rhizomes are horizontal and produce fronds at certain dis- growth position, albeit for the most part poorly preserved tances (Seward, 1899; Schweitzer, 1978). Two groups of in sandstones. long stipes from specimen PC IB UJ 109/2 may have orig- Karaszewski (1962) was the first to discover, describe and inated from one . In specimen PC IB UJ 109/1, the illustrate plants in the growth position from the Szydłowiec stipes are more scattered and one has pinnae still attached Sandstone in the Podkowiński Quarry. His specimens (Fig. 3). This sandstone block most probably preserves (Karaszewski, 1962, pl. 8, figs. 1–3) are very similar to the the upper parts of the stipes (Fig. 4). Karaszewski (1962) 144 g. pACYNA suggested that the dune area was quickly inundated after Active dunes are a very difficult habitat for plants plant burial, which allowed the plants to be fossilised and (Podbielkowski and Podbielkowska, 1992; Warren, 2013; preserved. One possible explanation of the lack of rhizomes Sudnik-Wójcikowska and Krzyk, 2015). They are endan- is that they would have been present in the soil layer be- gered by sand-blasting or burial in sand, and they have neath the frond-preserving sandstone and that they decayed problems establishing on a loose substrate. Sand dunes are after the plant was buried by sand and died. The presumed common features of shorelines. Beach dunes consist of the thin soil layer with rhizomes was not preserved. Fronds of foredune, the angled side of which faces the sea, the sand the fern Anomopteris sp. from the Early Triassic Eifel region plain at the top of the dune, and the backdune. Plants that in Germany were similarly preserved in the growth position thrive on the foredune are exposed to salt spray, strong wind, in sandstone of fluvial, not aeolian origin. Their rhizomes and burial by blowing and accumulating sand. Plants that were rooted in siltstone lying below the sandstone and poor- thrive on the broad dune plain and backdune grow together ly preserved, unlike the well-preserved fronds (Fuchs et al., into dense patches that hold the dune together. The presence 1991). In monotypic colonies of Clathropteris meniscioides of horizontal rhizomes of Matonia braunii could stabilise Brongniart (Dipteridaceae) preserved in situ in the Early a dune, like rhizomes of the modern grass Leymus arenarius Jurassic of central Patagonia, horizontal rhizomes are also (Linnaeus) Hochstetter on the Baltic coast (Podbielkowski lacking (Choo et al., 2016). and Podbielkowska, 1992; Sudnik-Wójcikowska and Krzyk, 2015). As no other fossil plants were found asso- ciated with Matonia braunii fronds, it probably formed Palaeoecology of Matonia braunii at Szydłowiec with monotypic stands on coastal dunes, similar to the monotyp- reference to the environmental preferences of ferns ic strands formed by Matonia today on sandy soils (Piggott Today, the Matoniaceae family is confined to tropical and Piggott, 1988). Thailand, Indo-Malaysia and Polynesia; the present distri- Generally ecological variability of ferns is narrower to- bution is a relict area. Matonia pectinata Brown in Wallich day than in the Mesozoic as a result of angiosperm compe- grows in exposed areas in relatively open scrub, on or tition (Rothwell and Stockey, 2008; Pšenička et al., 2020). near the tops of isolated mountains at 750–2,000 m a.s.l., However, extant ferns tolerate a wide range of environmen- but has also been found at sea level. It is a thicket-forming tal conditions (Mehltreter et al., 2010), and some species fern adapted to poor sandy soils (Piggott and Piggott, 1988; can colonize such highly disturbed habitats as lava flows, Kramer, 1990; Lindsay et al., 2003). dunes, landslides as well as areas of forests that have been The ecology of fossil Matoniaceae was more diversified damaged by burning. Living Matoniaceae are relictual and the family was more widespread (Van Konijnenburg- with a restricted number of species, ecology and range and van Cittert, 2002; Wang, 2002; Barbacka et al., 2014b) but certainly do not reflect ancient aptitudes of this fern fam- probably connected mainly with warm and humid climates. ily. Extant Matonia although adapted to poor sandy soils Forms preferring humid conditions (especially having large does not grow on coastal dunes (Piggott and Piggott, 1988; fronds) and those more stress-adapted were certainly pres- Kramer, 1990; Lindsay et al., 2003). At least one fossil ge- ent and growing in the understorey along riverbanks and on nus of Matoniaceae i.e. Weichselia, was salt tolerant and open plains, forming assemblages similar to today’s mead- grew in tidal environments (Silantieva and Krassilov, 2006). ows. Only some species were adapted to an arid climate and Some extant ferns demonstrate a wide range of ecological rather dry localities (e.g., Weichselia or Piazopteris Lorch) tolerance, including possibility of growth on dunes. Bracken and are frequently preserved as fusain (Tidwell and Ash, (Pteridium Scopoli) and scrambling ferns, Gleichenia 1994). There were also species with quite variable ecolo- Smith, Bernhardi and Presl, are gy, the most spectacular being Phlebopteris woodwardii often abundant on and characteristic of nutrient-poor sites Leckenby. This fern was present in such different environ- such as heaths, landslides, recent fire scars and coastal sand ments as coal-forming peatbogs, riverbanks, and fire-prone dunes (Mehltreter et al., 2010). Pteridium is one of the inland heaths (Raciborski, 1894; Harris, 1961). Matonia world’s most widespread invasive . Its suc- braunii also probably had a wide range of environmental cess is explained by its broad survival strategies, it grows in preferences. In the Lower Jurassic coal-bearing strata of woodlands and their edges, clearings, dunes, hedgerows and Hungary, this fern was interpreted as preferring the wet- roadside verges (Mehltreter et al., 2010). Southern bracken test, disturbed, alluvial habitats (Barbacka, 2011; Barbacka (Pteridium esculentum (Forster) Cockayne) grows on coast- et al., 2019). In the Polish age-coeval localities, it is present al sand dunes on the Kaitorete spit, southern New Zealand, in river and deltaic or lagoonal settings at Gromadzice and and in subtropical eastern Australia, another species colo- Chmielów. The environmental conditions of the specimens nises sand dunes and coastal cliffs in UK (Walker et al., from the Niekłań borehole are interpreted as flood-plain- 1981; Mehltreter et al., 2010). Fossil Matonia braunii might backswamp, located at the transition between floodplain and have had wide ecological amplitude and colonised such dis- lacustrine-back-swamp, associated with a fluvial environment turbed habitats as floodplains, varied alluvial and deltaic (Barbacka et al., 2016). The fern was found in a claystone settings and sand dunes. Numerous polypodiaceous ferns, bed separating two thin coal seams. The material described normally associated with woodlands, were found grow- here at Szydłowiec considerably broadens the environmental ing on lake dunes in the Indiana Dunes National Park, in tolerance of this species to include coastal dunes (very poor- Indiana, USA (Tryon, 1936). Among angiosperms are spe- ly developed sandy soils) and probably to include increased cies which grow not only on coastal dunes, but also in other salinity, which is typical for such environments. habitats, e.g., sand sedge (Carex arenaria Linnaeus) grow MATONIACEOUS FERNS PRESERVED IN GROWTH POSITION 145 mainly on North Sea and Baltic Sea coasts, but also in the and Brody-Lubienia boreholes; the outcrops have not valleys of larger rivers in forests with sandy soils (Sudnik- been analysed palynologically so far. These data provide Wójcikowska and Krzyk, 2015). a much wider picture of the vegetation. The following plant groups were confirmed by sporomorphs: bryophytes, lyco- phytes, ferns from the families Cyatheaceae, Dipteridaceae, Matonia braunii in the growth position at Podkowiński Dicksoniaceae, Gleicheniaceae, Matoniaceae, Osmundaceae Quarry in the context of Drzewica Formation biota and Schizeaceae, seed ferns, conifers from the families Additional data about plant cover during deposition of Taxodiaceae, Podocarpaceae and Pinaceae, cycads, bennet- Drzewica Formation relevant for discussed here fossils pro- titaleans and ginkgophytes (Rogalska, 1976; Pieńkowski vide nearby exposures and boreholes. The Śmiłów Quarry et al., 2016). Pollen grains from seed ferns and conifers pre- exploits fine-grained, very well sorted, quartz-cemented dominate over other sporomorphs. of Matoniaceae sandstones of the uppermost VIst sequence, which repre- (cf. Matonisporites sp.) are rather rare. Marine influences sent mostly nearshore facies (Pieńkowski, 2004, 2006). were confirmed by the presence of acritarchs (Rogalska, Aeolian reworking is present in the uppermost part of the 1976). profile, and undeterminable plants buried in growth posi- tion by barrier crest aeolian dunes were found (Pieńkowski, CONCLUSIONS 2004; Gierliński et al., 2006). They are not similar to the remains at the Podkowiński Quarry; they are larger, coali- The ecological range of the common Early Jurassic fern fied, up to 0.5 m high, and preserved only in the upright po- Matonia braunii (Goeppert) Harris was expanded to coastal sition. The remains may be interpreted as probably the basal dunes on the basis of specimens, preserved in the growth parts of plants, although no roots or rooting structures were position, from the late Pliensbachian Szydłowiec Sandstone described or illustrated, with numerous basal branches that of the Drzewica Formation in the Holy Cross Mountains. were interpreted by the authors as narrow bladed leaves. The fern formed monotypic stands on coastal dunes during This may be a fern with an upright rhizome producing nu- a marine highstand. From a taphonomical point of view, the merous fronds, from which only stipes are preserved, the fossil material is interesting as it is preserved well enough blades having fallen off before fossilisation. Skurczyńska- to be determined taxonomically. This is rather exceptional Garwolińska and Woroncowa-Marcinowska (2015, p. 251, for plants growing in such an unfavourable environment as fig. 2) erroneously identified as ?Neocalamites hoernensis a dunefield and preserved in such a type of sediment (me- (sic) some indeterminable drifted plant remains from this dium-grained sandstone). The leaf morphology (especially quarry. Casts of large logs of drifted trees and dinosaur foot- the relatively short, sterile pinnules) of the plants described, prints were also found. Dinosaur footprints in the Śmiłów which grew on dunes and were exposed to direct sun- Quarry were formed in the siliciclastic foreshore zone, in- light, resembles that of the leaves of the ecological type of terfingering with backshore dunes; it seems that the foot- Matonia braunii previously interpreted as shade growing. prints were formed on the coastal dunefield. On the basis of Fern thickets, formed by Matonia braunii on coastal dunes, this locality, Niedźwiedzki et al. (2009) recognised a new may have provided food for herbivorous dinosaurs crossing type of dinosaur track assemblage in the Early Jurassic of the dunefields. the Holy Cross Mountains (compare with Gierliński and Pieńkowski, 1999). This association is characterised by the Acknowledgments predominance of small- and medium-sized forms of thero- pods and early thyreophorans, but also contains footprints The paper is dedicated to the memory of W. Karaszewski and made by large sauropods (Gierliński and Niedźwiedzki, M. Reymanówna, who initiated modern research of the Early 2005). Fern thickets, formed by Matonia braunii on dunes, Jurassic flora of the Holy Cross Mountains and who first studied may have provided food for herbivorous dinosaurs crossing the material described here. Thanks go to D. Zdebska (Institute the coastal dune fields. of Botany, Jagiellonian University, Cracow) for bringing the spec- Szydłówek is a small quarry in the vicinity of Szydłowiec imens to my attention and for fruitful discussion, M. Barbacka exploiting the Szydłowiec Sandstone and showing a succes- and J. Ziaja (W. Szafer Institute of Botany, Polish Academy of sion comprised of nearshore-foreshore-backshore/eolian fa- Sciences, Cracow) for discussions and sharing important litera- cies. Fossils from this outcrop include numerous vertebrate ture, J. Ziaja for kindly revising the published palynological data tracks, impressions of woody trunks, plant root moulds, from the Gutwin borehole, and G. Pieńkowski (Polish Geological brackish bivalves, and limuloids (Pieńkowski, 2004, 2006). Institute – National Research Institute, Warsaw) for fruitful dis- Among the vertebrate tracks are well-preserved belonging cussions about the geological context and age of Early Jurassic to theropods, sauropods and ornithischians (Niedźwiedzki floras of the Holy Cross Mountains. I thank journal referees et al., 2008; Niedźwiedzki and Remin, 2008; Gierliński et G. Pieńkowski for correcting especially the geological part of the al., 2009). Impressions of drifted woody trunks, probably paper and kindly providing profile of the Holy Cross Mountains, of conifers, abundant in this outcrop, indicate the occur- Evelyn Kustatscher (Museum of Nature South Tyrol, Bolzano, rence of vast coniferous forests around the track site, but Department of Earth and Environmental Sciences, Paleontology have not been subjected to detailed palaeobotanical analysis & Geobiology, Ludwig-Maximilians-Universität München, (Pieńkowski, 2004, 2006). München and SNSB-Bayerische Staatssammlung für Paläontologie Palynological data about the plant cover during sedimen- und Geologie, München) and journal editor P. Filipiak (Faculty tation of the Drzewica Formation come from the Gutwin of Earth Sciences, University of Silesia, Sosnowiec) for helpful 146 g. pACYNA comments and suggestions. M. Szewczyk is thanked for pho- Falcon-Lang, H. J., 2015. A calamitalean forest preserved in tographs of the specimens, A. Sojka for preparing the figures. growth position in the Pennsylvanian coal measures of M. Jacobs and F. Simpson (School of the Environment, University South Wales: implications for palaeoecology, ontogeny and of Windsor, Windsor) skilfully corrected the language. The study taphonomy. Review of Palaeobotany and Palynology, 214: was financed by funds from the National Science Centre, Poland 51–67. (Grant No. 2017/25/B/ST10/01273) and the Institute of Botany of Florjan, S. & Worobiec, G., 2016. Skamieniałości roślinne. 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