New data about Matonia braunii (Göppert) Har ris from the Early Ju ras sic of Po land and its ecol ogy
Maria BARBACKA1, 2, *, Grzegorz PACYNA3, Grzegorz PIEÑKOWSKI4 and Jadwiga ZIAJA1
1 Pol ish Acad emy of Sci ences, W. Szafer In sti tute of Bot any, Lubicz 46, 31-512 Kraków, Po land
2 Hun gar ian Nat u ral His tory Mu seum, De part ment of Bot any, 1476 Bu da pest, P.O. Box 222, Hun gary
3 Jagiellonian Uni ver sity, In sti tute of Bot any, De part ment of Palaeobotany and Palaeoherbarium, Lubicz 46, 31-512 Kraków, Po land
4 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Rakowiecka 4, 00-975 Warszawa, Po land
Barbacka, M., Pacyna, G., Pieñkowski, G., Ziaja, J., 2016. New data about Matonia braunii (Göppert) Har ris from the Early Ju ras sic of Po land and its ecol ogy. Geo log i cal Quar terly, 60 (4): 857–868, doi: 10.7306/gq.1322
Fern re mains of matoniacean af fin ity were found in the Lower Hettangian strata of lac us trine/backswamp or i gin from the Niek³añ PGI 1 bore hole (cen tral Po land, Holy Cross Mts.). The pre served frag ments have been iden ti fied as Matonia braunii (Göppert, 1841) Har ris, 1980. The re mains sug gest a rather small, low-growth plant with palmately com pound fronds. The sori con tain at least 5 sporangia pre served with well-de vel oped annuli. The spores are tri an gu lar, trilete and kyrtomate, with a thin and smooth sur face cor re spond ing with dis persed Dictyophyllidites mortoni (de Jer sey, 1959) Playford et Dettmann, 1965. Based on the gross mor phol ogy of ster ile and fer tile pinnae, sug ges tions made by Har ris (1980) on the syn on ymy of Phlebopteris braunii (Göppert, 1841) Hirmer et Hörhammer, 1936 with P. muensteri Schenk, 1867 (Hirmer and Hörhammer, 1936) and their re fer ral to Matonia braunii is proved and con firmed in this pa per. The fern oc curs in strata in di cat ing a warm and hu mid cli mate and ap proach ing trans gres sion re sult ing in a high wa ter ta ble and the en hanced ac cu mu la tion of or ganic mat ter.
Key words: Po land, Holy Cross Mts., Hettangian, Matonia.
INTRODUCTION
Dur ing the Me so zoic, di ver si fied ferns con sti tuted a prom i - nent el e ment in the land veg e ta tion of the time (Van Konijnen - burg-van Cittert, 2002; Wang, 2002). Fam i lies such as Mato - niaceae, Dipteridaceae, Schizaeaceae, Gleicheniaceae, Hy - me no phyllaceae, Osmundaceae, Marattiaceae and Cya the a - ceae flour ished in Tri as sic and Ju ras sic times (Cleal, 1993). To - day, these fern fam i lies are mainly re stricted to trop i cal re gions and are rep re sented by a few taxa only (Seward 1899; Berry, 1919; Camus, 1990; Iwasuki, 1990; Kramer, 1990; Skog, 2001;
Roth well and Stockey, 2008; Tay lor et al., 2009). To day’s dom i - nant fern fam i lies like Polypodiaceae and Aspleniaceae (broa - dly con ceived) orig i nated dur ing the Cre ta ceous (Schnei der et al., 2004; Smith et al., 2006; Christenhusz and Chase, 2014). In the Early Ju ras sic, ferns were wide spread in many Eu ro pean lo cal i ties (Barbacka et al., 2014b) es pe cially in flu vial-deltaic coal-gen er at ing flo ras of the kind en coun tered in Swe den, Hun -
gary and Ro ma nia (Lundblad, 1950; Tralau, 1965; Popa, 1997;
Barbacka, 2011; Barbacka et al., 2014b, 2015).
Matoniaceae had their wid est dis tri bu tion dur ing the Me so - zoic, with re cords rang ing from Green land to Antarctica (Schenk, 1867; Seward, 1900; Berry, 1919; Har ris, 1931, 1961, 1980; Ar nold, 1956; Ash, 1969, 1972, 1991, 2001; Rushforth, 1970; Appert, 1973; Schweitzer, 1978; Ash et al., 1982; Czier, 1994; Skog and Litwin, 1995; Givulescu and Popa, 1998; Wang and Mei, 1999; Klavins et al., 2004; Mirzaie Ataabadi et al., 2005; Bartiromo et al., 2006; Nagalingum and Cantrill, 2006;
Schmidt and Dörfelt, 2007; Schweitzer et al., 2009; Kvaèek and Dašková, 2010; Naugolnykh and Pronin, 2015). About ten fos sil gen era of this fam ily have been de scribed from the Me so zoic based on their fo liage, and a sim i lar num ber based on pet ri fied rhi zomes and pet i oles (Hirmer and Hörhammer, 1936; Skog, 1988; Tidwell and Skog, 1992; Van Konijnenburg-van Cittert, 1993; Tidwell and Ash, 1994). Matoniacean fronds are pal - mately com pound or pen du lous, with pin nate, or more rarely bi - pin nate, pinnae. Sporangia are al ways ar ranged in sori (in a ring around the re cep ta cle), of ten with peltate indusium aris ing from the sorus cen tre. This is re garded as one of the di ag nos tic fea - tures of the fam ily (Brown in Wallich, 1830; Presl, 1848;
Kramer, 1990).
Matoniaceae are rep re sented in Pol ish Lower and Mid dle Ju ras sic flo ras by the gen era Phlebopteris and Matonidium (Raciborski, 1891, 1892, 1894; Makarewiczówna, 1928; Rey -
* Corresponding author, e-mail: maria.barbacka@gmail.com Received: September 12, 2016; accepted: October 26, 2016; first published online: November 10, 2016
ma nówna, 1963; Barbacka et al., 2010, 2014a). Ac cord ing to the lat est re view of Pol ish Lower Ju ras sic flo ras, which was un - der taken by Pacyna (2013), Phlebopteris angustiloba (Presl in Stern berg, 1838) Hirmer and Hörhammer 1936 is known from four lo cal i ties in the Holy Cross Mts. (Odrow¹¿, Huta, Groma - dzice and Chmielów), P. elegans (Presl in Stern berg, 1838) Go than and Weyland, 1954 from the Holy Cross Mts. (Groma - dzice, Szewna, Chlewiska, DŸwiertnia, Jêdrzejów and Chmie - lów) and also from Up per Silesia (Siewierz and Zawier cie), and P. muensteri Schenk, 1867 (Hirmer and Hörhammer, 1936) from the Holy Cross Mts. (DŸwiertnia and Chmielów) as well as Ciechocinek in the Kuyavian-Pom er a nian re gion (Barbacka et al., 2014a). Two spe cies, P. elegans and P. muensteri were re - ported from DŸwiertnia, which is lo cated near Niek³añ. Of these, ac cord ing to Pacyna (2013) P. elegans should be re ferred to P.
braunii (Göppert, 1841) Hirmer et Hörhammer, 1936.
In the Mid dle Ju ras sic flora of Grojec (Kraków re gion) the fol low ing spe cies have been de scribed: Phlebopteris angu - stiloba, P. muensteri (given by Raciborski, 1894) as Laccopteris mirovensis and re vised by Reymanówna (1963), Microdiction woodwardii Leckenby, 1864 (Raciborski, 1894) and Lacco - pteris phillipsii Zigno, 1856 (Raciborski, 1894).
GEOLOGICAL SETTING
The Niek³añ PGI 1 fully-cored bore hole (200 m) lo cated in cen tral Po land (the north ern slope of the Holy Cross Mts.) was drilled in 2010 and yielded a valu able, well-pre served core span ning the Up per Tri as sic – low er most Ju ras sic sec tion (Fig.
1), in clud ing a con tin u ous pro file through the Lower Hettangian sec tion with clay min eral data re flect ing im por tant cli ma tic changes (Pieñkowski et al., 2014; Brañski, 2014).
In Early Hettangian times, the Pol ish Ba sin (along with its depocentre, which is known as the Mid-Pol ish Trough) was lo - cated at a north ern sub trop i cal lat i tude (about 40–45°N). With a length of more than 700 km, the Mid-Pol ish Trough, which gen er - ally runs along the Teisseyre-Tornquist Zone (TTZ) and the Trans-Eu ro pean Su ture Zone (TESZ), is the larg est of many in - verted bas ins in west ern and cen tral Eu rope (Ziegler, 1990).
Rapid sub si dence in the trough com menced in the Early Hettangian (Pieñkowski, 2004). Up per Tri as sic strata are rep re - sented by red and var ie gated mudstones (Norian–Lower Rhaetian – Zb¹szynek and Wielichowo or var ie gated Parszów beds), cov ered with kaolinite-rich grey Parszów beds, re flect ing a pe riod of more hu mid con di tions and in tense chem i cal weath er - ing (Pieñkowski et al., 2014). The Tri as sic-Ju ras sic bound ary in the Niek³añ bore hole is lo cated at a depth of 162.2 m, at the ero - sional se quence bound ary cor re spond ing to the en hanced hy - dro log i cal cy cle and re ju ve nated palaeorelief, re sult ing in re - gional ero sion (Pieñkowski, 2004). The over lay ing Lower Hettan - gian al lu vial and lac us trine de pos its, to gether with the grey Parszów beds, are as signed to the Zagaje For ma tion (Pieñ - kowski, 2004). The Zagaje For ma tion starts the first depo sitional se quence (Hettangian in age) of the epicontinental Lower Ju ras - sic of Po land, di vided into 11 parasequences (Pieñ kowski, 2004), of which the lower 4 are rep re sented in the Niek³añ PIG 1 bore hole (Fig. 1). From a depth of 158.9 m up wards, char ac ter is - tic Ju ras sic palynomorphs (Pinuspolle nites–Trachy sporites as - sem blage) ap pear, in clud ing FAD (first ap pear ance da tum) of the in dex pol len grain Cerebropollenites thiergartii at a depth of 158 m (M. Hodbod, pers. comm., 2016). The strata be tween ca.
160 and ca. 135 m of the Niek³añ PIG 1 bore hole ap prox i mately cor re spond to the well-known So³tyków (also called Odrow¹¿) out crop (Pieñkowski et al., 2014) ex pos ing the low er most
Hettangian al lu vial plain de pos its (Pieñ kowski, 2004), which con - tain rich floristic re mains (Reyma nówna, 1992; Ziaja, 2006;
Barbacka et al., 2007, 2010). The ear li est Hettangian age of the out crop is in di cated by se quence strati graphic cor re la tion (Pieñ - kowski, 2004), macroflora dom i nated by the co ni fer Hirme riella muensteri (Schenk, 1867) Jung, 1968 and Podo zamites spp.
(Reymanówna, 1992; Wcis³o -Luraniec, 1992; Bar backa et al., 2010), the palyno morphs Classopolis torosus (Reissinger) Couper and Aratri spo rites minimus Schulz (Ziaja, 1992, 2006), as well as the conchostracans Bulbilimnadia kilianorum Kozur, Weems et Lucas, 2010. The tetrapod ichno fauna of the Zagaje For ma tion (Hettangian) ex posed at the So³tyków clay-pit shows high ichno taxonomic di ver sity (Pieñ kowski and Gierliñski, 1987;
Gier liñski and Pieñkowski, 1999; Gierliñski et al., 2001, 2004;
NiedŸwiedzki, 2011).
The Hettangian strata pro file in Niek³añ, like other pro files of the Zagaje For ma tion in the Holy Cross Mts., is of bi par tite char - ac ter – the al lu vial part (parasequence Ia, Fig. 1) con tain ing nu - mer ous sand stone lay ers de pos ited in flu vial chan nels is over - lain by parasequence Ib, dom i nated by much finer sed i ments.
The bound ary be tween parasequences Ia and Ib is cor re lated with the Early Hettangian step-wise trans gres sion ob served in NW and cen tral Po land (Pieñkowski, 2004; Pieñkowski et al., 2012). A rapid base-level rise had a pro nounced ef fect on sed i - men ta tion in the con ti nen tal ba sin lo cated fur ther to the south - -east – al lu vial de po si tion was rap idly re placed by fine-grained de po si tion and con se quently, the nonmarine cor re la tive sur face (time equiv a lent) of the transgressive sur face can be ob served within con ti nen tal de pos its. The ma rine trans gres sion in the re - gion com menced later. This is doc u mented by the trans - gressive sur face at the bot tom of parasequence Ic (Fig. 1 – bot - tom of the next Sk³oby For ma tion). The most fre quent litho - facies of the up per part of the Zagaje For ma tion (parasequence Ib) is rep re sented by dark, or ganic-rich, lam i nated mudstones and claystones. These mudstones and claystones are dark grey to black, some times ol ive-grey, lam i nated or of a mas sive ap pear ance, with nu mer ous plant roots. They are usu ally de - pos ited in shal low lakes or backswamps, so, as far as the lithofacies is con cerned, the floodplain depositional sys tem and lac us trine depositional sys tem are of ten very sim i lar or iden ti cal.
Plant growth and pedogenic pro cesses of ten oblit er ate pri mary struc tures. Lev els with abun dant small sid er ite con cre tions (“sid e r ite sphaerulites”) rep re sent a char ac ter is tic, early dia - genetic prod uct of pedogenic pro cesses in this per ma nently sat u rated soil. Plants rep re sent largely typ i cal wet land as so ci a - tion, also of a “reed swamp” char ac ter. The shal low wa ter ta bles oc cur ring in backswamps or ox bow lakes cre ated fa vour able con di tions for the de po si tion and pres er va tion of plant de bris typ i cal of the Gleysol type of palaeosol (Arndorff, 1993). The Gleysol type of palaeosol con tain ing abun dant plant roots and well-pre served plant frag ments trans formed into coal in ter ca la - tions is char ac ter is tic of this depositional sys tem.
The con tin u ously ris ing sea level led to a fur ther de crease in the de liv ery of clastic sed i ment and en hanced ac cu mu la tion of peat, which can be ob served in the strata just be low the trans - gressive sur face (Fig. 1, bot tom of parasequence Ic). Con tact be tween peat (or coal) and clastic sed i ments usu ally rep re sents a con sid er able hi a tus in clastic de po si tion (McCabe, 1984). For peat to form, the rate of in crease in ver ti cal ac com mo da tion space must equal the rate of ac cu mu la tion of the peat, and clastics must be ex cluded from the en vi ron ment (Bohacs and Suter, 1997). Peat ac cu mu la tion prob a bly oc curred in the wa - ter-logged re duc ing en vi ron ment of a per ma nently sat u rated palaeosol (Gleysol) where plant growth was caught up with sed i - men ta tion. The fre quent coal seams at the top of the Zagaje For - ma tion (and parasequence Ib) her ald an ap proach ing ma rine
858 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
Fig. 1. Pro file of the Niek³añ PGI 1 bore hole with po si tion of the fern Matonia braunii (star) and its palaeo geo graphi cal and palaeoenvironmental back ground
Gen er al ized pro file, li thol ogy: c – con glom er ate, s – sand stone, h – heteroliths, m – mudstone, c – claystone;
de tailed pro file to the right: s (c) – coarse to me dium-grained sand stone, s (f) – fine-grained sand stone, h – heteroliths, m – mudstone, cl – claystone
trans gres sion. The fern was found at a depth of 41.8 m, a metre be low a thin, dis tal cre vasse splay of a mini-delta char ac ter (the type 2 “progradational” cre vasse – Aslan and Autin, 1999) in a claystone bed sep a rat ing two thin coal seams (Fig. 1).
MATERIAL AND METHODS
The plant re mains de scribed in the pres ent pa per orig i nate from a borehole 10 cm in di am e ter. The length of the frond frag - ments is of ten re stricted by the di am e ter of the borehole. The fern is rep re sented by 7 spec i mens pre served in mudstone as im pres sions/com pres sions with well-pre served, coalified mat - ter. The larg est parts of the fronds are fer tile, while ster ile pinnae are more frag mented. Be sides these, some basal frag - ments of both types of pinnae are found.
Spores were ob tained in situ by mac er at ing coalified frag - ments of pinnules with sporangia in Schulze so lu tion (KClO3 + HNO3). These were then washed in 3% KOH and cen tri fuged af ter each stage of prep a ra tion.
SEM ob ser va tions were pro vided us ing a Hitachi 2360N scan ning elec tron mi cro scope (Hitachi Co., To kyo, Ja pan) for a gold coated sam ple im aged at 15 kV ac cel er at ing volt age.
The spec i mens are stored at the Geo log i cal Mu seum of the Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute in War - saw, and la belled with num bers 80.VI.150 A–D.
Tax o nom i cal clas si fi ca tion is given ac cord ing to Smith et al.
(2006).
RESULTS
SYSTEMATIC DESCRIPTION OF THE MACROREMAINS
Filicopsida
Or der: Gleicheniales Link, 1833 Fam ily: Matoniaceae Presl, 1847 Matonia R. Brown in Wallich, 1830 Matonia braunii (Göppert, 1841) Har ris, 1980 D e s c r i p t i o n. – Frond is palmately com pound with pin nate pinnae of rather slen der struc ture. The fer tile fronds were found sep a rately from the ster ile fronds, but both are of the same type and were in close prox im ity in the core (Fig. 2).
The pet i ole of the larg est pre served frond reaches 3 mm in width. The width of the pre served pet i ole in the other case is 2 mm, wid en ing at the pal mate branch ing to 5 mm. Two main basal arms arise (both rachises are 2–3 mm wide) from the pet i - ole with some pinnae in be tween [3 pinnae in spec i men no 80.VI.
151C (b)]. They have 1–1.2 mm wide rachises [Fig. 2B–D, G (a)].
The basal pinnules, about 2 x 2 mm large, be come more elon gated to wards the dis tal part of the pinna [Fig. 2B, Ea, F (a)]. The apex of the basiscopic pinnules is rather rounded [Fig.
2F (a)], be com ing sub acute to wards the dis tal end of the pinna (Fig. 3A–D).
The sur face of the pet i oles and rachises is very finely lon gi - tu di nally stri ated. Pinnules are op po site or subopposite. The lamina of neigh bour ing pinnules joins at the base, form ing at the rachis a 0.2 mm wide U-shaped edge.
The larg est ster ile pinna frag ment is 45 mm long. No ster ile pinnule is pre served in its en tirety. The lon gest ster ile pinnule frag ment is 10 mm long and 3 mm wide [Fig. 2G (b), H (b)].
Ve na tion is mainly only ev i dent as a dis tinct midrib run ning to the apex. Thin lat eral veins are hardly vis i ble in two ster ile
pinnules for a very short dis tance. They fork once just at the midrib (the basiscopic vein is al most at a right an gle to the midrib, acroscopic vein at 45° to the midrib) and at half of the pinnule width, they fork again (Fig. 4).
Fer tile pinnae are pre served in larger frag ments. The larg - est and most com plete pinna frag ment slightly nar rows to wards the apex and base. Rachis width ranges from 2 mm at the base of the frag ment to 1 mm at its top. Pinnules al most per pen dic u - lar to rachis, nar row. Their length is 11–25 mm, and their width, at 1–2 mm, is nar rower than that of the ster ile pinna frag ments.
The dis tance be tween the pinnule bases is 2–3 mm. The apex of the pinnules is sub acute (Fig. 3A–D).
In the fer tile pinnules, the veins are in vis i ble, with the ex cep - tion of the midvein run ning from the pinnule base to its apex.
The sori are ar ranged al ter nately in two rows on both sides of the midvein, half way be tween the midvein and mar gin (Fig. 3A, E–H). They start 2–4 mm from the base of the pinnule, end ing at the apex. They are cir cu lar, from 317 to 840 mm, av er ag ing about 550 mm in di am e ter, and in the cen tre, a trace of the stalk is vis i ble (about 130 mm in di am e ter); the dis tance be tween neigh bour ing sori is about 0.3 mm. The ex act num ber of sporangia per sorus is un known; the high est pre served num ber of annuli within one sorus is 3–5 (Fig. 5A, B). A num ber of sori are cov ered by indusia (Fig. 3H), but nu mer ous indusia are de - tached and re main on the coun ter part of the spec i men. They are up to 1 mm in di am e ter and their mar gins are widely si nu soi - dally in cised (prob a bly a re sult of com pres sion, Fig. 3E–G).
Some times, sin gle annuli are ad hered to these de tached indusia. In the cen tre from the in ner side, a track of the stalk is also vis i ble like on the sori (Fig. 3G). The annuli are well-de vel - oped. In ma jor cases, only the annuli of sin gle sporangia are ob - serv able in the sori along their whole length, while the rest of the sporangia have fallen off. They are 437–460 mm long in av er - age, the lon gest 851 mm. The width of the an nu lus ranges from 100.5 to 383 mm, av er ag ing about 212 mm. The lon gest pre - served an nu lus is formed by 12 cells with an ti cli nal walls form - ing ribs and periclinal walls col lapsed be tween them. The dis - tance be tween ribs (= width of the cells mea sured be tween the me dial long axes of the an ti cli nal walls) ranges from 14.9 to 72.9 mm, av er ag ing 41 mm (Fig. 5C–H).
DESCRIPTION OF SPORES IN SITU
In the mac er ated sporangia a to tal of 48 sep a rate spores and two clus ters con sist ing of about 20 and 40 spores (Fig. 5I) were found. Some times they were ad her ing to the sporangium cu ti cle shreds. Most of them are badly pre served and de formed with hardly re cog nis able de tails.
The spores are trilete and tri an gu lar in equa to rial out line with rounded api ces and con cave or straight sides in po lar view. They pos sess a trilete mark with el e vated and labrate commissures.
The kyrtome (interradial thick en ings, 3–5 mm across) is sub - parallel to the leasural ra dii and con nected on the api ces of the spores. In the equa to rial view, they are plano-con vex with a lon gi - tu di nal fold. The exine is thin and smooth (Fig. 5J, K).
The equa to rial di am e ter is 36–46 mm (15 best pre served spec i mens mea sured). The po lar di am e ter is 36–39 mm (4 spec i mens mea sured). The di men sions of clus ters mea sured in the wid est parts are: 172 x 162 mm and 215 x 215 mm.
The num ber of spores per sporangium is im pos si ble to de - ter mine pre cisely be cause of rel a tively small amount of spores found in the mac er ated sporangia. We es ti mate that one clus ter of spores con tains about 20 or 40 spores and prob a bly fills one sporangium.
860 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
DISCUSSION
Most of the fea tures de scribed above en able us to iden tify the fern from Niek³añ as Matonia braunii. How ever, we are un - able to be pre cise about the num ber of sporangia per sorus, their shape and the num ber of spores per sporangium due to
the poor state of pres er va tion of ma te rial. The fer tile fronds were prob a bly fully de vel oped and the sporangia were ripe, which con firms the po si tion of annuli en tirely spread and vis i ble along their full length. Usu ally only a sin gle an nu lus is pre served in place of the sorus. To be pre cise, of the ap prox i mately 520 sori pre served on spec i men Muz. Geol. PIG 80.VI.150A, about 140 have a sin gle an nu lus, and only 6 show more than one an - Fig. 2. Matonia braunii (Göppert) Har ris
A – spec i men Muz. Geol. PIG 80.VI.151C, frag ments of ster ile and fer tile pinne; B – spec i men 80 IV.151D, pet i ole with basal parts of pinnae; C – spec i men 80.IV.151B, pet i ole branch ing into fer tile pinnae; D – spec i men 80.IV.151C, pet i ole branch ing; E – spec i - men 80.VI.151B, shape vari abil ity of pinnules: fer tile pinna near base (a), in the mid dle part (b, d), ster ile pinna (e); F – de tail from Fig ure 2E, basal part with short pinnules (a), and fer tile pinnules (b); G – spec i men 80.VI.151C, pet i ole (a) as so ci ated with ster ile pinnae (b); H – de tail from Fig ure 2G, ster ile pinna (b)
nu lus (3–5). In Matonia braunii, the usual num ber of sporangia per sori is 10–12 and these are sit u ated ra di ally around the stalk of the indusium, as is usual in the case of Matoniaceae. The sporangia of the fern from Niek³añ do not form a ra dial struc - ture, but are strongly dis lo cated due to their state of pres er va -
tion. Vis i ble annuli are also in an un usual po si tion, not oblique, but flat tened by com pres sion.
We in ter pret the struc tures de tached and vis i ble on the spec i men coun ter part as si nu soi dally in cised coalified indu - sium. Their shape and cir cum fer ence is a lit tle larger than the 862 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
Fig. 3. Matonia braunii (Göppert) Har ris
A – spec i men 80.VI.150B, pos i tive im print; B – de tail from Fig ure 3A, rows of sori on pinnules; C – spec i men 80.VI.150A, coun ter - part of Fig ure 3A; D – de tail from Fig ure 3C; E – de tail from Fig ure 3A, indusia with stalks (in the cen tres); F – de tail from Fig ure 3A, indusia; G – mag ni fied de tail from Fig ure 3A, indusia; H – de tail from Fig ure 3C, sori cov ered by indusia
cir cum fer ence of the sori, and closely cor re spond to the indusia usu ally found in Matonia. The pinnule frag ments which do not show these struc tures on the coun ter part, have sori cov ered by a bulge of coalified mat ter (indusia). The de tached ones are de - formed by com pres sion and in cised by split ting. The pres ence of indusia ex cludes the Phlebopteris ge nus, sug gest ing Konijnenburgia Kvacek et Daskova, 2010 or Matonia. How - ever, Konijnenburgia dif fers from the fern from Niek³añ with re - gard to its ve na tion and wedge-shaped sporangia.
Due to the pre served fea tures of the fer tile and ster ile pinnules, the fern from Niek³añ could be in ter preted as Matonia braunii ac cord ing to Har ris’ dis cus sion (Har ris, 1980) of this spe cies from York shire. He first dis cussed and con firmed the uni fi ca tion of ster ile P. braunii fronds in ter preted by him as a shade form, with the ster ile and fer tile fronds of P. muensteri, in - ter preted as a sun form, into one plant, P. braunii. He no ticed the con stant co ex is tence of these two spe cies at many lo cal i - ties. He went on to as cribe P. braunii to the ge nus Matonia based on mor pho log i cal cor re spon dence, where only the pres - ence of indusia (Matonia) was in con sis tent. The whole Phle - bopteris ge nus is con sid ered to be exannulate, while the re cent Matonia ge nus pos sesses indusia. Orig i nally, Phlebopteris muensteri was de scribed as exanullate (Schenk, 1867), but Har ris (1980) rec og nized indusia on some spec i mens of this spe cies and con sid ered the rest of the spec i mens which lacked them to have lost their indusia. He no ticed that in re cent fer tile Matonia ma te rial, the indusia can de tach very eas ily, and come to closely re sem ble the ap par ently exanullate fronds of Phlebopteris muensteri.
Van Konijenburg-van Cittert did not fully agree with Har ris (1980) on this sub ject. In her opin ion, with out prov ing the pres - ence of indusia in spec i mens re ferred to Phlebopteris muensteri from clas sic lo cal i ties in Ger many and Rus sia, it was pre ma ture to fully ac cept P. muensteri with Matonia braunii as syn on y mous (apart from the ma te rial ex am ined and de scribed by Har ris, 1980). How ever, fer tile spec i mens from Niek³añ, with mor phol ogy typ i cal of P. muensteri, but pre served frag ments with sori and coun ter parts with de tached indusia, fully con firm Har ris’ state ment. Van Konijenburg-van Cittert (1993) also stressed the dif fer ences in pinnule shape and ve na tion in both spe cies. How ever the close prox im ity in the ex am ined core ma - te rial, of spec i mens hav ing pinnule mor phol ogy typ i cal of both P. muensteri and P. braunii strongly sup ports Har ris’ as sump - tion. Ac cord ing to Van Konijnenburg-van Cittert’s re search on matoniacean in situ spores, there are dif fer ences in spore di -
am e ter be tween P. muensteri and M. braunii. There fore, she treated P. muensteri and M. braunii as sep a rate spe cies. Based on the new ma te rial from Po land, we can con cur with Har ris (1980) that they are conspecific.
Un for tu nately, the small num ber of spec i mens known from the Pol ish Lower and Mid dle Ju ras sic as P. braunii from the Holy Cross Mts. and Up per Silesia, and P. muensteri from the Holy Cross Mts., Ciechocinek and the Mid dle Ju ras sic flora of Grojec (for ref er ences see In tro duc tion) are so badly pre served that it is im pos si ble to re vise them, and some of them (Up per Silesia) are prob a bly lost. We de cided to re tain these spec i - mens with their orig i nal iden ti fi ca tion.
The mor phol ogy of spores found ad hered to the thin pieces of cu ti cle in the mac er ated sporangia rep re sent typ i cal trilete, tri an gu lar forms with a smooth exine and are mainly known from the ferns of Cyatheaceae, Dipteridaceae, Dicksoniaceae, Matoniaceae and Gleicheniaceae (Balme, 1995). Such kind of spores hav ing thick en ings or folds more or less par al lel ing the leasural ra dii, are most sim i lar to fos sil (of the Late Tri as sic to Early Cre ta ceous) and ex tant matoniaceous spores de scribed by Van Konijnenburg-van Cittert (1993) and Van Konijnenburg - -van Cittert and Kurmann (1994), but they also re sem ble fos sil Dipteridaceae spores of Dictyophyllum rugosum (Couper, 1958; Balme, 1995). Van Konijnenburg-van Cittert (1993) and Van Konijnenburg-van Cittert and Kurmann (1994) de scribed and il lus trated in situ spores from Matonia braunii based on ma - te rial from York shire (Mid dle Ju ras sic) and Green land (Early Ju ras sic) as well as from re-ex am ined slides men tioned by Har - ris (1931, 1980). Van Konijnenburg-van Cittert (1993) de scri - bed Matonia braunii spores as valvate, with api ces usu ally slightly thick ened. This thick en ing is not vis i ble in the spores from Niek³añ, prob a bly due to the ma tu rity of these spores. The spores from Niek³añ are ma ture and ac cord ing to Van Konijnenburg -van Cittert and Kurmann (1994), the api cal thick - en ings in matoniaceous spores are much more pro nounced in im ma ture spores. The in situ spores from Matonia braunii de - scribed here are also slightly smaller (36–46 mm against 43–55 mm) than those de scribed by Van Konijnenburg-van Cittert (1993) and Van Konijnenburg-van Cittert and Kurmann (1994), but mor pho log i cally very sim i lar.
In Po land in situ fos sils spores of Matoniaceae have only been de scribed to date by Reymanówna (1963) from Phlebo - pteris angustiloba spec i mens found in Grojec near Kraków (south ern Po land).
Tri an gu lar, smooth, trilete fos sil spores in dis persed state can be at trib uted to sev eral gen era: Matonisporites Couper, 1958; Concavisporites Pflug in Thomson and Pflug, 1953;
Dictyophyllidites Couper, 1958; Deltoidospora Miner, 1935;
Cyathidites Couper, 1953; Phlebopterisporites Juhász, 1979 and Phanerosorisporites Juhász, 1979 (Miner, 1935; Couper, 1953, 1958; Thomson and Pflug, 1953; Juhász, 1979; Litwin, 1985; Van Konijnenburg-van Cittert, 1993; Balme, 1995). Dis - persed spores de scribed as Deltoidospora and Cyathidites are tri an gu lar and trilete with a smooth exine, but only some times have del i cate thick en ings near the trilete mark and do not have el e vated and labrate commissures. Dis persed Matoni sporites, Phlebopterisporites and Phanerosorisporites usu ally have a thick exine. Dis persed Concavisporites and Dictyo phyllidites, es pe cially Dictyophyllidites mortoni (de Jer sey, 1959) Playford et Dettmann, 1965, pos sess a trilete mark with el e vated and labrate commissures, thick en ings subparallel to the trilete mark and a rel a tively thin exine, so they are most sim i lar to the in situ spores from the Niek³añ fern.
Tri an gu lar, trilete dis persed spores with a smooth exine do not have clear sys tem at ics be cause of var i ous opin ions on the syn on ymy of these spore spe cies. Lund (1977) placed Leio - Fig. 4. Sche matic draw ing of pinnule ve na tion
[from spec i men 80.VI.151B (e)]
triletes Naumova, 1939, Cyathidites, Concavisporites and Dictyo phyllidites in Deltoidospora and treated Dictyophyllidites mortoni (de Jer sey, 1959) Playford et Dettmann, 1965 (= Leio - triletes mortoni de Jer sey, 1959) as a syn onym of Deltoido - spora toralis (Leschik, 1955) Lund, 1977 (Leschik, 1955; de Jer sey, 1959; Playford and Dettmann, 1965). Jansonius and Hills (1976) and Litwin (1985) treated these gen era as dis tinct.
Dis persed spores “sim i lar” to the spores of Matoniaceae, e.g. Dictyophyllidites mortoni (de Jer sey, 1959) Playford et Dettmann, 1965 (= spores de scribed as Clathropteris obovata var. magna Turatanova-Ketova in pa pers of Rogalska, 1954, 1956), Matonisporites equiexinus Couper, 1958, Matoni spo - rites phle bo pteroides Couper, 1958, Matonisporites sp. and spores cf. Matonia sp. or Matoniaceae in Rogalska, 1976 and
864 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
Fig. 5. Matonia braunii (Göppert) Har ris
A, B – spec i men Muz. Geol. PIG 80.VI.150A, sori, only annuli pre served af ter sporangia de com po si tion; C–F – spec i men 80.VI.150A, sori with only one an nu lus pre served; G – spec i men 80.IV.150A, an nu lus, SEM pic ture; H – de tail of an nu lus from Fig - ure 5G, vis i ble thick ened an ti cli nal walls form ing ribs, SEM pic ture; I – clus ter of about 20 spores; J, K – spores in situ in LM, prox i - mal view show ing trilete mark and kyrtome (interradial and api cal thick en ings)
Mamczar, 1986), have been de scribed from Rhaetian and Lower Ju ras sic sed i ments of west ern Po land (Or³owska - -Zwoliñska, 1983), Pomerania (Pieñkowski et al., 2012), and Lower Ju ras sic sed i ments of the Holy Cross Mts. re gion (Rogal ska, 1976; Ziaja, 2006), in clud ing the Mroczków - -Rozwady area (Rogalska, 1956). Be sides the Rhaetian-Lower Ju ras sic de pos its, these spores were also found in Mid dle Ju - ras sic sed i ments of the Holy Cross Mts. re gion (Rogalska, 1976), and Up per Ju ras sic and Lower Cre ta ceous sed i ments of the Kuyavian re gion (Mamczar, 1986).
PALAEOENVIRONMENT AND PALAEOECOLOGY OF MATONIA BRAUNII
The en vi ron men tal back ground of Matonia braunii from the Niek³añ bore hole can be con vinc ingly in ter preted as a flood - plain -backswamp en vi ron ment lo cated at the tran si tion be - tween floodplain (cre vasse splay-delta) and lac us trine -back - swamp zones. Spec i mens are pre served in a lam i nated clay - stone. The fern was found in the up per part of the Zagaje For - ma tion, be low the ma rine trans gres sion sur face. The trans - gres sion sur face marks the bot tom of the next parasequence and Sk³oby For ma tion (Fig. 1). The ap proach ing trans gres sion sig nif i cantly in flu enced sed i men tary pro cesses, rais ing the wa - ter ta ble and at the same time lim it ing de liv ery of clastic sed i - ments, which en abled the ac cu mu la tion of peat and coaly de - pos its. The plant prob a bly grew near the shore of a small floodplain lake and fos sil ized in mud de liv ered by the flood ing of a nearby river.
Some con clu sions about palaeoclimate can be de liv ered by clay min er al ogy. The clay min er als in the Niek³añ pro file are largely de tri tal and show a gen er ally in sig nif i cant diagenetic over print (Brañski, 2009, 2014; Pieñkowski et al., 2014) due to mod er ate burial and the fairly closed hydrologic sys tem. There is no dis tinct and sys tem atic evo lu tion of clay-min eral as sem - blages from top to bot tom of the stud ied bore holes that are in - dic a tive of burial diagenesis. In the Niek³añ pro file and the whole Zagaje For ma tion, kaolinite gen er ally pre dom i nates (with an av er age con tent of 49–54%) over illite (av er age con tent of 33–46%, re spec tively). Chlorite is sub sid iary to a sig nif i cant ex - tent, and smectite is al most ab sent. Clay min eral data are re - flected in the ma jor el e ment geo chem is try and in the val ues of al ter ation in di ces, which are high in the Up per Rhaetian and Lower Hettangian strata. The Chem i cal In dex of Al ter na tion (CIA) ranges from 79 to 96, but in the vast ma jor ity of cases it ex ceeds 85, and these are due to re sid ual clays (Nesbitt and Young, 1982). The CIA shows a loss of Ca2+, K+and Na+in re la - tion to Al2+ (Nesbitt and Young, 1982: cap tion of fig. 2). Fluc tu a - tions in both the kaolinite/illite ra tio and CIA are very use ful for rec og niz ing cli mate fluc tu a tions (Brañski, 2009, 2014; Pieñ - kowski et al., 2014). In ter est ingly, at the in ter val of the oc cur - rence of the fern (41.8 m), these in di ces are char ac ter ized by a slightly lower kaolinite/illite ra tio (one of three min ima within the Zagaje For ma tion in Niek³añ), and the lower CIA (P. Brañski,
pers. comm., 2013) points to some what cooler (pre cisely, less hot) con di tions, while the hu mid ity re mained very high. The hab i tat of Matonia braunii can be char ac ter ized as wet and warm (but not hot) cli mate con di tions, cor re spond ing with the gen er ally sim i lar en vi ron men tal re quire ments of other Me so zoic ferns (Van Konijnenburg-van Cittert, 2002; Wang, 2002).
In Po land, Ju ras sic ferns are usu ally as so ci ated with al lu vial and lac us trine depositional sys tems in con ti nen tal set tings (Odrow¹¿, Huta OP-1 and Gromadzice – Holy Cross Mts.) or with deltaic depositional sys tems in coastal set tings (Szewna, DŸwiertnia, Chlewiska, Jêdrzejowice, Chmielów – Holy Cross Mts.; Siewierz and Zawiercie – Up per Silesia) (Pieñkowski, 2004; Barbacka et al., 2010, 2014a). This ac cords with gen eral ten den cies in fern en vi ron men tal pref er ences, since in the Early Ju ras sic of Eu rope, ferns were most di verse in al lu vial and deltaic en vi ron ments (Barbacka et al., 2014b).
CONCLUSIONS
– Matonia braunii (Göppert) Har ris has been re ported for the first time from Po land, from a bore hole in a new lo cal ity, Niek³añ. This ma te rial for the first time fully con firms Har ris’ sug - ges tions about the tax on omy of this spe cies.
– Tri an gu lar, smooth, trilete, kyrtomate spores iso lated from sporangia are most sim i lar to in situ spores ob tained from dif fer - ent fern spe cies of Matoniaceae and dis persed Dictyo - phyllidites mortoni (de Jer sey) Playford et Dettmann.
– Thanks to a de tailed sedimentological study of the bore - hole in Niek³añ, the en vi ron men tal re quire ments of this spe cies were pre cisely de fined for the first time. The fern oc cu pied high mois ture ar eas of the lake shore.
– Slightly lower kaolinite/illite ra tio in the in ter val of the oc - cur rence of the fern points to some what cooler (pre cisely, less hot) con di tions, while the hu mid ity re mained very high.
– The fern’s hab i tat was pro posed to be wet and warm (but not hot).
Ac knowl edge ments. The au thors ex press their thanks to K. Bóka for tak ing SEM pic tures of the sporangia, A. Sojka for her draw ings and M. Szewczyk for his ex cel lent macro pho to graphs.
We are also thank ful to B. Bomfleur for his will ing ness to dis cuss the mor phol ogy of sori and annuli. We are thank ful to M. Hodbod for in for ma tion on palynomorph as sem blage in the Niek³añ core and to R. Pointer and P. Palmer for lin guis tic cor rec tions. The au - thors grate fully ac knowl edge com ments by the Re view ers, E. Kustatscher and J.H.A. van Konijnenburg-van Cittert, who con trib uted to the manu script im prove ment dur ing the peer re - view pro cess. The study was sup ported by a grant fi nanced from the re sources of the Pol ish Na tional Sci ence Cen tre, granted on the ba sis of de ci sion no. DEC-2012/06/M/ST10/00478 and by the W. Szafer In sti tute of Bot any, Pol ish Acad emy of Sci ences through its stat u tory funds. GP re search was fi nan cially sup - ported by the In sti tute of Bot any at the Jagiellonian Uni ver sity (K/ZDS/005396, K/ZDS/006302, K/ZDS/006305).
REFERENCES
Appert, O., 1973. Die Pteridophyten aus dem Oberen Jura des Manamana in Südwest-Madagaskar. Schweizerische Paläonto - lo gische Abhandlungen, 94: 1–62.
Arndorff, L., 1993. Lat eral re la tions of deltaic palaeosols from the Lower Ju ras sic Rrnne For ma tion on the is land of Born holm, Den mark. Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - ogy, 100: 235–250.
Ar nold, C.A., 1956. Fos sil ferns of the Matoniaceae from North Amer ica. Jour nal of the Palaeontological So ci ety of In dia, 1:
118–121.
Ash, S.R., 1969. Ferns from the Chinle For ma tion (Up per Tri as sic) in the Fort Wingate area, New Mex ico. U.S. Geo log i cal Sur vey Pro fes sional Pa per, 613-D: 1–52.
Ash, S.R., 1972. Piazopteris branneri from the Lower Ju ras sic, Egypt. Re view of Palaeobotany and Palynology, 13: 147–154.
Ash, S.R., 1991. A new Ju ras sic Phlebopteris (Plantae, Filicales) from the Wallowa Terrane in the Snake River Can yon, Or e gon and Idaho. Jour nal of Pa le on tol ogy, 65: 322–329.
Ash, S., 2001. The fos sil ferns of Pet ri fied For est Na tional Park, Ar i - zona, and their paleoclimatological im pli ca tions. In: Pro ceed - ings of the 6th Fos sil Re source Con fer ence (ed. V.L. Santucci):
3–10. Geo logic Re sources Di vi sion Tech ni cal Re port, NPS/NRGRD/GR DTR -01/01.
Ash, S., Litwin, R.J., Tra verse, A., 1982. The Up per Tri as sic fern Phlebopteris smithii (Daugherty) Ar nold and its spores. Palyno - logy, 6: 203–219.
Aslan, A., Autin, W.J., 1999. Evo lu tion of the Ho lo cene Mis sis sippi River floodplain, Ferriday, Lou i si ana: in sights on the or i gin of fine-grained floodplains. Jour nal of Sed i men tary Re search, 69:
800–815.
Balme, B.E., 1995. Fos sil in situ spores and pol len grains: an annnoted cat a logue. Re view of Palaeobotany and Palynology, 87: 81–323.
Barbacka, M., 2011. Biodiversity and the re con struc tion of Early Ju - ras sic flora from the Mecsek Moun tains (south ern Hun gary).
Acta Palaeobotanica, 51: 127–179.
Barbacka, M., Ziaja, J., Wcis³o-Luraniec, E., Reymanówna, M., 2007. Hirmeriella muensteri (Schenk) Jung from Odrow¹¿ (Po - land), with fe male and male cones, and in situ Classopollis pol - len grains. Acta Palaeobotanica, 47: 339–357.
Barbacka, M., Ziaja, J., Wcis³o-Luraniec, E., 2010. Tax on omy and palaeo ec ol ogy of the Early Ju ras sic macroflora from Odrow¹¿, cen tral Po land. Acta Geologica Polonica, 60: 373–392.
Barbacka, M., Pacyna, G., Feldman-Olszewska, A., Ziaja, J., Bodor, E.R., 2014a. Tri as sic-Ju ras sic flora of Po land; floristical sup port of cli ma tic changes. Acta Geologica Polonica, 64:
281–308.
Barbacka, M., Bodor, E.R., Jarzynka, A., Kustatscher, E., Pacyna, G., Popa, M.E., Scanu, G.G., Thévenard, F., Ziaja, J., 2014b. Eu ro pean Ju ras sic flo ras: sta tis tics and paleoenviron - mental prox ies. Acta Palaeobotanica, 54: 173–195.
Barbacka, M., Popa, M.E., Mitka, J., Bodor, E., Pacyna, G., 2015.
Re la tion ships be tween eco sys tems and plant as sem blages as re sponses to en vi ron men tal con di tions in the Lower Ju ras sic of Hun gary and Ro ma nia. Acta Palaeobotanica, 55: 3–17.
Bartiromo, A., Barone Lumaga, M.R., Bravi, S., 2006. First find ing of a fos sil fern (Matoniaceae) in the paleontological site of Pietraroja (Benevento, South ern It aly). Bollettino della Societ´
Paleontologica Italiana, 46: 29–34.
Berry, E.W., 1919. A new Matonidium from Col o rado, with re marks on the dis tri bu tion of the Matoniaceae. Bul le tin of the Torrey Bo - tan i cal Club, 46: 285–294.
Bohacs, K., Suter, J.R., 1997. Se quence strati graphic dis tri bu tion of coaly rocks: fun da men tal con trols and paralic ex am ples.
AAPG Bul le tin, 81: 1612–1639.
Brañski, P., 2009. In flu ence of palaeoclimate con di tions and green - house ef fect on the Hettangian clay min eral as sem blages (Holy
Cross Mts. area, Pol ish Ba sin). Geo log i cal Quar terly, 53 (3):
363–368.
Brañski, P., 2014. Cli ma tic di sas ter at the Tri as sic-Ju ras sic bound - ary – a clay min er als and ma jor el e ments re cord from the Pol ish Ba sin. Geo log i cal Quar terly, 58 (2): 291–310.
Camus, J.M., 1990. Marattiaceae. In: The Fam i lies and Gen era of Vas cu lar Plants. Vol. I. Pteri do phytes and Gym no sperms (eds.
K.U. Kramer and P.S. Green): 174–180. Springer, Berlin, Hei - del berg.
Christenhusz, M.J.M., Chase, M.W., 2014. Trends and con cepts in fern clas si fi ca tion. An nals of Bot any, 113: 571–594.
Cleal, C.J., 1993. Pteridophyta. In: The Fos sil Re cord 2 (ed. M.J.
Benton): 779–794. Chap man and Hall, Lon don.
Couper, R.A., 1953. Up per Me so zoic and Caino zo ic spores and pol - len grains from New Zea land. New Zea land Geo log i cal Sur vey.
Paleontological Bul le tin, 22: 1–77.
Couper, R.A., 1958. Brit ish Me so zoic micro spores and pol len grains. A sys tem atic and strati graphic study. Palaeontographica Abt. B, 103: 75–179.
Czier, Z., 1994. On a new re cord of Selenocarpus muensterianus (Presl) Schenk from the Fire clay For ma tion of ªuncuius (Ro ma - nia) and the lower Li assic age of the flora. Re view of Palaeo - botany and Palynology, 82: 351–363.
De Jer sey, N.J., 1959. Ju ras sic spores and pol len grains from the Rose wood Coal field. Queensland Gov ern ment Min ing Jour nal, 60: 346–366.
Gierliñski, G., Pieñkowski, G., 1999. Di no saur track as sem blages from Hettangian of Po land. Geo log i cal Quar terly, 43 (3):
329–346.
Gierliñski, G., NiedŸwiedzki, G., Pieñkowski, G., 2001. Gi gan tic foot print of a theropod di no saur in the Early Ju ras sic of Po land.
Acta Palaeontologica Polonica, 46: 441–446.
Gierliñski, G., Pieñkowski, G., NiedŸwiedzki, G., 2004. Tetrapod track as sem blage in the Hettangian of So³tyków, Po land, and its paleoenvironmental back ground. Ichnos, 11: 195–213.
Givulescu, R., Popa, M., 1998. Aninopteris formosa Givulescu et Popa, gen. et sp. nov., a new Li assic matoniaceous ge nus and spe cies from Anina, Banat, Ro ma nia. Re view of Palaeobotany and Palynology, 104: 51–66.
Göppert, H.R., 1841. Les Gen res des Plantes Fossiles. Lief. 1–2.
Verlag von Henry und Co hen, Bonn.
Har ris, T.M., 1931. The Fos sil Flora of Scores by Sound East Green - land. Part 1: Cryp to gams (Ex clu sive of Lycopodiales). Meddelel - ser om GrÝnland, 85: 1–102.
Har ris, T.M., 1961. The York shire Ju ras sic Flora I: Thallophyta – Pteridophyta. Trust ees of the Brit ish Mu seum (Nat u ral His tory), Lon don.
Har ris, T.M., 1980. The York shire Ju ras sic fern Phlebopteris braunii (Goeppert) and its ref er ence to Matonia R. Br. Bul le tin of the Brit ish Mu seum (Nat u ral His tory) Ge ol ogy Se ries, 33: 295–311.
Hirmer, M., Hörhammer, L., 1936. Morphologie, Systematik und geographische Verbreitung der fossilen und rezenten Mato - niaceen. Palaeontographica Abt. B, 81: 1–70.
Iwasuki, K.I., 1990. Hymenophyllaceae. In: The Fam i lies and Gen - era of Vas cu lar Plants. Vol. I. Pteri do phytes and Gym no sperms (eds. K.U. Kramer and P.S. Green): 157–162. Springer, Berlin, Hei del berg.
Jansonius, J., Hills, L.V., 1976. Gen era File of Fos sil Spores and Pol len. Spe cial Pub li ca tion, De part ment of Ge ol ogy Uni ver sity of Cal gary, Can ada.
Jarzynka, A., 2012. rodkowojurajska flora glinek grojeckich – taksonomia i paleoekologia (in Pol ish). Ph.D. the ses. Ar chives W. Szafer In sti tute of Bot any, Pol ish Acad emy of Sci ences, Kraków.
Juhász, M., 1979. Dis persed Matoniaceae spores from the Hun gar - ian Lower and Mid dle Cre ta ceous sed i ments. Acta Biologica Szegedensis, 25: 33–47.
866 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
Klavins, S.D., Tay lor, T.N., Tay lor, E.L., 2004. Matoniaceous ferns (Gleicheniales) from the Mid dle Tri as sic of Antarctica. Jour nal of Pa le on tol ogy, 78: 211–217.
Kozur, H.W., Weems, R.E., 2010. The biostratigraphic im por tance of conchostracans in the con ti nen tal Tri as sic of the north ern hemi sphere. Geo log i cal So ci ety Spe cial Pub li ca tions, 334:
315–417.
Kramer, K.U., 1990. Matoniaceae. In: The Fam i lies and Gen era of Vas cu lar Plants. Vol. I. Pteri do phytes and Gym no sperms (eds.
K.U. Kramer and P.S. Green): 183–185. Springer, Berlin, Hei - del berg.
Kvaèek, J., Dašková, J., 2010. Konijnenburgia, a new ge nus of the fern fam ily Matoniaceae. Re view of Palaeobotany and Palyno - logy, 158: 308–318.
Leschik, G., 1955. Die Keuperflora von Neuewelt bei Basel. II. Die Iso- und Mikrosporen. Schweizerische Paläontologische Abhan dlungen, 72: 1–70.
Litwin, R.J., 1985. Fer tile or gans and in situ spores of ferns from the Late Tri as sic Chinle for ma tion of Ar i zona and New Mex ico, with dis cus sion of the as so ci ated dis persed spores. Re view of Palaeobotany and Palynology, 44: 101–146.
Lund, J.J., 1977. Rha etic to Lower Li assic palynology of the on - shore south-east ern North Sea Ba sin. Danmarks Geologiske Undersrgelse II Rk., 109: 1–129.
Lundblad, A.B., 1950. Stud ies in the Rhaeto-Li assic flo ras of Swe - den. I. Pteridophyta, Pteridospermae and Cycadophyta from the min ing dis trict of NW Scania. Kungliga Svenska Vetenska psa - kademiens Handlingar, Ser., 4: 1–82.
Makarewiczówna, A., 1928. Etude sur la flore fossile du lias inférieur des en vi rons d’Ostrowiec (in Pol ish with French sum - mary). Prace Towarzystwa Przyjació³ Nauk w Wilnie, Wydzia³ Nauk Matematycznych i Przyrodniczych, Prace Zak³adu Geolo - gicznego Uniwersytetu St. Batorego w Wilnie, 4: 1–49.
Mamczar, J., 1986. Palynostratigraphy of the Up per most Ju ras sic and Lower Cre ta ceous in the Pol ish Low lands (in Pol ish with Eng lish sum mary). Prace Instytutu Geologicznego, 116: 1–53.
McCabe, P.J., 1984. Depositional en vi ron ments of coal and coal-bear ing strata. IAS Spe cial Pa per, 7: 13–42.
Miner, E.L., 1935. Palaeobotanical ex am i na tions of Cre ta ceous and Ter tiary coals: II. Cre ta ceous and Ter tiary coals from Montana. Amer i can Mid land Nat u ral ist, 16: 616–625.
Mirzaie Ataabadi, M., Djafarian, M., Mohammadalizadeh, J., 2005. Oc cur rence of Aninopteris (Matoniaceae) from the Mid dle Ju ras sic of East cen tral Iran. Palaeobotanist, 54: 99–106.
Nagalingum, N.S., Cantrill, D.J., 2006. Early Cre ta ceous Gleiche - niaceae and Matoniaceae (Gleicheniales) from Al ex an der Is - land, Antarctica. Re view of Palaeobotany and Palynology, 138:
73–93.
Naugolnykh, S.V., Pronin, A.P., 2015. A new matoniaceous fern from the Up per Tri as sic of the Cas pian De pres sion in the con - text of florogenetic pro cesses of tran si tion from the Pa leo zoic to Me so zoic. Paleontological Jour nal, 49: 326–336.
Nesbitt, H.W., Young, G.M., 1982. Early Pro tero zoic cli mates and plate mo tions in ferred from ma jor el e ment chem is try of lutites.
Na ture, 299: 715–717.
NiedŸwiedzki, G., 2011. Di no saur tracks from the Early Ju ras sic eco sys tem of So³tyków, Holy Cross Moun tains (in Pol ish with Eng lish sum mary). Biuletyn Pañstwowego Instytutu Geolo - gicznego, 447: 49–98.
Or³owska-Zwoliñska, T., 1983. Palynostratigraphy of the up per part of Tri as sic epicontinental sed i ments in Po land (in Pol ish with Eng lish sum mary). Prace Instytutu Geologicznego, 104:
1–89.
Pacyna, G., 2013. Crit i cal re view of re search on the Lower Ju ras sic flora of Po land. Acta Palaeobotanica, 53: 141–163.
Pieñkowski, G., 2004. The epicontinental Lower Ju ras sic of Po - land. Pol ish Geo log i cal In sti tute Spe cial Pa pers, 12: 1–122.
Pieñkowski, G., Gierliñski, G., 1987. New finds of di no saur foot - prints in Li assic of the Holy Cross Moun tains and its palaeo - environmental back ground. Przegl¹d Geologiczny, 35: 199–205.
Pieñkowski, G., NiedŸwiedzki, G., Waksmundzka, M., 2012.
Sedymentological, palynological and geo chem i cal stud ies of
the ter res trial Tri as sic-Ju ras sic bound ary in north west ern Po - land. Geo log i cal Mag a zine, 149: 308–332.
Pieñkowski, G., NiedŸwiedzki, G., Brañski, P., 2014. Cli ma tic re - ver sals re lated to the Cen tral At lan tic mag matic prov ince caused the end-Tri as sic bi otic cri sis – ev i dence from con ti nen tal strata in Po land. GSA Spe cial Pa per, 505: 263–286.
Playford, G., Dettmann, M.E., 1965. Rhaeto-Li assic plant microfossils from the Leigh Creek Coal Mea sures, South Aus - tra lia. Senckenbergiana Lethaea, 46: 127–181.
Popa, M., 1997. Li assic ferns from the Steierdorf For ma tion, Anina, Ro ma nia. Mededelinger Nederlands Instituut voor Toegepaste Geowetenschappen TNO, 58: 139–147.
Presl, K.B., 1848. Die Gefässbündel im stipes der farrn. Abhand - lungen der Königlichen Böhmischen Gesellschaft der Wissen - schaften, 5: 307–356.
Raciborski, M., 1891. Flora retycka pó³nocnego stoku Gór Œwiê - tokrzyskich (in Pol ish). Rozprawy Wydzia³u matematyczno -przy - rodniczego Akademii Umiejêtnoœci w Krakowie, 23: 292–326.
Raciborski, M., 1892. Przyczynek do flory retyckiej Polski (in Pol - ish). Rozprawy Wydzia³u matematyczno-przyrodniczego Aka - demii Umiejêtnoœci w Krakowie, 22: 345–360.
Raciborski, M., 1894. Flora kopalna ogniotrwa³ych glinek krako - wskich. Czêœæ I. Rodniowce (Archaegoniatae) (in Pol ish).
Pamiêtnik matematyczno-przyrodniczy Akademii Umiejêtnoœci, 18: 1–101.
Reymanówna, M., 1963. The Ju ras sic flora from Grojec near Cra - cow in Po land. Part I. Acta Palaeobotanica, 4: 9–48.
Reymanówna, M., 1992. Two co ni fers from the Li assic flora of Odrow¹¿ in Po land. In: Palaeovegetational de vel op ment in Eu - rope and re gions rel e vant to its palaeofloristic evo lu tion (ed. J.
Kovar-Eder): 307–311. Pro ceed ings Pan-Eu ro pean Palaeobo - ta nical Con fer ence. Mu seum of Nat u ral His tory, Vi enna.
Rogalska, M., 1954. Spore and pol len anal y sis of the Li assic coal of Blanowice in Up per Silesia (in Pol ish with Eng lish sum mary.) Biuletyn Instytutu Geologicznego, 89: 1–46.
Rogalska, M., 1956. Spore and pol len anal y sis of the Li assic de pos - its of the Mroczków-Rozwady area in the Opoczno dis trict (in Pol ish with Eng lish sum mary). Biuletyn Instytutu Geologiczego, 104: 1–89.
Rogalska, M., 1976. Stra tig ra phy of the Lower and Mid dle Ju ras sic in the Pol ish Low lands on the ba sis of spore and pol len anal y sis (in Pol ish with Eng lish sum mary). Prace Instytutu Geolo gi czne - go, 78: 1–61.
Roth well, G.W., Stockey, R.A., 2008. Phy log eny and evo lu tion of ferns: a paleontological per spec tive. In: Bi ol ogy and Evo lu tion of Ferns and Lycophytes (eds. T.A. Ranker and Ch.H. Haufler):
332–366. Cam bridge Uni ver sity Press, Cam bridge.
Rushforth, S.R., 1970. Notes on the fern fam ily Matoniaceae from the West ern United States. Brigham Young Uni ver sity Ge ol ogy Stud ies, 6: 3–34.
Schenk, A., 1867. Die fossile flora der Grenzschichten des Keupers und Lias Frankens. Kreidel’s Verlangen, Wiesbaden.
Schmidt, A.R., Dörfelt, H., 2007. Ev i dence of Ce no zoic Mato - niaceae from Bal tic and Bitterfeld am ber. Re view of Palaeo - botany and Palynology, 144: 145–156.
Schnei der, H., Schuettpelz, E., Pryer, K.M., Cranfill, R., Magallón, S., Lupia, R., 2004. Ferns di ver si fied in the shadow of an gio sperms. Na ture, 428: 553–557.
Schweitzer, H-J., 1978. Die Räto-Jurassischen Floren des Iran und Afganistans. 5. Todites princeps, Thaumatopteris brauniana und Phlebopteris polypodioides. Palaeontographica Abt. B, 168: 17–60.
Schweitzer, H-J., Schweitzer, U., Kirchner, M., Van Konijnen - burg -van Cittert, J.H.A., Van der Burgh, J., Ashraf, R.A., 2009. The Rhaeto-Ju ras sic flora of Iran and Af ghan i stan. 14.
Pterophyta – Leptosporangiatae. Palaeontographica Abt. B, 279: 1–108.
Seward, A.C., 1899. On the struc ture and af fin i ties of Matonia pectinata, R. Br., with notes on the geo log i cal his tory of the Matonineae. Philo soph i cal Trans ac tions of the Royal So ci ety B:
Bi o log i cal Sci ences, 191: 171–209.
Seward, A.C., 1900. The Ju ras sic flora I. The York shire coast. Cat a - logue of the Me so zoic Plants in the De part ment of Ge ol ogy, Brit - ish Mu seum (Nat u ral His tory), 3: 1–341.
Skog, J.E., 1988. Re as sign ment of Aspidium heterophyllum to a new ge nus in the fam ily Matoniaceae. Amer ica Jour nal of Bot - any, 75: 1120–1129.
Skog, J.E., 2001. The biogeography of Me so zoic leptosporangiate ferns re lated to ex tant ferns. Brittonia, 53: 236–269.
Skog, J.E., Litwin, R.L., 1995. Re ex am i na tion of the fos sil fern ge - nus Dryopterites Berry based on re pro duc tive char ac ter is tics.
Re view of Palaeobotany and Palynology, 89: 235–256.
Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schnei der, H., Wolf, P.G., 2006. A clas si fi ca tion for ex tant ferns. Taxon, 55:
705–731.
Tay lor, T.N., Tay lor, E.L., Krings, M., 2009. Paleobotany: the bi ol - ogy and evo lu tion of fos sil plants. Sec ond Edi tion. Ac a demic Press, Burlington.
Thomson, P.W., Pflug, H., 1953. Pol len und Sporen des Mitteleuro - päischen Tertiärs. Palaeontographica Abt. B, 94: 1–138.
Tidwell, W.D., Ash, S.R., 1994. A re view of se lected Tri as sic to Early Cre ta ceous ferns. Jour nal of Plant Re search, 107: 417–442.
Tidwell, W.D., Skog, J.E., 1992. Two new fos sil matoniaceous stem gen era from Tas ma nia, Aus tra lia. Re view of Palaeobotany and Palynology, 70: 263–277.
Tralau, H., 1965. Phlebopteris angustiloba (Presl) Hirmer et Hörhammer (Matoniaceae) from “Olstorp” Shaft, Bjuv, Scania.
Botaniska Notiser, 118: 373–376.
Wallich, N., 1830. Plantae Asiaticae Rariores, or, De scrip tions and Fig ures of a Se lect Num ber of Un pub lished East In dian Plants.
Part 1. Treuttel and Würtz, Lon don.
Wang, Y., 2002. Fern eco log i cal im pli ca tions from the Lower Ju ras - sic in West ern Hubei, China. Re view of Palaeobotany and Palynology, 119: 125–141.
Wang., Y., Mei, S., 1999. Fer tile or gans and in situ spores of matoniaceous fern from the Lower Ju ras sic of West Hubei. Chi - nese Sci ence Bul le tin, 44: 1333–1337.
Wcis³o-Luraniec, E., 1992. Flora from Odrow¹¿ in Po land – a typ i cal Lower Li assic Eu ro pean flora. In: Palaeovegetational de vel op - ment in Eu rope and re gions rel e vant to its palaeofloristic evo lu - tion (ed. J. Kovar-Eder): 331–335. Pro ceed ings Pan-Eu ro pean Palaeo botanical Con fer ence. Mu seum of Nat u ral His tory, Vi enna.
Van Konijnenburg-van Cittert, J.H.A., 1993. A re view of the Matoniaceae based on in situ spores. Re view of Palaeobotany and Palynology, 78: 235–267.
Van Konijnenburg-van Cittert, J.H.A., 2002. Ecol ogy of some Late Tri as sic to Early Cre ta ceous ferns in Eur asia. Re view of Palaeobotany and Palynology, 119: 113–124.
Van Konijnenburg-van Cittert, J.H.A., Kurmann, M.H., 1994.
Com par a tive ultrastructure of liv ing and fos sil matoniaceous spores. In: Ultrastructure of Fos sil Spores and Pol len (eds. M.H.
Kurmann and J.A. Doyle): 67–86. Royal Bo tanic Gar den, Kew.
Ziaja, J., 1992. The Lower Li assic microflora from Odrow¹¿ in Po - land. In: Palaeovegetational de vel op ment in Eu rope and re - gions rel e vant to its palaeofloristic evo lu tion (ed. J. Kovar-Eder):
337–340. Pro ceed ings Pan-Eu ro pean Palaeobotanical Con fer - ence. Mu seum of Nat u ral His tory, Vi enna.
Ziaja, J., 2006. Lower Ju ras sic spores and pol len grains from Odrow¹¿, Me so zoic mar gin of the Holy Cross Moun tains, Po - land. Acta Palaeobotanica, 46: 3–83.
Ziegler, P.A., 1990. Geo log i cal At las of West ern and Cen tral Eu rope – 2nd edi tion. Shell Int. Petr. Maatsch. B.V.
868 Maria Barbacka, Grzegorz Pacyna, Grzegorz Pieñkowski and Jadwiga Ziaja
