ArticlePDF Available

Abstract and Figures

A new palaeobotanical site was recovered at artificial excavations within the city district of Ústí nad Labem called Mojžíř in northern Bohemia (Czech Republic) that yielded slabs of Oligocene tephritic pyroclastic rocks with plant impressions. The fossiliferous layer belongs to the Děčín Formation of the České středohoří Mts. dated to the Oligocene (30.8–24.7 Ma). One of the recovered fossils belongs to a new leptosporangiate fern matching modern representatives of Polystichum Roth (Dryopteridaceae Herter). These frond fragments are partly fertile and allow studying the position of sori and details of indusia and spores. This fossil fern is described here as a new species of Polystichum, P. pacltovae Kvaček sp. nov., which matches the modern European species of Polystichum , namely Polystichum braunii (Spenner) Fée . Accompanying plant fossils include another fern frond described as " Aspidium " elongatum Heer (non Swartz) of unknown affinities, fragmentary needles assigned to Pinus cf. hepios (Unger) Unger of the Pinaceae and leaf impressions of Betulaceae comparable to Betula brongniartii Ettingshausen.
Content may be subject to copyright.
E
N. Jb. Geol. Paläont. Abh.
www.schweizerbart.deE. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany
1. Introduction
Fern fossils comparable to the Dryopteridaceae are
quite rarely recorded in the Palaeogene. A new locality


(Czech Republic) has yielded fertile foliage worth of
a detailed study including sporangia. Previous records
-

were explored since the 18th
macrofossils described by  (1866),
 (1869),  (1891), -
 (1898),  (1909) and others from this
part of the Czech Republic were sterile or not treated
in detail. Only studies of Polypodium ( 2001)
      
this respect. This study of the newly recovered spec-
imens treated below is a continuation of such efforts.
2. Material and methods
        from

        

    
light grey trachybasaltic pyroclastics containing plant
impressions. It does not crop out in the surround-
ings. According to the local geological map of the
 -
 
        
2000

sensu  (2000) or
 et al. (1999
and  et al. (2001), more spread near the town

Stuttgart,
DOI:
A new Oligocene fern of Dryopteridaceae from the České středohoří
Mts (Czech Republic)
Zlatko Kvaček and Vasilis Teodoridis
With 3 figures
Abstract: 
-


fossils belongs to a new leptosporangiate fern matching modern representatives of Polystichum 
(Dryopteridaceae ). These frond fragments are partly fertile and allow studying the position
of sori and details of indusia and spores. This fossil fern is described here as a new species of Polys-
tichum, P. pacltovae  sp. nov., which matches the modern European species of Polystichum,
namely Polystichum braunii () . Accompanying plant fossils include another fern frond
Aspidium elongatum  (non 
assigned to Pinus cf. hepios ()     
comparable to Betula brongniartii .
Key words: Polystichum, Dryopteridaceae, fertile foliage, Oligocene, Czech Republic.
295/1 (2020), 9–16
January 2020
Article
© 2020
10.1127/njgpa/2020/0864 $ 2.00/2020/08640077-7749
eschweizerbart_xxx
10 Z. Kvaček and V. Teodoridis
       
switched, which is a typical preservation for plant

foliage of a pine and dicotyledonous plants has been
recovered. The leaf material was described using the
current morphological terminology published by 
et al. (2009). The collection was passed over into the

     -
ment details of the fern fossils by various reproduc-
tion technics and the palynological analysis. Despite
repeated maceration of samples by  ,
no in situ spores have been obtained.
3. Systematic palaeobotany

-
gene of Europe. The accompanying plant elements are
additionally reported with short comments.
Polypodiopsida ,  &
W. 
Dryopteridaceae 
Polystichum 
Polystichum pacltovae  sp. nov.
Figs. 11.8, 2.1, 2.2
Etymology: Epithet commemorating Czech palynologist,
-
ty, who died suddenly in Prague on the 2nd February 2019.
Holotype:
11.8.
Other material:
2.1, 2

Type locality:
Type horizon:      

Description: Fern foliage fragmentary, preserved parts of
-
row elongate, up to 60 mm long and 20 mm broad, pinnules
almost of the same size, elongate or slightly falcate, gradu-
ally diminishing in size towards tips of pinnae, arranged in
  
-
date to obliquely truncate, symmetrical, rounded or shortly
decurrent at base, densely spiny toothed on margin, vena-
tion simple free, midrib straight, secondary veins craspe-
  

in 1 row on each side of midrib, in up to 6 pairs or solitary
 
in diameter, centrally attached, partly leaving only rounded
traces.
Discussion:  
and rounded sporangia with peltate indusia refer this fern
to the genus Polystichum. The spiny- toothed margin of ses-
sile pinnules places it near the modern species Polystichum
braunii () , a circumboreal montane to sub mon-
tane evergreen fern growing in deciduous forests (see Fig.
22.6). Different in our opinion are also similar Polys-
tichum setiferum ()  ex , a small
      
  
and strong asymmetry of basal pinnules, and Polystichum
aculeatum ()  ex  (syn. Polystichum
lobatum () ) from western Asia and most of
Europe, which is characterized by differently sized pinnules
within the pinna. Preservation of the fossils at hand does not
       -
quence data of the whole plant. Therefore, it would be futile
to attempt a more satisfactory comparison with the 200 ex-
tant species distributed worldwide ( & 
;  I 2016).
Polypodiopsida familiae et genera indeterminate
Aspidiumelongatum  sensu 
Fig. .1, .2
 Aspidium elongatum , p. 96.
Material:
Description: A single fern frond studied is sterile, of the in-



length, rounded at tips. The rachis is straight, midrib of pin-
Fig. 1. 1–8Polystichum pacltovae  
  

1
sorus from Fig. 1
eschweizerbart_xxx
11
A new Oligocene fern of Dryopteridaceae from the České středohoří Mts (Czech Republic)
eschweizerbart_xxx
12
nae slightly sinuous, venation of pinnules not well visible,
probably reticulate.
Discussion:        
-
       ;
 2001
Pinopsida 
Pinaceae 
Pinus
Pinus cf. hepios () 
Fig. .6, 
Material:

Description: Needle leaves in pairs, incomplete in length,
ca. 1 mm broad, entire- margined.
Fig. 2. 1, 2Polystichum pacltovae 
3–5Polystichum braunii () , Rossia subcarpatica,
& 

Z. Kvaček and V. Teodoridis
eschweizerbart_xxx
13
Fig. 3. 1, 2“Aspidium” elongatum 
3–5Betula brongniartii 
-
6, 7Pinus cf. hepios ()  and Betula brongniartii ,

A new Oligocene fern of Dryopteridaceae from the České středohoří Mts (Czech Republic)
eschweizerbart_xxx
14
Discussion: The sporadically occurring pine foliage is
     
-
        
et al. 2018  
 1901). The nee-
dles are associated with cones of Pinus ornata ()
 (&  2008


Betula 
Betula brongniartii 
Fig. 
Material:       
         

Description:


apex acute, margin double sharply serrate, venation craspe-
dodromous, midrib straight, moderate, strong, secondary
veins alternate or subopposite at basal part, thinner, straight,
  -
served, tertiary veins alternate percurrent, straight or curved,
venation of the higher orders poorly visible, regular polyg-
onal reticulate.
Discussion:      Betula
brongniartii match in most respects fossils of birch foli-

 et al. 2018
( et al. 2001-
sil species is characterized particularly by strongly double
serrate margin recalling hornbeam foliage.
4. Notes on the occurrences of ferns in
the Palaeogene of North Bohemia and
adjacent Saxony
Fern fossils are rare in the magmatic complex of the
-
ony. No monographic treatment is available so far, but
     
and partly illustrations of fern macrofossilis besides
       -
ered ferns from there are two: a thelypterid species
called Pronephrium stiriacum () &
  and newly assigned to Cyclosorus
stiriacus ()  &  (
2001&
 2011) and Oligocene sites Kundratice
( &  1998  -
 &     et al.
), as well as Osmunda lignitum () 
 & -
 2011), Kundratice ( &  1998)
& -

as Woodwardia muensteriana ( ) 
   et al. 2018), Polypodium radonii
      2001,
 et al. 2006) and Rumohra recentior ()
&  2011-
& -

       -
 (1891, ) and  (1909) is so far
incompletely revisited. None of the so far described
ferns is related to Polystichum pacltovae, which is ob-
viously a novelty.
According to the accompanying leaf assemblage,
Polystichum pacltovae can be interpreted as a zonal
fern of the ground cover in the broad- leaved decidu-

substrate connected with slope vegetation in volcanic
area.
Acknowledgements


  , who tried to obtain
spores in situ. Curators of the palaeontological collections
 

are also the suggestions and notes made by two reviewers,
namely   and  , on the

        
         

Z. Kvaček and V. Teodoridis
eschweizerbart_xxx
15
References
      

14
 ,   &     
       -

18

      -
75
 ,  ,  ,   & -
      
9
  (2001): Cainozoic ferns and their dis-
   53    

, , , , -
,  & 

Press).

 -

Dresden, 1891
      
-
15
   


1
   (1866): Die fossile Flora des Ter-
       -
    
naturwissenschaftliche Klasse, 26
   (1869): Die fossile Flora des Ter-
       -
    
naturwissenschaftliche Klasse, 29
,  &  (2001): 19th century’s
palaeobotanical types and originals of the Hungarian
-

  &     -
 
42
  (2001): A new fossil species of Polypodium
 -
112

 &  (1998): The Oligocene volcanic
      
-
54
 & -
        
      

60
 & 

67
,  &  (2018): Review

 
74
 & -
-
nus Polystichum
90    


-

Dresden, 1900
       
 
54


       
26
      
-
     

 ,   &   (2006): Oligo-
cene megafossil plant remains and environment from

     -
47:

  &      -
       -
       -
tologie, Abhandlungen, 250   

, ,  & -


27
 &   
      
63

       -

A new Oligocene fern of Dryopteridaceae from the České středohoří Mts (Czech Republic)
eschweizerbart_xxx
16
Addresses of the authors:
-

  

  (corresponding author), Depart-
 
     -
-

Z. Kvaček and V. Teodoridis
eschweizerbart_xxx
Article
A comprehensive anatomical and mineralogical study of fossil wood fragments from fields in the vicinity of Bečov and Břvany villages (NW Bohemia, Czech Republic) indicates that Taxodioxylon gypsaceum (Cupressaceae s.l.) predominates, but also identifies another coniferous wood: Pinuxylon parryoides (Pinaceae) and three angiosperms Quercoxylon böckhianum, Castanoxylon bavaricum and Lithocarpoxylon sp. (all Fagaceae). This paper therefore presents the first occurrence of Pinaceae and Fagaceae fossil wood in the volcanic rocks of the České Středohoří Mts. as well as its youngest palaeobotanical record in general, late Oligocene in age (26.56 ± 0.38 Ma). The samples were buried by alkaline pyroclastic deposits and were mineralized by carbonates. Two distinct depositional processes burying the fossil woods were identified. Closer to the vent, the woods occur in a nearvent pyroclastic fall deposits of the former pyroclastic cone, whereas more distant sites consist of pyroclastic flow deposits. Carbonate mineralization mostly consists of dolomite, but subordinate amounts of magnesite (likely the first time this is documented in fossil wood) as well as calcite and siderite are present. Only one sample collected in the same area, bearing clear signs of riverbed transport (Lithocarpoxylon sp.), was perfectly silicified, but its origin remains unclear.
Article
Full-text available
This complete list of publications of Professor RNDr. Zlatko Kvaček, DrSc. (1937 – 2020) is an addition and supplementary continuation of the memorial volumes of Fossil Imprint (2021/2 and 2022/1) devoted to his work and life. The present list is compiled from three published sources: (1) in Acta Musei Nationalis Pragae, Ser. B – Historia Naturalis in 2007 (Kvaček, J.: Bibliography of Prof. RNDr. Zlatko Kvaček, DrSc. – Acta Musei Nationalis Pragae, Ser. B – Historia Naturalis, 63(2-4): 75–83), (2) in Fossil Imprint in 2018 (Sakala, J.: Bibliography of Prof. RNDr. Zlatko Kvaček, DrSc. (2007 – 2017). – Fossil Imprint, 74(1-2): 4–7) and (3) in Fossil Imprint in 2020 (Sakala, J.: Bibliography of Prof. RNDr. Zlatko Kvaček, DrSc. (2017 – 2020). – Fossil Imprint, 76(2): viii).
Article
Full-text available
The Oligocene palaeontological locality on Matrý Hill near Sebuzín in the České středohoří Mts., North Bohemia, belongs to the Děčín Formation and is dated to 30.8-24.7 Ma according to the regional stratigraphy. It has yielded, in addition to insects from the Heteroptera group, a fossil bee Apis petrefacta and palaeobatrachid frogs, also numerous plant remains. Their recovery began in 1996. The plant fossil assemblage consists mostly of leaf impressions, occasionally accompanied by casts or impressions of fruits. Noteworthy are records of a fern Woodwardia muensteriana, conifers Pinus cf. rigios, P. cf. hepios, Calocedrus suleticensis, Tetraclinis salicornioides, Torreya bilinica, cf. Cephalotaxus parvifolia and numerous angiosperms, e.g. Liriodendron haueri, Daphnogene cinnamomifolia, Platanus neptuni, Cercidiphyllum crenatum, Sloanea artocarpites, Ulmus pyramidalis, Celtis pirskenbergensis, Carya fragiliformis, C. quadrangula, Betula brongniartii, B. dryadum, Alnus rhenana, A. cf. kefersteinii, Carpinus grandis, Ostrya atlantidis, Acer crenatifolium, A. cf. palaeosaccharinum, A. integrilobum and Craigia bronnii. Several angiosperm foliage specimens of both monocots and dicots have not yet been identified to a particular genus and species. The fossil plant assemblage at Matrý corresponds to two vegetation types, i.e. a zonal riparian forest and zonal mixed mesophytic forest, as corroborated by the Integrated Plant Record vegetation analysis. The vegetation thrived under a humid climate, characterized by average values of MAT (13.4 °C), WMMT (23.8 °C), CMMT (3.6 °C) and MAP (1,117 mm). The Matry fossil flora is similar in composition to the flora of Žichov from the same Oligocene Děčín Formation in the České středohoří Mts.
Article
Full-text available
Phylogeny has long informed pteridophyte classification. As our ability to infer evolutionary trees has improved, classifications aimed at recognizing natural groups have become increasingly predictive and stable. Here, we provide a modern, comprehensive classification for lycophytes and ferns, down to the genus level, utilizing a community-based approach. We use monophyly as the primary criterion for the recognition of taxa, but also aim to preserve existing taxa and circumscriptions that are both widely accepted and consistent with our understanding of pteridophyte phylogeny. In total, this classification treats an estimated 11 916 species in 337 genera, 51 families, 14 orders, and two classes. This classification is not intended as the final word on lycophyte and fern taxonomy, but rather a summary statement of current hypotheses, derived from the best available data and shaped by those most familiar with the plants in question. We hope that it will serve as a resource for those wanting references to the recent literature on pteridophyte phylogeny and classification, a framework for guiding future investigations, and a stimulus to further discourse
Article
This contribution proposes the lithostratigraphy of the Tertiary volcanosedimentary complex of the Ceske stredohori Mts. The superficial volcanic products of the Ceske stredohori Mts. are divided into three formations. The Usti Fm. (36.1 - 25.5 Ma) includes basanites and their concomitant volcaniclastics including fossiliferous sediments, and represents products of early rift development - the filling of a rift valley. The Decin Fm. (30.8 - 24.7 Ma) is composed of trachybasalts and trachybasaltic resedimented tuffs, which are erosional relics of a composite volcano - the product of more advanced stage of rift evolution. The Dobrna Fm. (24.0 - 19.3? Ma), represented by basanitic lavas, was produced as a result of a possible remobilization of the magma chamber. All these formations represent one volcanic cycle (uppermost Eocene - lowermost Miocene, with a maximum in the Oligocene). The basanitic intrusions penetrating the Most Fm. sediments are comprised in the fourth lithostratigraphic unit, the Strbice Fm. (13.4 - 9.0? Ma). It is most probably a product of a younger volcanic cycle (Middle Miocene) and extends to the Most Basin, too. The Usti Fm is largely correlated with the Strezov Fm. of the Most Basin.
Article
The site of Oligocene plants, the Holý Kluk hill near Proboštov, which has been lately recovered for new collecting after more than 100 years, is characterised in terms of the flora, vegetation and environment. The flora includes 2 ferns, 3 conifers and more than 60 angiosperms. The reconstructed vegetation corresponds to the Mixed Mesophytic Forest with a considerable representation of thermophilic elements surviving from the latest Eocene. The site arose in a lake filled with volcanogenic deposits during a calm period of the volcanic activity about 29-30 MA according to the radiometric data. The composition of the flora and the aspect of the vegetation corroborate the subtropical climate of this time interval with frostless winters and a high amount of the annual precipitation.
Article
The phytogeographic distribution of Cainozoic ferns is reported based upon a critical re-appraisal of the macrofossil and mesofossil record also taking account of evidence from a few highly diagnostic spores. Well-documented circum-Arctic Cainozoic floras show ferns (Woodwardia, Onoclea, Osmunda, Coniopteris and to a lesser extentAzolla) distributed around the pole to very high paleolatitudes. Some ferns are shared between the mid-paleolatitudes of North America and Europe as would be predicted from the distributions of other biota. Evidence for the composition of Cainozoic fern floras is minimal in some regions (e.g., Antarctica, Central and South America, Africa, India, South East Asia), so the absence of fern fossils from these areas has no biogeographical significance. Matoniaceae were abundant in the preceding Mesozoic. However, the absence of Cainozoic macrofossils, and the fact that no CainozoicMatonisporites spores areMatonia-like, indicates that Matoniaceae had attained their modern relict distribution by, or very early in, the Cainozoic. The important Mesozoic families Marattiaceae and Dipteridaceae are also not represented by Cainozoic macrofossils. They probably also showed Cainozoic restriction but spores are not sufficiently diagnostic to enable testing of this hypothesis. Other ferns, which were also important in the Mesozoic (e.g., Dicksoniaceae, Gleicheniaceae), have patchy, equivocal, or inadequately published Cainozoic records. The dispersed spore record may provide an opportunity to track Cainozoic Gleicheniaceae but this approach is not without problems. Most well-represented Cainozoic fern families, genera and subgenera show widespread Cainozoic ranges, typically with considerable range extensions over their living relatives, both onto other continents and north and south to higher paleolatitudes. These include Schizaeaceae (Lygodium, Anemia, and the extinctRuffordia), Osmundaceae (Osmunda), Pteridaceae (Acrostichum), Thelypteridaceae (Cyclosorus), Lophosoriaceae (Lophosoria), Cyatheaceae (theCnemidaria/Cyathea decurrens clade) and the heterosporous water fernAzolla (Azollaceae). A few well-represented ferns show Cainozoic distributions similar to those of the present day (e.g.,Salvinia [Salviniaceae] andCeratopteris [Pteridaceae] (the latter by the Neogene and based only on spores]) but even these had slightly broader ranges in the Cainozoic. Some Cainozoic ferns have apparently local distributions, e.g.,Blechnum dentatum (Blechnaceae) in Europe; and others are so far represented at only one or few sites, e.g.,Dennstaedtiopsis (Dennstaedtiaceae),Botrychium (Ophioglossales),Grammitis (Grammitidaceae), andMakotopteris andRumohra (Dryopteridaceae). Cainozoic fossils assigned toDryopteris (and some other dryopteroids) require revision along with those of Thelypteridaceae, the latter having high potential to provide useful paleobiogeographic evidence, at least of theCyclosorus group. Cainozoic records of Hymenophyllaceae and Polypodiaceae are here considered unconfirmed.