Kladno Formation

The Kladno Formation is a Carboniferous formation in the Czech Republic which preserves a wide array of temnospondyls, lepospondyls and fish,^[1] among other fauna.

Kladno Formation
Stratigraphic range: Westphalian
Type	Geological formation
Underlies	Týnec Formation
Location
Region	Pilsen
Country	Czech Republic

Paleobiota

Invertebrates

Color key Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon		Notes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Invertebrates
Genus	Species	Higher taxon	Notes	Images
Necymylacris	N. scudderi	Blattodea	Formerly included within Phylloblatta[2]
Bohemiatupus	B. elegans	Meganeuridae[3]	One of the larger meganisopterans
Bojophlebia	B. prokopi	Palaeoptera	Largest insect by wingspan other than palaeodictyopterans or griffinflies.[4]
Carbotriplura	C. kukalovae	Dicondylia	Sister group to Pterygota, formerly classed as a Bojophlebia nymph.[5]
Pronaidites	P. carbonarius, P. arenivorus	Tubificidae	”P. crenulatus” actually an ichnofossil[6]
Microconchus	M. vorax, M. minimus	Tentaculita	Formerly classed as a spirorbid[6]
Prolimulus	P. woodwardi	Bellinuridae	Likely clades with other genera like Alanops.[7]
Eolycosa[8]	E. lorenzii	Mesothelae incertae sedis[9]	Spider affinities doubted in a 1996 paper[10]
Geralycosa[8]	G. fricii	Arthromygalidae	May not be a spider[10]
Rakovnicia[8]	R. antiqua	Tetrapulmonata incertae sedis[9]
Pyritaranea	P. tubifera	Araneae incertae sedis[9]	Synonymised with Eopholcus
Nyranytarbus[9][11]	N. hofmanni, N. longipes	Trigonotarbida	Formerly placed within Hemiphrynus
Doubravatarbus	D. krafti	Aphantomartidae	Likely arboreal, due to being found on a liana-like plant and having relatively gracile legs.[12]
Tynecotarbus	T. tichaveki	Trigonotarbida	Collected alongside two more fragmentary indeterminate fossils[13]
Palaranea	P. borassifolia	Araneae incertae sedis	Known from a different locality to other Kladno fauna[9]
Anthracomartus[9]	A. carcinoides, A. bohemica, A. elegans	Trigonotarbida		A. trilobitus fossil (from North America)
Isobuthus	I. nyranensis[14]	Scorpiones
Prothelyphonus	P. bohemicus[15]	Uropygi	Synonymous with Geralinura[16]	Geralinura fossil (from North America)
Dinopilio	D. gigas	Arachnida incertae sedis[9]	Very large fossil, hence the name
Geratarbus[8]	G. bohemicus	Phalangiotarbida
Orthotarbus[8]	O. nyranensis	Phalangiotarbida
Pleurolycosa	P. prolifera	Arthropoda incertae sedis[9]
Nyranius	N. costulatus, N. tabulatus	Xyloiuloidea	Formerly within Xylobius[17]	N. costulatus fossil
Pleurojulus	P. biornatus, P. levis	Pleurojulida	Also known from Mazon Creek[18]	P. biornatus fossil
Isojulus	I. constans	Pleurojulida	Synonymised with two Pleurojulus species[18]
Purkynia	P. lata	Zosterogrammida	Very poorly preserved[19]	Illustration of a P. lata fossil

Vertebrates

Vertebrates
Genus	Species	Higher taxon	Notes	Images
Archaeothyris	A. sp	Ophiacodontidae	Only known from a jaw fragment[20]	Archaeothyris reconstruction
Brouffia	B. orientalis	Eureptilia	Formerly placed within Gephyrostegus[21]	Brouffia skull reconstruction
Coelostegus	C. prothales[21]	Eureptilia	Recovered as the basalmost eureptile[22]	Coelostegus skull reconstruction
Gephyrostegus	G. bohemicus[23]	Gephyrostegidae	Formerly included another species, G. watsoni[24]	Gephyrostegus reconstruction
Solenodonsaurus	S. janenschi	Reptiliomorpha?	Resembles diadectomorphs, yet likely not closely related to amniotes[25]	Solenodonsaurus reconstruction
Urocordylus	U. angularis[26]	Urocordylinae	Also known from Ireland	Urocordylus reconstruction
Sauropleura	S. scalaris	Sauropleurinae	Formerly included in Urocordylus[27]	Sauropleura life reconstruction
Scincosaurus	S. crassus	Scincosauridae	Formerly synonymised with Keraterpeton,[26] related to diplocaulids[28]	Scincosaurus reconstruction
Keraterpeton	K. galvani[26]	Diplocaulidae	Bore an unusually long tail	Keraterpeton reconstruction
Hyloplesion	H. longicostatum	”Microsauria”	Synonymous with “Orthocosta” and “Seeleya”[26][29]	Hyloplesion reconstruction
Microbrachis	M. pelikani	Recumbirostra	Synonymised with three other species in the same genus[29]	Microbrachis reconstruction
Ricnodon	R. copei	Recumbirostra?	May be related to Saxonerpeton[29]
Crinodon	C. limnophyes	Tuditanidae	Name derives from an anagram of Ricnodon[29]
Sparodus	S. validus	Gymnarthridae?	Resembles Hylerpeton[29]
Diplovertebron	D. punctatum	Embolomeri	Synonymous with “Nummulosaurus”[26]	Diplovertebron reconstruction
Capetus	C. palustris[26]	Temnospondyli	One of the most basal temnospondyls[30]	Capetus reconstruction
Cochleosaurus	C. bohemicus	Cochleosauridae	Adult specimens only found in 2003[31]	Cochleosaurus life restoration
Mordex	M. calliprepes[26]	Trematopidae	Unclear position within the family[32]
Mattauschia	M. laticeps	Trematopidae	Formerly included in Mordex,[32] includes “Potamochoston” and several “Limnerpeton” species[33]
Nyranerpeton[34]	N. amilneri	Micromelerpetontidae	Also known from the Montceau-les-Mines lagerstätte[35]
Limnogyrinus	L. elegans	Micromelerpetontidae	Formerly included within “Limnerpeton”[33]
Oestocephalus[36]	O. granulosum, O. nanum	Aistopoda	Formerly included within Ophiderpeton[26]	Oestocephalus reconstruction
Phlegethontia	P. longissima, “Dolichosoma scutiferum”	Aistopoda	Formerly included within the nomen nudum “Dolichosoma”[37][26]	Phlegethontia reconstruction
Baphetes	B. orientalis	Baphetidae	Formerly known as “Loxomma bohemicum”[38]
Stambergichthys	S. macrodens	Actinopterygii	Likely a high-level predator in its ecosystem[1]
Sceletophorus	S. biserialis, S. verrucosus	Trissolepididae[39]
Pyritocephalus	P. sculptus	Haplolepiformes[39]	Synonymous with other Pyritocephalus species

Plants

Non-seed plants

Non-seed plants
Genus	Species	Higher taxon	Notes	Images
Kladnostrobus	K. clealii, K. psendae	Lycopsida	Within the monotypic family Kladnostrobaceae due to differing sporangia structure[40]
Lepidophloios	L. acerosus	Lepidodendrales	Mistakenly spelled “Lepidofloios” in the paper reporting its presence[41]	Lepidophloios fossil (from Scotland)
Lepidodendron	L. acutum, L. simile,[41] L. aculeatum, L. ophiurum,[42] L. lycopodioides[43]	Lepidodendrales	One of the most abundant Carboniferous plants	Lepidodendron restoration
Flemingites	F. sp	Lepidodendrales	Cone taxon[41]
Lepidocarpon	L. majus[42]	Lepidodendrales	Sporangium taxon[41]
Sigillaria	S. rugosa, S. diploderma	Lepidodendrales	S. diploderma only known from a small fragment[41]	Sigillaria reconstruction
Syringodendron	S. sp	Lepidodendrales	Stem taxon[41]
Omphalophloios	O. feistmantelli[42]	Isoetales	Questionable specimen known from a fragment of a fertile axis[41]	Omphalophloios fossil
Stigmaria	S. ficoides[41]	Lepidodendrales	Form taxon representing roots	Stigmaria fossil
Calamites	C. carinatus, C. distachya?, C. cistii?[42]	Calamitaceae	Second species is closest to C. distachya, but poor preservation means its affinity is unclear[41]	Reconstruction of the entire Calamites tree
Annularia	A. radiata[41]	Calamitaceae	Leaf taxon of Calamites stems	A. stellata fossil
Asterophylites	A. longifolius, A. equisetiformis, A. grandis[43]	Calamitaceae	One specimen preserves the tip of a branch with dense leaves[41]
Sphenophyllum	S. cuneifolium, S. pseudoaquense, S. ?majus,[43] S. priveticense[44]	Sphenophyllales	Very common, but often poorly preserved[41]	Sphenophyllum reconstruction
Bowmanites	B. priveticensis, B. myriophyllus, B. pseudoaquensis, B. brasensis[45]	Sphenophyllales	Cones of Sphenophyllum[44]
Lobatopteris	L. aspidioides	Marattiales	Only one small fragment preserved[41]	L. corsinii fossil (from Spain)
Corynepteris	C. sternbergii, C. angustissima, C. essinghii[42]	Zygopteridales	C. sternbergii may be synonymous with C. angustissima[41]	C. angustissima fossil
Senftenbergia	S. plumosa	Tedeleaceae (Filicales)	Common elsewhere, yet rare in Kladno[41]	S. dentata fossil (from Spain)
Zeilleria	Z. delicatula	Filicales incertae sedis[41]	Shares its name with a Jurassic brachiopod
Renaultia	R. crepinii	Filicales incertae sedis	Only known from a fragment[41]	R. lebachensis fossil (from Spain)
Sphenopteris	S. rotundifolia, S. mixta, S. delicatula, S. acutiloba, S. ujezdensis, S. pulcherrima,[42] S. cirrhifolia,[46] S. spinosa[43]	Filicales incertae sedis (rotundifolia), Lyginopteridales (others)	Includes seed plants and a fern[41]	S. rotundiloba fossil
Aphlebia	A. sp	Indeterminate	Artificial taxon; may belong to another species preserved.[41]	A. crispa fossil (accompanied by Polymorphopteris)
Noeggerathia	N. foliosa, N. intermedia[47]	Noeggerathiales	Most common plant in the sampled fossils[42]	Drawings of N. expansa fossils
Archaeonoeggerathia	A. schatzlarensis[47]	Noeggerathiales
Discosoropteris	D. zlatkokvacekii, D. chlupatum	Filicales incertae sedis	Likely a short, upright plant[48]
Spencerites	S. leismanii, S. havlenae[43]	Lycopsida incertae sedis	A member of the “sub-arborescent” lycophytes[49][46]
Dendraena	D. pinnatilobata	Anachoropteridaceae (Filicales)	Likely grew near river margins,[50] a very common plant[46]
Kidstonia	K. heracleensis	Filicales	Likely a small understorey fern[46]
Desmopteris	D. alethopteroides, D. longifolia[43]	Filicales	Likely bore procumbent (growing along the ground) rhizomes[46]
Selaginella	S. gutbieri,[43] S. labutae[51]	Selaginellaceae	Extant genus	The extant S. selaginoides (from Estonia)
Palaeostachya	P. distachya, P. gracilima	Calamitaceae	Cone taxon,[43] although with a complete plant known from the early Permian of China[52]	P. sp fossil (from Mazon Creek)
Pecopteris	P. aspidioides, P. tuffitica[53]	Psaroniaceae	Likely a mid-sized tree fern[43]	P. sp fossil (from Mazon Creek)
Sonapteris	S. pilsensis, S. barthelii, S. bekii[43]	Botryopteridaceae (Filicales)	Possibly a liana-like plant?[53]
Oligocarpia	O. lindsaeoides	Gleicheniales?	Probably a liana-like plant[43]	O. gutbieri fossil (from Germany)
Adiantites	A. sp	Progymnospermopsida	Likely a low-lying plant[43]
Echinosporangites	E. libertite	Filicopsida?	Sporangium taxon, parent plant unknown[54]
Polysporia	P. drabekii, P. rothwelii, P. radvanicensis, P. robusta	Isoetales	A sub-arborescent lycophyte[55]
Huttonia	H. spicata	Equisetales	Cone taxon[56]
Thomasites	T. serrata, T. elongatus	Lycopsida	T. elongatus formerly placed in Selaginellites[57]

Seed plants

Seed plants
Genus	Species	Higher taxon	Notes	Images
Alethopteris	A. lonchitica, A. distantinervosa, A. pseudograndinioides, A. kettneri, A. serlii	Medullosales	A. pseudograndinioides is a replacement name for the illegitimate “A. grandinioides”[58]	A. lonchitica fossil
Neuromariopteris	N. scandens	Callistophytales	Likely a creeping plant growing on ridge slopes[59]
Eusphenopteris	E. nummularia	Lyginopteridales	While a large (>1 m) frond was found, it could not be transported from the site.[41]	E. striata fossil
Lonchopteris	L. rugosa	Medullosales	Possible record of L. chandesrisii (although based on a poorly preserved specimen) also known[58]	L. rugosa fossil
Macroneuropteris	M. scheuchzeri	Medullosales	Formerly placed within Neuropteris[58]	M. scheuchzeri frond reconstruction
Neuropteris	N. plicata	Medullosales	Placement of this species is unclear in relation to N. ovata.[58]	N. ovata fossil (from Kansas)
Odontopteris	O. reichiana	Medullosales	May be synonymous with O. brardii[58]	O. genuina fossil
Callipteridium	C. rubescens, C. armasii	Medullosales	May be synonymous with C. jongmansii[58]	C. gigas fossil
Laveineopteris	L. loshii, L. tenuifolia, L. bohemica, L. lubnensis, L. hollandica	Medullosales	L. loshii is abundant elsewhere, while some other species are endemic to the Kladno Formation[58]	L. rarinervis fossil
Palaeoweichselia	P. defrancei	Medullosales	Described from Kladno in a 2020 overview paper despite specimens being already known prior[58]
Linopteris	L. neuropteroides, L. obliqua, L. weigelii	Medullosales	L. neuropteroides is split into two forms, neuropteroides and minor.[58]	L. subbrongniarti fossil
Paripteris	P. linguaefolia	Medullosales	Included alleged records of P. gigantea[58]
Havlenaea[60]	H. coriacea, H. stradonitzensis	Medullosales	Formerly placed within Neuropteris[58]
Mixoneura	M. muensterifolia	Medullosales	Too poorly preserved to place within a new genus, yet does not belong to Mixoneura either[58]
Rhacopteris	R. elegans	Pteridospermatophyta	Only known from one isolated pinnule (leaf cluster)[42]	R. elegans fossil (from Italy)
Cordaites	C. theodorii, C. wartmannii, C. kladnoensis,[61] C. borassifolius[43]	Cordaitales	C. theodorii is only known from this formation[42]	C. foliatus fossil (from France)
Palmatopteris	P. furcata	Lyginopteridales	Probably a liana-like plant[43]	P. furcata fossil (from Germany)
Mariopteris	M. muricata	Lyginopteridales	Likely a vining plant, as its fossils are all associated with Lepidodendron.[43]	M. muricata fossil (from Spain)
Fortopteris	F. radnicensis	Medullosales	Not given a type specimen in its original description within Mariopteris.[62]
Rhodeites	R. gutbieri	Lyginopteridales	Likely a canopy-dwelling climber[63]