Geo log i cal Quar terly, 2016, 60 (3): 737–745 DOI: http://dx.doi.org/10.7306/gq.1301

The first “osteolepiform” tetrapodomorph (Sarcopterygii) from the Pa leo zoic se quences of the Moravian Karst (Czech Re pub lic)

Hedvika POUKAROVÁ1, 2, * and Tomáš WEINER1, 2

1 Masaryk Uni ver sity, Depart ment of Geolog i cal Sci ences, Fac ulty of Sci ence, Kotláøská 2, 611 37 Brno, Czech Re pub lic 2 In sti tute of Ge ol ogy of the Czech Acad emy of Sci ences, v.v.i., Rozvojová 269, 165 00 Prague 6, Czech Re pub lic

Poukarová, H., Weiner, T., 2016. The first “osteolepiform” tetrapodomorph (Sarcopterygii) from the Pa leo zoic se quences of the Moravian Karst (Czech Re pub lic). Geo log i cal Quar terly, 60 (3): 737–745, doi: 10.7306/gq.1301

The first tetrapodomorph spec i men from the Pa leo zoic se quences of the Moravian Karst (Moravo-Silesian Ba sin, Bohe mian Mas sif, Czech Re pub lic) is de scribed. The well-pre served, cosmine-cov ered lat eral extrascapular bone co mes from the Up- per De vo nian (Famennian) hemipelagic Køtiny Lime stone of the Líšeò For ma tion. The af fin ity to the “osteolepiforms” is in- ferred from the cosmine his tol ogy and mor pho log i cal fea tures of the bone. As sign ment to the Megalichthyiformes seems most prob a ble. The as so ci ated fauna, e.g., clymenids, orthocone nautiloids, thin shelled bivalves , trilobites and cri noids, clearly dem on strate a marine envi ron ment .

Key words: Tetrapodomorpha, “Osteolepiformes”, cosmine, Famennian, Moravo-Silesian Ba sin, Moravian Karst.

INTRODUCTION To date, the De vo nian and Lower Car bon if er ous se- quences of the Moravian Karst (Moravo-Silesian Ba sin, Bo he - mian Mas sif, Czech Re pub lic; see Figs. 1A, B and 2) have pro - Sarcopterygian fish are es pe cially in ter est ing be cause of vided poor ev i dence of sarcopterygian fish com pris ing their close re la tion to the tetra pods (Rosen et al., 1981; onychodontiforms only (see Ginter, 1991; Smutná, 1994, 1996; Thomson, 1993; Cloutier and Ahlberg, 1996; Ahlberg and Kumpan, 2013). Johanson, 1998; Long and Gordon, 2004; Clack, 2006). The Re cently, a cosmine-cov ered der mal bone was ob tained integumentary skel e ton of the sarcopterygians is from a small aban doned quarry sit u ated in the south ern part of plesiomorphically char ac ter ized by cosmine (e.g., Sire et al., the Moravian Karst near the road con nect ing Brno-Líšeò and 2009). This unique tis sue com plex con sists of a ca nal sys tem Ochoz at Brno (Fig. 1C). This quarry might cor re spond to a fos- en closed in an enamel/enameloid-coated sin gle layer of sil site de scribed by Rzehak (1910). A sec tion about two metres odontodes (Gross, 1956; Thomson, 1975, 1977; Meinke, 1984; thick is com posed of the Famennian hemipelagic Køtiny Lime - Zhu et al., 2006) and an up per most part of spongy bone stone of the Líšeò For ma tion (Weiner and Kalvoda, 2013). This (Thomson, 1975, 1977; Borgen, 1992). Flask-shaped pore-ca - strongly con densed suc ces sion (Weiner and Kalvoda, 2016) nals, which open to the sur face by min ute pores, are con nected cor re sponds to the Hostìnice fa cies de vel op ment (Rez et al., by mesh-ca nals to each other and by cross-ca nals to the pulp 2011). The bone co mes from a darker grey bioclastic lime stone cav i ties (Gross, 1956; Thomson, 1975, 1977; Meinke, 1984). (Fig. 3) de vel oped closely above the black lime stone lenses of The ca nal sys tem is con tin u ous with the vas cu lar ca nals of the the Lower Annulata Event (Up per Palmatolepis rugosa un der ly ing spongiosa (Thomson, 1977; Borgen, 1992) and is trachytera cono dont Zone, see Weiner and Kalvoda, 2016). sup posed to pos sess vas cu lar func tions in volved in the de po si - The pres ence of Polygnathus styriacus Hinde, 1900 in the grey tion of dentine and enamel/enameloid (Bemis and Northcutt, bioclastic lime stone (microfacies F sensu Weiner and Kalvoda, 1992; Borgen, 1992; Zhu et al., 2006). The cosmine cover in 2016) might in di cate the base of the Lower Palmatolepis crown sarcopterygians is sig nif i cant for its uniphase de po si tion perlobata postera cono dont Zone (e.g., Hartenfels, 2011, (Zhu et al., 2006, 2010). Sea sonal re sorp tion and redeposition Weiner and Kalvoda, 2016). Cono donts in di cate the of this spe cial tis sue com plex is sup posed to al low growth of the palmatolepid-polygnathid biofacies (Weiner, 2013; Weiner and an i mal (e.g., Westoll, 1936; rvig, 1969; Thomson, 1975, 1977; Kalvoda, 2016), which is gen er ally sup posed to oc cupy the up - Borgen, 1989, 1992; Fox et al., 1995). per to mid dle part of a ba sin slope en vi ron ment (e.g., Kalvoda et al., 1999: 144; Fig. 3). The microfacies cor re spond ing to wackestone to packstone was in ter preted as de pos ited above or slightly be low storm wave base (microfacies F sensu Weiner * Corresponding author, e-mail: [email protected] and Kalvoda, 2016). The layer con tains a rich fauna in clud ing ammonoids, orthocone nautiloids, thin-shelled bi valves, Received: February 1, 2016; accepted: April 7, 2016; first published ostracods, cri noids and trilobites (Weiner and Kalvoda, 2013, online: June 28, 2016 738 Hedvika Poukarová and Tomáš Weiner

Fig. 1A – position of the area studied in the Czech Republic and the Bohemian Massif; B – simplified map of the Upper Devonian carbonate platform of the Moravo-Silesian Basin in the subsurface and at outcrop, modified after Bábek et al. (2007); C – geographic position of the locality studied (marked by a black arrow)

Fig. 2. Stratigraphic scheme of the southern part of the Moravian Karst (after Kalvoda, 1996 in Rez et al., 2011)

Lst. – limestone, Fa. – Famennian, Tn. – Tournaisian The first “osteolepiform” tetrapodomorph (Sarcopterygii) from the Paleozoic sequences of the Moravian Karst (Czech Republic) 739

and drowned dur ing the Frasnian to Late Famennian in ter val, and this was con nected to the on set of the halfgraben sed i men - ta tion of the Líšeò For ma tion (Kalvoda and Melichar, 1999; Bábek et al., 2007; Kalvoda et al., 2008). In the Famennian in ter - val, two fa cies de vel opments of the Líšeò For ma tion may be roughly distin guished in the south ern part of the Moravian Karst (Fig. 2): the Horákov fa cies de vel op ment is char ac ter ized by rel - atively thick sequences of calciturbidites (Hády–Øíèka Lime - stone), which were de pos ited in deeper parts of the slope and basin, whereas the Hostìnice fa cies de vel op ment com prises con densed suc ces sions of nod u lar hemipelagites (Køtiny Lime - stone), rep re sent ing a shal lower en vi ron ment on the by pass slope (Kalvoda et al., 1996; Rez et al., 2011).

MATERIAL AND METHODS

The cosmine-cov ered der mal bone is pre served together with in deter min able fish remains in the same sam ple. The bone is well-pre served but the cosmine sur face es pecially is af fected by nu mer ous microcracks. The bone, bro ken in the field into two parts, was me chan i - cally prepared us ing a pneu matic en grav ing pen and sub se - quently chem i cally treated with di lute ace tic acid. The small area at the crack di vid ing the bone into two parts was polished. De tailed pho to graphs were taken us ing a Nikon SMZ 1500 bin - oc u lar mi cro scope with a Nikon DXM dig i tal cam era and NIS-El e ments soft ware. In some cases, the sam ple was sub - merged in water for better re sults, and in oth ers, a coat ing of am mo nium chlo ride was ap plied. De tailed mea sure ments were taken us ing JMicroVision soft ware (Nicolas Roduit, Switzer - land). A coat ing of gold was ap plied to a small area of the cosmine sur face be fore us ing scan ning elec tron mi cros copy.

SYSTEMATIC PART

Fig. 3. Lithostratigraphy and biostratigraphy of the section Osteichthyes Huxley, 1880 The strati graphic po si tion of the “osteolepiform” dermal bone is Sarcopterygii Romer, 1955 marked by an ar row; Cl – calcilutite, Ca – calcarenite, f – fine Rhipidistia Cope, 1887 sensu Cloutier and Ahlberg, 1996 grained, m – me dium grained; mod i fied af ter Weiner and Kalvoda Tetrapodomorpha Ahlberg, 1991 (2016) “Osteolepiformes” indet. (Figs. 4A, 5 and 6) 2016). The in sol u ble re sid uum also yielded fish re mains be - long ing mostly to chondrichthyans and “palaeoniscids”. The cosmine-cov ered bone is housed in the Col lec tions of Material. – One cosmine-cov ered right lat eral the Czech Geo log i cal Sur vey, Prague, in ven tory num bers extrascapular from the Køtiny Lime stone of the Líšeò Forma - HP108a and HP108b. tion; Famennian (the span of the Lower postera to Lower expansa cono dont Zones); “Ochoz sec tion” (sensu Weiner and Kalvoda, 2016) be tween Brno and Ochoz at Brno. GEOLOGICAL SETTING Description. – The right lat eral extrascapular has a subtriangular out line and is about 2 cm in size. Three slightly The Mid dle De vo nian to Lower Car bon if er ous se quences of con cave over lapped ar eas are de vel oped at its ante rior mar gin. the Moravian Karst were de pos ited in the Moravo-Silesian Basin The main sen sory canal passes through the middle of them. at the south ern mar gin of Laurussia (e.g., Kalvoda et al., 2002, Other mar gins lack over lapped ar eas. The cor ner be tween the 2008). In a re gional geo log i cal con text, these se quences be long slightly con vex lat eral to posterolateral mar gin and the con cave to the Moravo-Silesian zone of the Bo hemian mas sif (Fin ger et posteromedial mar gin is rounded, with out a notch. The me dial al., 2000) and cor re spond to the Rhenohercynian zone of the mar gin is straight to very slightly concave (Fig. 4A). Cen tral Eu ro pean Variscides (Hladil et al., 1999). The Moravian The exter nal sur face of the bone bears nu mer ous open ings Karst fa cies de vel op ment rep re sents one of sev eral pre-flysch for tubes lead ing to the main and supratemporal commissural fa cies do mains in the Moravo-Silesian Ba sin (e.g., Zukalová and sen sory canals. The even dis tri bu tion of these open ings does Chlupáè, 1982; Hladil, 1992; Kalvoda et al., 2002, 2008). The not al low one to trace the course of the sen sory canals from the car bon ate plat form (Macocha For ma tion) of the Moravian Karst bone’s ex ter nal sur face (Fig. 4A). Nev er the less, these sen sory de vel op ment be gan to be pro gres sively tec toni cally dis in te grated ca nals with a di am eter of about 0.6 mm are vis i ble at a crack 740 Hedvika Poukarová and Tomáš Weiner go ing through the bone. A short pit-line with fo ram ina reach ing pore-ca nals. The shape of the pore-ca nals is rem i nis cent of an ap prox i mately 57 mm in di am e ter is de vel oped (Figs. 4A and oast-house chim ney. These ca nals reach a height of around 5F). The cosmine cover is devoid of Westoll lines. 137 mm (Fig. 5A, B, E), and their sur face open ings are shaped A layer approx i mately 0.4 mm thick of lamellar bone is over - like wide fun nels. The di am eter of these open ings reaches on lain by a layer about 1.3 mm thick of spongy bone. Buried av er age 16 mm at the top (based on 70 mea sure ments), but it odontodes are ab sent. The enamel/enameloid coated dentine can be re duced to 5 mm at the base of the fun nel. The dis tance layer is about 0.14 mm thick and en closes the pore-ca nal net - betw een open ings is about 153 mm (140 mea sure ments were work (Fig. 5E). The enamel/enameloid does not en ter the taken). The mesh-canals reach their great est di am eter at their con tact with the pore-ca nals, but their thick ness of ten sig nif i - cantly decreases in their cen tral part (Fig. 5B). The lower mesh-canals are miss ing. Bun dles of roughly radi ally arranged dentine tu bules are de vel oped be tween the flask-shaped pore-ca nals. The or ange col our of the dentine tu bules might be caused by iron oxides/ oxyhydroxides. The cosmine cover is brown, and its trans par ency enables one to ob serve the dis tri - bution of the pore-ca nals and dentine tubules, es pecially when the spec i men is sub merged in water (Fig. 5A, D, F). Us ing an opti cal mi cro scope, a very fine pat tern at the cosmine sur face is ap par ent (Fig. 5C); nev er the less, this pat tern was not clearly ob served us ing the scan ning elec tron mi cro scope (see Fig. 6 show ing also the de tails of the sen sory tube opening). The par - tial ab sence of cosmine is prob ably caused by dam age rather than by resorp tion.

DISCUSSION

The cosmine his tol ogy in vari ous groups of sarcopterygian fish has been dis cussed e.g., by Gross (1956), Rosen et al. (1981), Meinke (1984), Schultze (1986), Chang and Smith (1992), Sire et al. (2009) and Zhu et al. (2010). The Moravian spec i men pos sesses the com bi na tion of cosmine fea tures in - clud ing the ab sence of Westoll lines and bur ied odontodes, the enamel/enameloid not ex tend ing into the pore-ca nals, and the shape of the pore- and mesh-ca nals, which matches the con di - tions com mon in “osteolepiforms” (see Rosen et al., 1981; Meinke, 1984; Chang and Smith, 1992). The spec i men also lacks the lower mesh-ca nals known in the Mid dle De vo nian “osteolepiforms” such as Osteolepis Agassiz, 1835 and many early dip no ans (e.g., Dipterus Sedg wick and Murchi son, 1829), which is usual in youn ger forms of “osteolepiform” tetrapodomorphs, e.g., Megalichthys Agassiz, 1935 or Ectosteorhachis Cope, 1880 (see Thomson, 1977). A sim ilar shape of pore- and mesh-canals was re corded es pecially in Cladarosymblema Fox et al., 1995 and Megalichthys Agassiz, 1935 (see Fox et al., 1995; Gross, 1956). The di am eter of the pore-ca nal open ings in the Moravian spec i men is close to the av er age of 10 mm re corded in the “osteolepiforms” by Thomson (1977). The height of the pore-ca nals and their spac ing also

Fig. 4A – right lateral extrascapular of “osteolepiform” tetrapodomorph from the Moravian Karst; the speci men was coated with am mo nium chlo ride; B – po si tion of right lateral extrascapular (ar rowed) on the skull and exoskeletal shoul der gir dle re con struc tion of an “osteolepiform” fish (Osteolepis macrolepidotus Agassiz, 1835), modi fied after Jarvik (1948) Ana tom i cal abbre vi a tions: Et – extratemporal, Exsc.l. – lat eral extrascapular, Exsc.m. – me dian extrascapular, msc – main sen - sory canal, od.Et – area on lat eral extrascapular over lapped by extratemporal, od.Ppa – area on lat eral extrascapular over lapped by postparietal, od.Ta – area on lateral extrascapular over lapped by tabu lar, Op – operculum, pl – pit-line, Pop – preoperculum, Ppa – postparietal, Pt – posttemporal, Sq – squamosal, stcc – supratemporal commissural ca nal, sto – sen sory tube opening, Ta – tab u lar The first “osteolepiform” tetrapodomorph (Sarcopterygii) from the Paleozoic sequences of the Moravian Karst (Czech Republic) 741

Fig. 5A – detail of the cosmine surface observed at an acute angle; B – detail of polished cross-section through the cosmine cover; C – detail of the cosmine surface showing very fine granulation; D – detail of the cosmine surface; E – polished cross-section through the dermal bone with the cosmine cover; F – detailed view of the pit-line from the cosmine surface

dl – dentine layer, e – enamel/enameloid, lb – lamellar bone, mc – mesh-canal, msc – main sensory canal, pc – pore-canal, pco – pore-canal opening, pl – pit-line, sb – spongy bone, sto – sensory tube opening, vc – vascular canal 742 Hedvika Poukarová and Tomáš Weiner

oped posteromedial part of this bone (Jarvik, 1948). On the other hand, a spec i men of Thursius moy-thomasi Jarvik, 1948 lack ing an in de pend ently de vel oped me dian extrascapular is also known (Jarvik, 1948). The me dial mar gin of the Moravian spec i men lacks an over lapped area sug gest ing that the lat eral extrascapulars must have over lapped the me dian extrasca - pular. This con di tion is usual in “osteolepiform” tetrapodo - morphs (Jarvik, 1980). In porolepiforms, lat eral extrascapulars are over lapped by the median one (Jarvik, 1980), and the sit ua - tion in dip noans is un clear be cause of the vari abil ity in the num - ber of extrascapulars (Miles, 1977). The vari abil ity in shape of “osteolepiform” lat eral extrascapulars was clearly dem on - strated by Säve-Söderbergh (1933) in Mid dle De vo nian spec i - mens from Scot land. The pit-line can be pres ent or ab sent in these bones, and this con di tion can also vary even on both sides of one an i mal (Jarvik, 1948). Nu mer ous and evenly dis- trib uted open ings of the main and supratemporal commissural ca nals in the Moravian spec i men rep re sent a fea ture known es- Fig. 6. Scanning electron micrograph of the cosmine surface pecially in younger forms of “osteolepiform” tetrapodomorphs with the sensory tube opening such as Megalichthys Agassiz, 1835 or Eusthenopteron Whiteaves, 1881 from the Car bon if er ous and Perm ian suc ces - sions (Jarvik, 1948). In Mid dle De vo nian forms such as cor re spond to the 150–250 mm mea sured in “osteolepiforms” Thursius macrolepidotus (Sedg wick and Murchi son, 1829) or (Thomson, 1977). All these val ues are con sid er ably higher in Osteolepis macrolepidotus Agassiz, 1835, the branch ing of porolepiforms and dip noans (Thomson, 1977). The pore-ca - tubes ris ing from these ca nals is weak, and the tu bule open ings nals of Porolepis Wood ward, 1891 are ap prox i mately 250 mm are of ten ar ranged in a sin gle row (Jarvik, 1948). An in ter me di - high, their openings reach 60–70 mm in di am eter, and the dis - ate con di tion was re corded e.g., in the lower most Up per De vo - tance be tween them is about 300 mm (Thomson, 1977). In the nian Latvius grewingki (Gross, 1933) with tu bule open ings ar - dip no an ge nus Ganorhynchus Traquair, 1873 the pore-ca nals ranged in dou ble to tri ple rows (Jarvik, 1948). are ap prox i mately 650 mm high, their open ings reach 400 mm in “Osteolepiform” tetrapodomorphs have been ob tained from di am eter, and the dis tance be tween them is about 500 mm fresh water and brack ish water as well as marine strata, and the (Thomson, 1977). Bemis and Northcutt (1992) re corded the di - pos si bil ity of their anadromy has been dis cussed (see am e ter of dip no an pore-ca nal open ings as fol lows: 83 mm in Thomson, 1969, 1980). Ac cord ing to Long et al. (1997), the Sunwapta grandiceps Thomson, 1967; 20–160 mm with an av - sen sory pore groups, which are dis tinctly devel oped mainly in er age of 64 mm in Dipterus valenciennesi Sedg wick and spec i mens from freshw ater de pos its and ab sent in the ma rine Murchi son, 1829 and the open ings as small as 45 mm in Gogonasus andrewsae Long, 1985 could in di cate sa lin ity. The Chirodipterus aus tra lis Miles, 1977. spec i men from the Moravian Karst derives from a rel atively Few com ments have been made on the fine cosmine sur- deeper water en vi ron ment with clearly marine fauna. Un for tu - face or na men ta tion be tween the pores. Chang and Smith nately, the lat eral extrascapulars seem not to be suit able for di - (1992) re corded thin scal loped ridges of unknow n char ac ter agno sis of the pres ence of sen sory pore groups, so the sug ges - anterolateral to the pores in one spec i men of Diabolepis Chang, tion by Long et al. (1997) can not be con firmed nor dis proved. 1984. A hex ag o nal pat tern re flect ing the ar range ment and size More over, many “osteolepiforms” were ob tained from red bed of ep i the lial cells was observed in the dip noans Chirodipterus fa cies, the in ter pre ta tion of which as regards sa lin ity is still prob - aus tra lis Miles, 1977 (Smith, 1977) and Dipterus sp. (Schultze, lem atic (Laurin and Soler-Gijón, 2010). 1977) us ing a mag ni fi ca tion of 400 to 2000 times. The cosmine The “osteolepiforms” have been sub jected to numer ous sur face of the Moravian spec i men shows a very fine pat tern anal y ses (e.g., Long, 1985a, b; Ahlberg, 1991; Young et al., (Fig. 5C); nev er the less, the scan ning elec tron mi cro scope 1992; Ahlberg and Johanson, 1998; Coates and Fried man, failed to re veal such a hex ag o nal pat tern us ing cor re spond ing 2010; Swartz, 2012; Witzmann and Schoch, 2012; Hol land, mag ni fi ca tions (see Fig. 6). Säve-Söderbergh (1941) and 2013) lead ing to dif fer ent in ter pre ta tions of re la tion ships within Jarvik (1948) described an al ter na tion of dark and light bands in this group. Swartz (2012) used the term “osteolepiform” to en - the dentine layer of sev eral (mainly black) spec i mens of cap su late the grade of tetrapodomorphs, in clud ing the “osteolepiform” tetrapodomorphs, es pecially if they were sub - stem-based Canowindridae, Megalichthyiformes and merged in water or al co hol. A brown speci men from the Tristichopteridae. Moravian Karst does not show these bands. Cosmine cover is com pletely absent in tristichopterids (e.g., Der mal bones of “osteolepiform” tetrapodomorphs show a Snitting, 2008), and canowindrids are sup posed to be a uni form over lap pat tern (Jarvik, 1980). Over lapped ar eas at the Gondwanan group (Young et al., 1992). It is prob able that the an te rior mar gin of the lat eral extrascapular in the Moravian spec i men from the Moravian Karst be longs to the spec i men could have served for ar tic ula tion with the megalichthyiforms sensu Swartz (2012). Young et al. (1992) postparietal, tab ular and prob ably also with the in de pend ently described the shape of lat eral extrascapulars in var i ous de vel oped extratemporal bone (in ferred from the tri par tite an te - “osteolepiform” groups (the char ac ter “C2: equi lat eral shape of rior mar gin, see Säve-Söderbergh, 1933; see Fig. 4). However, extrascapulars” in their cladistic anal y sis). The con di tion of bones of the postparietal shield and extrascapular series can three-sided lat eral extrascapulars al most meet ing in the midline vary in devel opment even on both sides of one indi vid ual in canowindrids and megalichthyids vs. four-sided lat eral (Jarvik, 1948). A com pound char ac ter of the lat eral extrascapulars well sep arated in the an te rior midline in extrascapulars was rec og nized in a spec i men of Osteolepis tristichopterids and “osteolepidids” such as Osteolepis Agassiz, macrolepidotus Agassiz, 1835 with an in depend ently devel - 1835 or Gyroptychius M’Coy, 1848 was later dis cussed by The first “osteolepiform” tetrapodomorph (Sarcopterygii) from the Paleozoic sequences of the Moravian Karst (Czech Republic) 743

Johanson and Ahlberg (1997). These au thors pointed out that and Kladno–Rakovník bas ins, the Klobouky Ho ri zon of the Línì the lat eral extrascapulars of the megalichthyid Cladaro - For ma tion in the Kladno–Rakovník Ba sin and the Perm ian Kalná symblema Fox et al., 1955 are four-sided and not three-sided Ho ri zon of the Proseèné For ma tion in the Krkonoše Piedmont as should be ex pected, and they revealed some con tra dic tion Ba sin (Štamberg and Zajíc, 2008). be tween the de scrip tions and real shapes of these bones in some other genera. Johanson and Ahlberg (1997: p. 49) di - vided the char ac ter into two: “three sided lat eral extrascapulars” CONCLUSIONS and “lat eral extrascapulars al most meet ing in midline” and rec - om mended fur ther con sid er ation. In this re spect, the shape of To date, De vo nian and Car bon if er ous de pos its of the the lat eral extrascapular of the Moravian spec i men is closer to Moravian Karst (Moravo-Silesian Ba sin, Bo he mian Mas sif) have megalichthyids than “osteolepidids”. Nev er the less, it is nec es - yielded poor mate rial of sarcopterygian fish, com pris ing only sary to treat such an as sign ment with cau tion. The dis tance be - onychodontiforms (see Ginter, 1991; Smutná, 1994, 1996; tween lat eral extrascapulars in the Moravian spec i men is not Kumpan, 2013). The newly recorded subtriangular cosmine- cov - known. In ad di tion, these fea tures are gener ally rather prob lem - ered right lat eral extrascapular co mes from the pre dom i nantly atic. Both groups are closely re lated (see Janvier et al., 2007) hemipelagic suc ces sion of the Famennian Køtiny Lime stone of and com prise megalichthyiforms ac cord ing to Swartz (2012). the Líšeò For ma tion. The com bi na tion of cosmine histological The spec i men de scribed in this pa per is the first recorded fea tures in clud ing an ab sence of Westoll lines and bur ied “osteolepiform” tetrapodomorph from the Pa leo zoic se quences odontodes, enamel/enameloid not ex tend ing into the pore-ca - of the Moravian Karst. World wide, other known Famennian nals, pore-ca nals in a shape rem i nis cent of oast-house chim - cosmine-bear ing “osteolepiforms” in clude e.g., Megapomus neys and the diam eter of the pore-canal open ings dem onstrat e Vorobyeva, 1977 and Cryptolepis Vorobyeva, 1975 (Baltica an “osteolepiform” af fin ity. This as sign ment is supported by the Prov ince), Megistolepis Obruchev, 1955 (Si beria Prov ince) and lack of an over lapped area at the me dial bone mar gin. The bone Sterropterygion Thomson, 1972 (Laurentia Province) (see bears nu mer ous evenly distrib uted open ings of the sen sory ca- Lebedev and Zakharenko, 2010). A Famennian or pos si bly a nals as is usual in stratigraphically later forms of “osteolepiform” Tournaisian age is also sup posed for Medoevia lata Lebedev, tetrapodomorphs (Jarvik, 1948), and it lacks the lower mesh-ca - 1995 which is from an un known lo cal ity (Lebedev, 1995). Al- nals pres ent es pecially in some Mid dle De vo nian forms though these genera could also pos ses nu mer ous and evenly (Thomson, 1977). The as sign ment to the “Mega lichthyiformes” dis trib uted open ings of sen sory canals, we avoid as sign ing the sensu Swartz (2012) seems to be most proba ble. Moravian spec i men to any of them be cause the mate rial com - The as so ci ated fauna has a clearly marine char ac ter and prises only one bone, and the lat eral extrascapulars are known con tains e.g., clymenids, orthocone nautiloids, thin-shelled bi - for their shape vari abil ity (see Säve-Söderbergh, 1933). valves, trilobites, cri noids, cono donts, ostracods, chondri - Its posi tion at the Laurussian con ti nen tal mar gin (Kalvoda chthyans and “palaeoniscids”. Com pared to nu mer ous and Bábek, 2010) and its presumed or i gin in a rel atively deeper “osteolepiforms” from red bed se quences, the Moravian spec i - water en vi ron ment on the bypass slope (see Weiner and men orig i nates from the rel a tively deeper-wa ter en vi ron ment of Kalvoda, 2016) sug gest as sign ment of this Moravian the bypass slope (see Weiner and Kalvoda, 2016). “osteolepiform” ac com pa nied by chondrichthyans and “Osteolepiforms” are re corded in the Paleo zoic se quences “palaeoniscids” to the con ti nental mar gin as sem blage of the of the Moravian Karst for the first time. In the Czech Re pub lic, Baltica Prov ince pe riph ery sensu Lebedev et al. (2010). How - the re mains of these tetrapodomorphs have pre vi ously only ever, there is still a lack of detailed data on the fish fauna from the been re ported from the Car bon if er ous and Perm ian limnic bas - Ochoz sec tion or from other Moravian Karst lo cal i ties rep re sent - ins (Štamberg and Zajíc, 2008). ing a sim ilar mid Famennian envi ron ment. From the Czech Re - public, remains of “osteolepiform” fish have been re ported from Ac knowl edge ments. Thanks are due to O. Fatka and limnic Up per Car bon if er ous and Lower Perm ian de pos its (Zajíc, J. Kalvoda for their help ful com ments. We are grate ful to 2000, 2008; Štamberg and Zajíc, 2008). Iso lated re mains of J. Štelcl for prep ara tion of the scan ning elec tron mi cro graphs Megalichthys nitens Fritsch, 1889 were re corded from the Car - and T. Viktorýn for his kind help in the field. Cor dial thanks are bon if er ous Kounov Mem ber of the Slaný For ma tion of the ad dressed to the re view ers J. Clack and M. Ginter for their use - Kladno–Rakovník Basin (Fritsch, 1889, 1893; Romer, 1945; ful sug ges tions. This con tri bu tion was sup ported by the Czech Štamberg and Zajíc, 2008). Re mains as signed to Sci ence Foun da tion pro ject GA14-18183S and In sti tute Re - “Osteolepiformes” indet. are known from the Car bonif er ous search Plan RVO67985831. Kounov Mem ber of the Slaný For ma tion in the Mšeno–Roudnice

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