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, De part ment of Geo log 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, Bo he 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 bi valves, trilobites and cri noids, clearly dem on strate a ma rine en vi ron ment.
Key words: Tetrapodomorpha, “Osteolepiformes”, cosmine, Famennian, Moravo-Silesian Ba sin, Moravian Karst.
INTRODUCTION
Sarcopterygian fish are es pe cially in ter est ing be cause of their close re la tion to the tetra pods (Rosen et al., 1981;
Thomson, 1993; Cloutier and Ahlberg, 1996; Ahlberg and Johanson, 1998; Long and Gordon, 2004; Clack, 2006). The integumentary skel e ton of the sarcopterygians is plesiomorphically char ac ter ized by cosmine (e.g., Sire et al., 2009). This unique tis sue com plex con sists of a ca nal sys tem en closed in an enamel/enameloid-coated sin gle layer of odontodes (Gross, 1956; Thomson, 1975, 1977; Meinke, 1984;
Zhu et al., 2006) and an up per most part of spongy bone (Thomson, 1975, 1977; Borgen, 1992). Flask-shaped pore-ca - nals, which open to the sur face by min ute pores, are con nected by mesh-ca nals to each other and by cross-ca nals to the pulp cav i ties (Gross, 1956; Thomson, 1975, 1977; Meinke, 1984).
The ca nal sys tem is con tin u ous with the vas cu lar ca nals of the un der ly ing spongiosa (Thomson, 1977; Borgen, 1992) and is sup posed to pos sess vas cu lar func tions in volved in the de po si - tion of dentine and enamel/enameloid (Bemis and Northcutt, 1992; Borgen, 1992; Zhu et al., 2006). The cosmine cover in crown sarcopterygians is sig nif i cant for its uniphase de po si tion (Zhu et al., 2006, 2010). Sea sonal re sorp tion and redeposition of this spe cial tis sue com plex is sup posed to al low growth of the an i mal (e.g., Westoll, 1936; rvig, 1969; Thomson, 1975, 1977;
Borgen, 1989, 1992; Fox et al., 1995).
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 - vided poor ev i dence of sarcopterygian fish com pris ing onychodontiforms only (see Ginter, 1991; Smutná, 1994, 1996;
Kumpan, 2013).
Re cently, a cosmine-cov ered der mal bone was ob tained from a small aban doned quarry sit u ated in the south ern part of the Moravian Karst near the road con nect ing Brno-Líšeò and Ochoz at Brno (Fig. 1C). This quarry might cor re spond to a fos - sil site de scribed by Rzehak (1910). A sec tion about two metres thick is com posed of the Famennian hemipelagic Køtiny Lime - stone of the Líšeò For ma tion (Weiner and Kalvoda, 2013). This strongly con densed suc ces sion (Weiner and Kalvoda, 2016) cor re sponds to the Hostìnice fa cies de vel op ment (Rez et al., 2011). The bone co mes from a darker grey bioclastic lime stone (Fig. 3) de vel oped closely above the black lime stone lenses of the Lower Annulata Event (Up per Palmatolepis rugosa trachytera cono dont Zone, see Weiner and Kalvoda, 2016).
The pres ence of Polygnathus styriacus Hinde, 1900 in the grey bioclastic lime stone (microfacies F sensu Weiner and Kalvoda, 2016) might in di cate the base of the Lower Palmatolepis perlobata postera cono dont Zone (e.g., Hartenfels, 2011, Weiner and Kalvoda, 2016). Cono donts in di cate the palmatolepid-polygnathid biofacies (Weiner, 2013; Weiner and Kalvoda, 2016), which is gen er ally sup posed to oc cupy the up - 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 and Kalvoda, 2016). The layer con tains a rich fauna in clud ing ammonoids, orthocone nautiloids, thin-shelled bi valves, ostracods, cri noids and trilobites (Weiner and Kalvoda, 2013,
* Corresponding author, e-mail: h.poukarova@seznam.cz
Received: February 1, 2016; accepted: April 7, 2016; first published online: June 28, 2016
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
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 the Czech Geo log i cal Sur vey, Prague, in ven tory num bers HP108a and HP108b.
GEOLOGICAL SETTING
The Mid dle De vo nian to Lower Car bon if er ous se quences of the Moravian Karst were de pos ited in the Moravo-Silesian Ba sin at the south ern mar gin of Laurussia (e.g., Kalvoda et al., 2002, 2008). In a re gional geo log i cal con text, these se quences be long to the Moravo-Silesian zone of the Bo he mian mas sif (Fin ger et al., 2000) and cor re spond to the Rhenohercynian zone of the Cen tral Eu ro pean Variscides (Hladil et al., 1999). The Moravian Karst fa cies de vel op ment rep re sents one of sev eral pre-flysch fa cies do mains in the Moravo-Silesian Ba sin (e.g., Zukalová and Chlupáè, 1982; Hladil, 1992; Kalvoda et al., 2002, 2008). The car bon ate plat form (Macocha For ma tion) of the Moravian Karst de vel op ment be gan to be pro gres sively tec toni cally dis in te grated
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 op ments of the Líšeò For ma tion may be roughly dis tin 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 - a tively thick se quences of calciturbidites (Hády–Øíèka Lime - stone), which were de pos ited in deeper parts of the slope and ba sin, 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 to gether with in de ter min able fish re mains in the same sam ple. The bone is well-pre served but the cosmine sur face es pe cially is af fected by nu mer ous microcracks.
The bone, bro ken in the field into two parts, was me chan i - cally pre pared 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 pol ished.
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 wa ter 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, Swit zer - 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
Osteichthyes Huxley, 1880 Sarcopterygii Romer, 1955
Rhipidistia Cope, 1887 sensu Cloutier and Ahlberg, 1996 Tetrapodomorpha Ahlberg, 1991
“Osteolepiformes” indet.
(Figs. 4A, 5 and 6)
M a t e r i a l. – One cosmine-cov ered right lat eral extrascapular from the Køtiny Lime stone of the Líšeò For ma - 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.
D e s c r i p t i o n. – The right lat eral extrascapular has a subtriangular out line and is about 2 cm in size. Three slightly con cave over lapped ar eas are de vel oped at its an te rior mar gin.
The main sen sory ca nal passes through the mid dle of them.
Other mar gins lack over lapped ar eas. The cor ner be tween the slightly con vex lat eral to posterolateral mar gin and the con cave posteromedial mar gin is rounded, with out a notch. The me dial mar gin is straight to very slightly con cave (Fig. 4A).
The ex ter nal sur face of the bone bears nu mer ous open ings for tubes lead ing to the main and supratemporal commissural sen sory ca nals. The even dis tri bu tion of these open ings does not al low one to trace the course of the sen sory ca nals from the bone’s ex ter nal sur face (Fig. 4A). Nev er the less, these sen sory ca nals with a di am e ter of about 0.6 mm are vis i ble at a crack Fig. 3. Lithostratigraphy and biostratigraphy of the section
The strati graphic po si tion of the “osteolepiform” der mal bone is marked by an ar row; Cl – calcilutite, Ca – calcarenite, f – fine grained, m – me dium grained; mod i fied af ter Weiner and Kalvoda (2016)
go ing through the bone. A short pit-line with fo ram ina reach ing ap prox i mately 57 mm in di am e ter is de vel oped (Figs. 4A and 5F). The cosmine cover is de void of Westoll lines.
A layer ap prox i mately 0.4 mm thick of lamellar bone is over - lain by a layer about 1.3 mm thick of spongy bone. Bur ied odontodes are ab sent. The enamel/enameloid coated dentine layer is about 0.14 mm thick and en closes the pore-ca nal net - work (Fig. 5E). The enamel/enameloid does not en ter the
pore-ca nals. The shape of the pore-ca nals is rem i nis cent of an oast-house chim ney. These ca nals reach a height of around 137 mm (Fig. 5A, B, E), and their sur face open ings are shaped like wide fun nels. The di am e ter of these open ings reaches on av er age 16 mm at the top (based on 70 mea sure ments), but it can be re duced to 5 mm at the base of the fun nel. The dis tance be tween open ings is about 153 mm (140 mea sure ments were taken). The mesh-ca nals reach their great est di am e ter at their con tact with the pore-ca nals, but their thick ness of ten sig nif i - cantly de creases in their cen tral part (Fig. 5B). The lower mesh-ca nals are miss ing. Bun dles of roughly ra di ally ar ranged 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 ox ides/oxyhydroxides. The cosmine cover is brown, and its trans par ency en ables one to ob serve the dis tri - bu tion of the pore-ca nals and dentine tu bules, es pe cially when the spec i men is sub merged in wa ter (Fig. 5A, D, F). Us ing an op ti 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 open ing). The par - tial ab sence of cosmine is prob a bly caused by dam age rather than by re sorp tion.
DISCUSSION
The cosmine his tol ogy in var i 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 i lar shape of pore- and mesh-ca nals was re corded es pe cially in Cladarosymblema Fox et al., 1995 and Megalichthys Agassiz, 1935 (see Fox et al., 1995; Gross, 1956). The di am e ter 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 lat eral extrascapular of “osteolepiform”
tetrapodomorph from the Moravian Karst; the spec i men was coated with am mo nium chlo ride; B – po si tion of right lat eral extrascapular (ar rowed) on the skull and exoskeletal shoul der gir dle re con struc tion of an “osteolepiform” fish (Osteolepis macrolepidotus Agassiz, 1835), mod i fied af ter Jarvik (1948) An a tom i cal ab bre vi a tions: Et – extratemporal, Exsc.l. – lat eral extrascapular, Exsc.m. – me dian extrascapular, msc – main sen - sory ca nal, 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 lat eral extrascapular over lapped by tab u lar, Op – operculum, pl – pit-line, Pop – preoperculum, Ppa – postparietal, Pt – posttemporal, Sq – squamosal, stcc – supratemporal commissural ca nal, sto – sen sory tube open ing, Ta – tab u lar
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
cor re spond to the 150–250 mm mea sured in “osteolepiforms”
(Thomson, 1977). All these val ues are con sid er ably higher in porolepiforms and dip no ans (Thomson, 1977). The pore-ca - nals of Porolepis Wood ward, 1891 are ap prox i mately 250 mm high, their open ings reach 60–70 mm in di am e ter, and the dis - tance be tween them is about 300 mm (Thomson, 1977). In the dip no an ge nus Ganorhynchus Traquair, 1873 the pore-ca nals are ap prox i mately 650 mm high, their open ings reach 400 mm in di am e ter, and the dis tance be tween them is about 500 mm (Thomson, 1977). Bemis and Northcutt (1992) re corded the di - am e ter of dip no an pore-ca nal open ings as fol lows: 83 mm in Sunwapta grandiceps Thomson, 1967; 20–160 mm with an av - er age of 64 mm in Dipterus valenciennesi Sedg wick and Murchi son, 1829 and the open ings as small as 45 mm in Chirodipterus aus tra lis Miles, 1977.
Few com ments have been made on the fine cosmine sur - face or na men ta tion be tween the pores. Chang and Smith (1992) re corded thin scal loped ridges of un known char ac ter anterolateral to the pores in one spec i men of Diabolepis Chang, 1984. A hex ag o nal pat tern re flect ing the ar range ment and size of ep i the lial cells was ob served in the dip no ans Chirodipterus aus tra lis Miles, 1977 (Smith, 1977) and Dipterus sp. (Schultze, 1977) us ing a mag ni fi ca tion of 400 to 2000 times. The cosmine sur face of the Moravian spec i men shows a very fine pat tern (Fig. 5C); nev er the less, the scan ning elec tron mi cro scope failed to re veal such a hex ag o nal pat tern us ing cor re spond ing mag ni fi ca tions (see Fig. 6). Säve-Söderbergh (1941) and Jarvik (1948) de scribed an al ter na tion of dark and light bands in the dentine layer of sev eral (mainly black) spec i mens of
“osteolepiform” tetrapodomorphs, es pe cially if they were sub - merged in wa ter or al co hol. A brown spec i men from the Moravian Karst does not show these bands.
Der mal bones of “osteolepiform” tetrapodomorphs show a uni form over lap pat tern (Jarvik, 1980). Over lapped ar eas at the an te rior mar gin of the lat eral extrascapular in the Moravian spec i men could have served for ar tic u la tion with the postparietal, tab u lar and prob a bly also with the in de pend ently de vel oped extratemporal bone (in ferred from the tri par tite an te - rior mar gin, see Säve-Söderbergh, 1933; see Fig. 4). How ever, bones of the postparietal shield and extrascapular se ries can vary in de vel op ment even on both sides of one in di vid ual (Jarvik, 1948). A com pound char ac ter of the lat eral extrascapulars was rec og nized in a spec i men of Osteolepis macrolepidotus Agassiz, 1835 with an in de pend ently de vel -
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 me dian one (Jarvik, 1980), and the sit u a - tion in dip no ans 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 - pe cially in youn ger forms of “osteolepiform” tetrapodomorphs 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 Thursius macrolepidotus (Sedg wick and Murchi son, 1829) or Osteolepis macrolepidotus Agassiz, 1835, the branch ing of tubes ris ing from these ca nals is weak, and the tu bule open ings are of ten ar ranged in a sin gle row (Jarvik, 1948). An in ter me di - ate con di tion was re corded e.g., in the low er most Up per De vo - nian Latvius grewingki (Gross, 1933) with tu bule open ings ar - ranged in dou ble to tri ple rows (Jarvik, 1948).
“Osteolepiform” tetrapodomorphs have been ob tained from fresh wa ter and brack ish wa ter as well as ma rine strata, and the pos si bil ity of their anadromy has been dis cussed (see Thomson, 1969, 1980). Ac cord ing to Long et al. (1997), the sen sory pore groups, which are dis tinctly de vel oped mainly in spec i mens from fresh wa ter de pos its and ab sent in the ma rine Gogonasus andrewsae Long, 1985 could in di cate sa lin ity. The spec i men from the Moravian Karst de rives from a rel a tively deeper wa ter en vi ron ment with clearly ma rine fauna. Un for tu - nately, the lat eral extrascapulars seem not to be suit able for di - ag no sis of the pres ence of sen sory pore groups, so the sug ges - tion by Long et al. (1997) can not be con firmed nor dis proved.
More over, many “osteolepiforms” were ob tained from red bed fa cies, the in ter pre ta tion of which as re gards sa lin ity is still prob - lem atic (Laurin and Soler-Gijón, 2010).
The “osteolepiforms” have been sub jected to nu mer ous anal y ses (e.g., Long, 1985a, b; Ahlberg, 1991; Young et al., 1992; Ahlberg and Johanson, 1998; Coates and Fried man, 2010; Swartz, 2012; Witzmann and Schoch, 2012; Hol land, 2013) lead ing to dif fer ent in ter pre ta tions of re la tion ships within this group. Swartz (2012) used the term “osteolepiform” to en - cap su late the grade of tetrapodomorphs, in clud ing the stem-based Canowindridae, Megalichthyiformes and Tristichopteridae.
Cosmine cover is com pletely ab sent in tristichopterids (e.g., Snitting, 2008), and canowindrids are sup posed to be a Gondwanan group (Young et al., 1992). It is prob a ble that the spec i men from the Moravian Karst be longs to the megalichthyiforms sensu Swartz (2012). Young et al. (1992) de scribed the shape of lat eral extrascapulars in var i ous
“osteolepiform” groups (the char ac ter “C2: equi lat eral shape of extrascapulars” in their cladistic anal y sis). The con di tion of three-sided lat eral extrascapulars al most meet ing in the midline in canowindrids and megalichthyids vs. four-sided lat eral extrascapulars well sep a rated in the an te rior midline in tristichopterids and “osteolepidids” such as Osteolepis Agassiz, 1835 or Gyroptychius M’Coy, 1848 was later dis cussed by Fig. 6. Scanning electron micrograph of the cosmine surface
with the sensory tube opening
Johanson and Ahlberg (1997). These au thors pointed out that the lat eral extrascapulars of the megalichthyid Cladaro - symblema Fox et al., 1955 are four-sided and not three-sided as should be ex pected, and they re vealed some con tra dic tion be tween the de scrip tions and real shapes of these bones in some other gen era. Johanson and Ahlberg (1997: p. 49) di - vided the char ac ter into two: “three sided lat eral extrascapulars”
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 the lat eral extrascapular of the Moravian spec i men is closer to megalichthyids than “osteolepidids”. Nev er the less, it is nec es - sary to treat such an as sign ment with cau tion. The dis tance be - tween lat eral extrascapulars in the Moravian spec i men is not known. In ad di tion, these fea tures are gen er ally rather prob lem - atic. Both groups are closely re lated (see Janvier et al., 2007) and com prise megalichthyiforms ac cord ing to Swartz (2012).
The spec i men de scribed in this pa per is the first re corded
“osteolepiform” tetrapodomorph from the Pa leo zoic se quences of the Moravian Karst. World wide, other known Famennian cosmine-bear ing “osteolepiforms” in clude e.g., Megapomus Vorobyeva, 1977 and Cryptolepis Vorobyeva, 1975 (Baltica Prov ince), Megistolepis Obruchev, 1955 (Si be ria Prov ince) and Sterropterygion Thomson, 1972 (Laurentia Prov ince) (see Lebedev and Zakharenko, 2010). A Famennian or pos si bly a Tournaisian age is also sup posed for Medoevia lata Lebedev, 1995 which is from an un known lo cal ity (Lebedev, 1995). Al - though these gen era could also pos ses nu mer ous and evenly dis trib uted open ings of sen sory ca nals, we avoid as sign ing the Moravian spec i men to any of them be cause the ma te rial com - prises only one bone, and the lat eral extrascapulars are known for their shape vari abil ity (see Säve-Söderbergh, 1933).
Its po si tion at the Laurussian con ti nen tal mar gin (Kalvoda and Bábek, 2010) and its pre sumed or i gin in a rel a tively deeper wa ter en vi ron ment on the by pass slope (see Weiner and Kalvoda, 2016) sug gest as sign ment of this Moravian
“osteolepiform” ac com pa nied by chondrichthyans and
“palaeoniscids” to the con ti nen tal mar gin as sem blage of the Baltica Prov ince pe riph ery sensu Lebedev et al. (2010). How - ever, there is still a lack of de tailed data on the fish fauna from the Ochoz sec tion or from other Moravian Karst lo cal i ties rep re sent - ing a sim i lar mid Famennian en vi ron ment. From the Czech Re - pub lic, re mains of “osteolepiform” fish have been re ported from limnic Up per Car bon if er ous and Lower Perm ian de pos its (Zajíc, 2000, 2008; Štamberg and Zajíc, 2008). Iso lated re mains of Megalichthys nitens Fritsch, 1889 were re corded from the Car - bon if er ous Kounov Mem ber of the Slaný For ma tion of the Kladno–Rakovník Ba sin (Fritsch, 1889, 1893; Romer, 1945;
Štamberg and Zajíc, 2008). Re mains as signed to
“Osteolepiformes” indet. are known from the Car bon if er ous Kounov Mem ber of the Slaný For ma tion in the Mšeno– Roudnice
and Kladno–Rakovník bas ins, the Klobouky Ho ri zon of the Línì For ma tion in the Kladno–Rakovník Ba sin and the Perm ian Kalná Ho ri zon of the Proseèné For ma tion in the Krkonoše Piedmont Ba sin (Štamberg and Zajíc, 2008).
CONCLUSIONS
To date, De vo nian and Car bon if er ous de pos its of the Moravian Karst (Moravo-Silesian Ba sin, Bo he mian Mas sif) have yielded poor ma te rial of sarcopterygian fish, com pris ing only onychodontiforms (see Ginter, 1991; Smutná, 1994, 1996;
Kumpan, 2013). The newly re corded subtriangular cosmine- cov - ered right lat eral extrascapular co mes from the pre dom i nantly hemipelagic suc ces sion of the Famennian Køtiny Lime stone of the Líšeò For ma tion. The com bi na tion of cosmine histological fea tures in clud ing an ab sence of Westoll lines and bur ied odontodes, enamel/enameloid not ex tend ing into the pore-ca - nals, pore-ca nals in a shape rem i nis cent of oast-house chim - neys and the di am e ter of the pore-ca nal open ings dem on strate an “osteolepiform” af fin ity. This as sign ment is sup ported by the lack of an over lapped area at the me dial bone mar gin. The bone bears nu mer ous evenly dis trib uted open ings of the sen sory ca - nals as is usual in stratigraphically later forms of “osteolepiform”
tetrapodomorphs (Jarvik, 1948), and it lacks the lower mesh-ca - nals pres ent es pe cially in some Mid dle De vo nian forms (Thomson, 1977). The as sign ment to the “Mega lichthyiformes”
sensu Swartz (2012) seems to be most prob a ble.
The as so ci ated fauna has a clearly ma rine char ac ter and con tains e.g., clymenids, orthocone nautiloids, thin-shelled bi - valves, trilobites, cri noids, cono donts, ostracods, chondri - chthyans and “palaeoniscids”. Com pared to nu mer ous
“osteolepiforms” from red bed se quences, the Moravian spec i - men orig i nates from the rel a tively deeper-wa ter en vi ron ment of the by pass slope (see Weiner and Kalvoda, 2016).
“Osteolepiforms” are re corded in the Pa leo zoic se quences of the Moravian Karst for the first time. In the Czech Re pub lic, the re mains of these tetrapodomorphs have pre vi ously only been re ported from the Car bon if er ous and Perm ian limnic bas - ins (Štamberg and Zajíc, 2008).
Ac knowl edge ments. Thanks are due to O. Fatka and J. Kalvoda for their help ful com ments. We are grate ful to J. Štelcl for prep a ra tion of the scan ning elec tron mi cro graphs and T. Viktorýn for his kind help in the field. Cor dial thanks are ad dressed to the re view ers J. Clack and M. Ginter for their use - ful sug ges tions. This con tri bu tion was sup ported by the Czech Sci ence Foun da tion pro ject GA14-18183S and In sti tute Re - search Plan RVO67985831.
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