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Home , Brachythoraci, Dinichthys, Dunkleosteus, Eastmanosteus, Holonema, Holonematidae

Comments on the Late placoderms from the Holy Cross Mountains (Poland)

ALEXANDER IVANOV andMICHAŁ GINTER

Ivanov, A. & Ginter,M. 1997. Comments on the Late Devonian placoderms from the Holy Cross Mountains (Poland).- Acta Palaeontologica Polonica 4,3,4I34f6.

Taxonomy of the Late Devonian placoderm remains from the Holy Cross Mountains, Poland, described by Gorizdro-Kulczycka (L934,1950) and Kulczycki (1956, 1957),is revised. Several recently found specimens are also mentioned. The old collections are composed of representatives of Ptyctodontidae, , Plourdosteidae, Pholi- dosteidae, Selenosteidae, Titanichthyidae and Dinichthyidae, the latter with an unde- scribed of . Newly found specimens belong to Ptyctodontidae, Plourdosteidae and Dinichthyidae. The occurrence of the Antiarcha in the Late Devonian of the Holy Cross Mountains, suggestedby former authors, has not been confirmed.

K e y w o rd s : ,Late Devonian, Holy Cross Mountains, Poland.

Alexander Ivanov [[email protected]], Laboratory of Paleontology, Institute of the Earth Crust, Sankt-Petersburg University, 16 Liniya 29, 199178 St.Petersburg, Russia. Michał Ginter [email protected]], InsĘtut Geologii Podstawowej, Uniwersytet War szaw ski, ul. Zw irki i Wi gury 9 3, 02 -089 War szaw a, P oland.

Introduction

The main goals of this study are to revise placodermsfrom the Late Devonian of the Holy Cross Mountains collected by Jan Czarnocki, Julian Kulczycki and Zinuda Gorizdro-Kulczycka in the first half of the century (the material now housed in the Museum of the Polish Geological Instituteand in the Museum of the Earth in Warsaw), and to describe a fęw new specimenscollected recently by Jerzy Dzik and Grzegorz Racki. The old collections have formerly been described in part by Gorizdro- -Kulczycka(1934,1950) and Kulczycki (1956,1957).The specimensin thesecollec- tions come from Wietrznia Quarry, Kadzielnia Quany and Psie Górki Hills in Kielce, and from the vicinities of Gałęzicein the westernpart of the Holy Cross Mountains. The new specimens were found in Wietrznia Quarry, Kowala (south of Kielce), ostńwka Quarry tnGatęzice region, and Płuckiin the vicinities of Łagów, easternpart of the Holy Cross Mountains(Fig. 1). 414 Late Devonian placoderms.' IVANOV & GINTER

m Early and Middle Devonian

f-777 t) Late Devonian and Cańoniferous

0 10km

Fig. 1. Location of outcropsmentioned in the paper on a sketchmap of Palaeozoic core of the Holy Cross Mountains; K _ Kadzielnia, Kw _ Kowala, Ł+Pł _ł-agów and Pfucki, o+G _ ostrówka and Gałęzice,P_ Psie Górki, W - Wietrznia.

Following abbreviationsof the institutionnames arc used herein:GruS - Geologi- cal Institute,University of Silęsia, Sosnowiec; MZ - Museum of the Earth, Polish Academy of Sciences, Warsaw; PIG - Polish Geological Institute,Warsaw; ZPN- - Instituteof Palaeobiology,Polish Academy of Sciences,Warsaw.

Stratigraphy

Any analysis of the stratigraphicaldistribution of placodermsin the Late Devonian of the Holy Cross Mountains faces considerabledifficulties. First of all, the specimens from old collections were datedaccording to the subdivisionof the Late Devonian used by Czarnocki (in Gorizdro-Kulczyck a 1934:p. 14;see also Czarnocki 1947; Gorizdro- Kulczycka 1950; Kulczycki 1957). Only the part of that subdivision conesponds roughly to modern zonations, based on goniatites or . The was subdivided into three local zones (Polish 'poziom'): zone I, with Hypo- thyridina procuboides; zone II, with H. coronula; and zone III, with H. cuboides and Mąnticocerasintumescens; Kulczycki (1957)used the terms .Lower', .Middle' and 'UpperFrasnian', respectively.There is no confirmationthatthe characteristic brachio- pod and goniatite speciesmentioned above have ever been found togetherwith, or at least in the same horizon as the specimensof placodermsunder study.Moreover, it is possible thatCzarnocki applied a superficiallithostratigraphic correlation to subdivide the rocks in a particular quarry or such a correlation might even have been used betweenthe quarries (Szulczewski, personal communication 1994).This could have lead to very confusing results,since Frasnian and lower Famennianrocks of the Holy Cross Mountains were depositedin complicatędconditions of Synsedimentaryblock tectonics (Szulczewski 1989).In Wietrznia Quarry a single lithostratigraphicunit can span differentnumbers of conodontzones even at a distanceof severaltens of meters. ACTA PALAEONTOLOGTCA POLONTCA (42) (3) 415

The same lithostratigraphicunit may representeither the late Frasnian or an interval from the late Frasnian through the early Famennian.Thus, for instance,if a museum label reads that a specimencomes from the Frasnian zone III, one cannotbe sure that it was found togetherwith 1L cuboidesor M. intumencens,andone cannot even be sure whetherit was really found in Frasnian rocks. Famennianspecimens come from the so-called'Cheiloceras-beds'and 'Clymenia- -beds'. In this case it is possible to determine that the specimens found in the Cheiloceras-beds are of the unspecified early Famennian age. Those found in the Clymenia-beds are of late Famennian age, probably from the interval of the Early Pąlmatolepis marginifera throughthe Middle P. praesulcata conodontZones. The precision of dating of the new material is different in particular cases. The specimens from Wietrznia (undeterminedMesotąxis asymmetrica Zone), Kowala road cut and Pfucki (Late P rhenana-P. linguifurmis Zones) come from samples whose age was determinedby conodonts.The specimenfrom ostrówka has no exact age determination,but it is known that it comes from the upper Famennian part of the Quarry (J. Dzik, personalcommunication 1994). The revised and updateddistribution of placodermsconsidering the new findings is shownin Table 1 andZ.For thedetailed stratigraphy of theoutcrops see Szulczewski (1971,1989),Szulczewski et al. (1996),and Racki (1993).

Order Gross, I93f Family Ptyctodontidae Woodward, 1891 Gorizdro-Kulczycka(1934) describedtwo new speciesof Ptyctodus:P. czarnockii and P kielcensis,as well as P. obliquusPander, 1858, P. sp.,and a new speciesRhynchodus marginnlisbased on the isolatedtoothplates. Ptyctodus czarnockii, P. obliquus,P sp., andRhynchodus marginalis were reportedfrom the zone II of the Frasnianof Wietrznia Quany. The remains of P kielcensis and P obliquus occur in the zoneIII of Wietrznia and Kadzielnia Quarries, respectively. Ptyctodus czarnockii is representedby two tooĘlates' one of them (Gorizdro-Kulczycka 1934:pl. I: 1, 2) belongsto the upper jaw tooĘlate (supragnathalplate) with curved oblique anteriorpart and flat posterior part of the biting surface,characteristic of Rłynchodus,and is very similar to thatof R. tetrodon(Jaekel, 1903).Second specimen(Gorizdro-Kulczycka 1934:pl. I: 3) is a fragment of lower tooĘlate which is difficult to attribute to a definite of ptyctodonts.The toothplateof Ptyctoduskielcensls (Gorizdro-Kulczycka 1934: pl. II: 5) should be referredto Rhynchodusbased on the shape of biting surface.Under the name Ptyctodus obliquus she also described the upper toothplate of Rhynchodus (Gorizdro-Kulczycka 1934: pl. tr: 4) with curved oblique biting surface and well developed anterior dorsal process' as well as ,Ptyctodus'-typetooĘlates (unillus- tratedspecimens from the collection of the PIG museum).The tooĘlates referredto as Rhynchodusmarginalis (Gorizdro-Kulczycka 1934:pl. II: 7, 8) were not found in the museum collections. The determination of their taxonomic position from thę illustrations alone cannotbe done. Thus, in the original material published by Gorizdro-Kulczycka (1934), ptycto- donts are representedby tootĘlates of Rhynchoduscf . tetrodon(Jaekel, 1903),Rhyn- 'Ptyctodus' chodussp., and sp. Afew new specimenswere found in the lower Frasnian of Wietrznia Quarry (M. asymmetricaconodont Zone; new zonation:Early P. falsio- 416 Late Devonian placoderms; IVANOV & GINTER

Table 1. Distribution of placoderms in the Late Devonian of the Holy Cross Mountains: collections of J. Czarnocki, Z. Gorizdro-Kulczycka and J. Kulczycki; taxa revised by presentauthors. Conodont zones: tri - Palmatolepis triangularis, cre - P. crepida, rho - P. rhomboidea, mar - P. marginifera, tra - P. trachytera,pos-P.postera,exp-P.expansa,prae-P.praesulcata,E-early,M-Middle,L-Late, L* - latest.

E a o o PLACODERM (UCD EE TAXA U) sŃ _L_ prae !t!

E ___l __l I t_ a I exp I $ o I o o E -o ---I (U L- .EI 'ao pos I I E Ę I I .C z o --'l t--- .E L tr I I .9 z tra E \ I c z o I _-..1 ó UJ -L:

mar L_ I q) E I i LJL t--- l- - i '-- o I s c; rho -L- o I o a N O) E o ro I o (Ą -o t- a- - ci' 'o, g-.=' I -L: tĄ Ę ,, cĄ a .Ą g eN Śs € (Ą s o CĄ cre l- :B 5 g o o i'o ś< Ę M o s b Ę ci U) ś (U o o ro € s a E r_ -o. -: 'oe 3 s q) s_ CĄ 'a * L il t\ b ro s (Ą Ę o -$ .€ -t 'os -(Ąr-o- :B $- tri (Ąrt I ict .9 M o s e (Ą s Ę ut b $ (U o o lp E s -t o t- .Ę -Ę .o rO' rA- t- o o a o rO i= o Ę Ę a- E rp o s rJ I o: Zorrelll o eo g o ro I .ei s I Ę Ę I o Ę io- I I o; t_ --L-- Bi_ z I u)t l I ,l 5r I Q)r z I a)l a Zonell I or I q)l I ś(: t --r--I LL t- t- Il t-- I ll I ai Zone I I I I

1) seeGorizdro-Kulczycka (1934: p. 14; 1950).

valis-Early P. hassi Zones). These are toothplatesand fragmentsof the exoskeleton. Among the first type there are few 'Ptyctodus'-typetoothplates and abundanttooth- ACTA PALAEONTOLOGICA POLONIC A (42) (3) 4r7

Table 2. New findings of placodermsin the Late Devonian of thę Holy Cross Mountains: collections of J. Dzik (7FN-) and G. Racki (GIUS). E - early, M - Middle, L - late, L* - latest.

STAGES PLACODERM TAXA AND AND LOCALITIES CONODONTZONES

L I I I I I I Palmatotepis praesulcata M I I I I I I I I E I I I I I I L I I I I P. expansa M I tl I :- I tl I tr I L _ _ _ _ _l_ _l L I I I I P. postera I I I I z E I I I I I P trachytera L I z E I z -+-- L* I LJJ I = P. marginifera L I I E I lFiullt-t-l LL -'T-- --i- -l--$-- . L I ć--i-E P ńomboidea I l6ilĘ,= E -'T--I --r-ta,rrr;'9 - q--t- -'-t--|,-- L* I I .(o id i #io P. crepida L I c .Ol iEisiF; M El N |Ęt(Dl|.. Nt o lQrĘlĘ E - -i-'I -i--s--!- € -:--E-- L .9 = - -lĘ -J-. _i__E _l__s _i_E P. triangularis M o-.iuJ |ó di o ls E a (Ą rrLLiX - -J-. i s P. linguiformis (Ą| B - 5 -r-- o -i--l--i---[--i-d -- L bl s P rhenana €| o z E Ol rttl - *$ -+-'€ - F - - - - -t- -t------P.iamieae $! t tl z -'t-- - |------t- -l ------L I rtl a P fiassi I ltl rtl E ------t - - F t- t P. punctata ttl ------r - - tL F r- ttl P transitans ------r - - L i- r- p falsiovalis ttl E tll li ttl plates of Rhynchodus, mostly the upper ones (Figs. 2A-C). The upper toothplates of Rhynchodu,s resemble those of R. rostratus Eastman, 1898 or R. SeCcłnSNewberry, 1873 (Eastman 1898a), but specimens from the Holy Cross Mountains are much smaller (4_6 mm long). The anterior dorsal process is thin and thę groove for attach- ment to the autopalatine is located close to the anterior edge. The biting surface is nalrow and a small cusp is present on the lingual side. Of the exoskeletal plates there is a fragment of a median dorsal spine (Fig. 2D) and an incomplete median ventral plate (Fig. 2E). The former resembles median dorsal 418 I-ąte Devonian placoderłzs.. IVANOV & GINTER

Fig. f. A-C, E. Rhynchodussp., Frasnian, Mesotaxis asymmetricaconodont Zone; Wietrznia Quarry.A-C. Uppertoothplatesinlingualviews:A-ZPALP.VVI,x 15,B -ZPN,P.yUf,x 16,C -ZPN-P.W3,x22: E - median ventral plate in ventral view, ZPALP.VV4, x 17. D. ftyctodontidae gen. indet., fragmentof median dorsal spine in lateral vięw, ZPAI' P.VV5; Frasnian, M' asymmetrica conodontZone; Wieffznia Quarry; x 40.

spines of Campbellodus (Long 1987), the latter belongs to the exoskeleton of Rhyn- chodus (Ivanov I99f). Most of Rhynchodus species are mainly known from the Eifelian-Frasnian of Germany and (Denison 1978). Latest representatives of this genus on the East European Platform were recorded from the early Famennian (Ivanov 1992). ,Ptyctodus,-type tooĘlates are very cofirmon in the Middle and Late Devonian carbonate sections.

Order Woodward, 1891 Family HolonematidaeObruchev, 1933 Kulczycki (1957) described and illustrated radiatum Obruchev, 1933, as well as new genera and species Deveonema obruchevi and Operchallosteus vialovi from the zone II of the Frasnian. The attribution of the incomplete anterior dorsolateral plate to H. radiatum is apparently correct. The identification of the middle part of the median dorsal plate as Deveonema obruchevi is rather doubtful. A tuberculated orna- ment and narrow central part between the overlap areas are main characters of the ACTA PALAEONTOLOGTCA POLONTCA (4f) (3) 419 alleged new genus.However, the smaller individuals of ,F/.westolli Miles, 1971 have a mediandorsal platecovered with the tubercleswhich are alrangedin therow in places (Miles I971: fig. 19).The overlaps on the visceral surfaceof the median dorsal plate are broader in these individuals than those in large ones and respectively the central part betweenthe internalmargins of overlapsis narrower(Miles l97I: p. 156).It is possible, therefore,that the fragment of median dorsal plate from the Holy Cross Mountains belongs to a small individual of Holonema. The diagnosis of Operchnllosteus vialowi Kulczycki, 1957 was based on the characters of ornament of a plate fragment, identified by Kulczycki (1957) as a posteńor ventrolateralplate.This fragmentwas redescribedby obruchev (|964) and Miles (1971) as a posterior part of an anterior ventrolateralplate of Gyroplacosteus vialowt. Most probably this specimenshould be referredto as Gyroplacosteuspanderi obruchev, 1933 which is characterisedby the similar outline of anteńor ventrolateral plate and the ornamentof ventral plates. This being the case' the representativesof Holonematidaefound in thę Holy Cross Mountains should be referred to the following taxa: Holonema radiatum, Holonem.a sp. and Gyroplacosteussp. The first was reportedfrom the upperpart of lower Frasnian of the Main Devonian Field (obruchev 1933), the Frasnian of kan (Lelićvre et al, 1981), as well as possibly from the Eifelian of Spain (lrlidvre et aI. 1990).The representativesof Gyroplacosteusoccur in the the upper lower Frasnian of the Main Devonian Field in associationwith Holonema radiatum, theFrasnian of the Kuznetsk Basin (Obruchev 1933)and the lower Frasnian of Germany (Miles 1971).

Family Plourdosteidae Yózina, 1990 Kulczycki (1957)described Plourdosten.r sp. on the basis of two plates:inferognathal and marginal from the Frasnian zone II of Wietrznia Quarry. From the same beds he establisheda new genus and species Tomaiosteusgrossi, which is representedby a fragmentof skull roof and a posteriorsupragnathal plate. Denison (1978)synonymised this genus with Plourdostens. The inferognathal and posterior supragnathalplates belong to Plourdosteus, while the skull roof fragment and the marginal plate are attributedto some plourdosteidarthrodires. The supraorbitalsensory line divides the preorbital plate into the central and lateral parts.The central part of the preorbital in this specimenis long and of similar size to the lateralpart. Such shapeof the preorbital and shapeof dorsal and visceral surfaceof the pineal plate are characteristicof some plourdosteids:Harrytoombsia (Miles & Dennis 1979), Janiosteus (Ivanov 1988), Mcrnmara,qpls (Long 1995) and Torosteus(Gardiner & Miles 1990). In addition, Kulczycki (1957) mentionedthe incomplete median dorsal plate from the same beds as oxyosteł.'sp. This plateprobably belongsto coccosteidor plourdosteidarthrodires. There are new findings of Plourdosteusin the Holy Cross Mountains: a posterior dorsolateralplate from the late Frasnian,the Late P. rhenaru-P linguifurmis Zones of Kowala road cut and an anterior ventrolateralplate from the late Frasnian of Pfucki. The posterior dorsolateralplate has a broad overlap area for the median dorsal plate, relatively n.uTowoverlap area for the anterior dorsolateralplate and a cavity for thę posteriorlateralplate. Margins of the frst two form an almostńght angle.The posteńor margin of the plate is curved and shghtly convex. There is no groove for canals on the external surface. The anterior ventrolateralplate is rounded in shape, its length is 4f0 Late Devonian placoderrzs: IVANOV & GINTER

.Zone'III;Psie Fig. 3. A_G. Malerosteus7orizdroae Kulczycki, 1957,Frasnian, Górki; coll. J. Kulczycki. A-C. Nasal capsulein anterior(A), posterior(B) and lateral(C) views; MZYIIWp-3Z/I, x 2.8.D-G. Left postnasalplate in right lateral (D, nasal notch),visceral (E), external(F), and oblique dorsal (G) views, MZ YInNp-3212, x 2.8. similar to breadth.A groove for the ventral transversesensory line is presentin the anteriorpart of the plate.

Family PholidosteidaeGross, 1932 Kulczycki(1957) erected a newgenus and species Malerosteus gorizdroae based on the incomplete nuchal plates, articulated paranuchal, marginal and postorbital plates, articulated preorbital and suborbital plates. These specimens were found in the Fras- nian zone III of Psie Górki. There are two unpublished specimens of Malerosteus gorizdroae, which Kulczycki has extracted together with the described preorbital and suborbital plates: a nasal capsule (Fig. 3A-C) and a postnasal plate (Fig. 3D-G) in the collection described by Kulczycki (1957). These bones rarely occur źImongthe isolated arthrodiran remains and are new for Malerostezs. The nasal capsule is frustum-shaped. The broad anterior surface is concave and oval. It bears numerous small foramina (Fig. 3A). It is the ACTA PALAEONTOLOGTCA POLONTCA (42) (3) 421 cribrosal region of the postnasalwall. The posterior side of the bone has a rounded depressionwith the openings of various size (Fig. 3B). The largestof them probably fitted the olfactory nerve. The lateral surface of the nasal capsule is spongious with multiple fine canals.The mentionedspecimen resembles the nasal capsuleof Eastma- nosteuscalliaspis Dennis-Bryan, 1987,but the latterhas a high conical shapeand deep depressionon the posteriorside (Dennis-Bryan1987: fig. 16). The postnasalplate has a trapezoidalexternal surface with broad ventraland narrow dorsal edges.The latter is a contactface for the preorbital plate. The margin of fairly large nasal notch extendson the visceral side as a long interfenestralprocess (Fig. 3D). The narrow cavity for the contactof the suborbitalplate is placed on the oppositeside to the nasal notch. There is no groove for canals on the externalsurface. The visceral side of the plate bears a transverselyelongate groove (Fig. 3E).

Family DinichthyidaeNewberry, 1885 Kulczycki (1956, 1957)described various materialasDinichthys pustulosusEastman, 1897.The largest part of the collection comes from the Frasnian, mainly the zone II, of Wietrznia: fragmentsof skull roof, nuchal, paranuchaland anterior lateral plates, median dorsal plates and parasphenoid.The most completepart of the skull roof was found in the zone I of the Frasnian, and two fragmentsof median dorsal plates in the zone III. A single anterior supragnathalplate was reported from the Famennian 'Cheiloceras beds' of Kadzielnia. Most of the isolated plates are incomplete and it is difficult to identify the genus,but all of them belong to Dinichthyidae.TWo specimens of the skull roof, paranuchalplate and parasphenoidundoubtedly belongto Eastma- nosteus.They differ from Eastmnnosteus() pustulosłs (Eastman' 1897) from the Givetian-early Frasnian of North America (Eastman1898b, 1907), E. callias- pis Dennis-Bryan, 1987 and other species of Eastmanosteusby the shape of nuchal, paranuchal and central plates on the external surface.The lateral lobe of the central plate of the North American speciesis much shorterand broaderthan it is in Eastmą- nosteus from the Holy Cross Mountains. The contacts of the central plate with postorbitaland marginal ones in the Polish specimensnearly overlie on one line. The suturebetween the central and marginal plates is longer in the Kulczycki's specimens (MZ VtrWp-ll, MZ VIII/I/p-L2). The anterior margin of the nuchal plate in the American speciesis concaveand bearstwo large anterolateralprojections, but the same margin in Eastmanosteusfrom Poland has three small projectionswhich are equal in length.The posterolateralparts of the nuchal plate in the skull roof from theHoly Cross Mountains bear transversegrooves. The nuchal plate of E. pustulosus described by Eastman has a breadthof posterior edge to breadthof anterior edge ratio of 3.1. Eastmanosteusfrom Poland has this ratio of 2.6. T\e paranuchalof this speciesis longer and narrowerthan thatin the American species,and it has a broaderpostnuchal process.Thus, Eastmanosteusfrom the Holy Cross Mountains should be referredto a new species.The formal descriptionof the specieswill be presentedin a separatestudy. In the new placoderm material there is a skull roof fragment (Fig. 4A) with inferognathalplate of thejuvenile individuals from the Late P. rhenanaconodontZone of Płucki (nearŁagów), which can be attributedto the same' new Speciesof Eastma- nosteus. Aan łLL Inte Devonian placoderms: TVANOV & GINTER

Fig. 4. 4,.Eastmnnosteus sp. n., juvenile, fragment of skullroof in dorsalview, anterior margin is up;ZPAL P'YU7 Frasnian,Late Palmatolepis rhenana conodont Zone;Łagów, Płucki; x 1.8.B. Dunkleoste'J'sp., nuchalplatein ventralview;ZPAL P.W6; upperFamennian; ostrówka Quarry; x 1.

Kulczycki (195'7) established two new species of Dinichthys: D. ceretus and D. denisoni based on the incomplete median dorsal plates. Both taxa occur in the Famen- 'Cheiloceras nian, beds' of Kadzielnia, and the latter has also been found in'Clymenia beds' of Gałęzice. The median dorsal plate of first species belongs to DinichĘidae and resembles that of (Heintz I93Z) and Heintzichthys (Carr 1991). Denison (1978) referred Dinichthys denisoni to the genus Dunkleosteus. ACTA PALAEONTOLOGTCA POLONTCA (42) (3) 423

The inferognathalplate describedby Kulczycki as Dinichthys cf. tuberculatus Newberry,1888 from theFamennian, ,Clymeniabeds' of Gałęzice(Kulczycki 1957: pl. VIII: 2) is similar to Dunkleosteusterrelli (Newberry, 1873) and Dinichthys herzeri Newberry, 1868, and possibly pertainsto Dunkleosteus.Another specimen from the same beds, the fragment of the posterior ventrolateral plate (Kulczycki 1957: pl. VIII: 3) referred to Dinichthys cf . tuberculatusby Kulczycki does not belong to Dinichthyidae.However, it is difficult to identify its taxonomicposition. A single nuchal plate of Dunkleosteushas recently been found in the late Famennian of ostrówka Quarry Gig. 4B). This plate has a rectangularanterior margin and deep overlap areasfor the central and paranuchalplates on the visceral surface,which arę characteristicfeatures of Dunkleosteus.Itresembles more closely that of Dunkleos- teus marsaisiLehman,1954 from the early Famennianof (Lehman 1956) by the shape of dorsal and visceral surfaces of the nuchal, especially by the long posterolaterallobes of the plates. Apart from the placoderms mentionedabove, Kulczycki (1957) described a new species Titanichthyskozlowskii and Pachyosteus bulla Jaekel, 1903 from 'Clymenia beds' of Gałęzice.Aspidichthys ingens Koenen, 1883 identified by Kulczycki as Anomalichthys ingens (Koenen, 1883) was found in zone III of the Frasnian, as well as in the late Frasnian Manticocerąs beds of Rheinland, Germany, the Frasnian of Morocco and Iran (Denison 1978).

Order Antiarcha Cope, 1885 In her lists of ichthyofauna,Gorizdro-Kulczycka (1950)reported Bothriolepis panderi Lahusen, 1880, B. cf. m.aximaGross, 1933 and B. sp. from the Frasnian zone II of Wietrznia. Apart from antiarchs in the same list, she mentioned the occurrence of Onychodusjaekeli, Dipterus sp. and Osteolepissp. from the samelocality. Theseforms are presentin the PIG collection, however,we did not find any antiarchremains there. Kulczycki (1957)figured one fragmentof a plate and assignedit as Bothriolepis. That specimenshould probably be referredto one of the representativesof brachythoracid arthrodires.This shows that thus far no confirmed antiarchremains were found in the Holy Cross Mountains.

Relations to other regions

Most of the studied placoderms belong to widely distributed taxa. Only one genus: Malerosteus and the species of Eastmanosteus, , and Dunkleosteus are unique for the Holy Cross Mountains. There occur species typical for Rhenish facies, Ike Asptdichthys ingens and Pachyosteus bulla, together with Holoneł?w radiatum,, which is known from several localities on the East European Platform (Main and Central Devonian Fields, North Timan) and was also found in the eastern part of Iran. Many forms which apparently coexisted in the Holy Cross Mountains were found together in some other regions, too: Aspidichthys ingens and Pachyosteus bulla in the Frasnian, Mąnticocerąs beds of Rheinland (Gross 193f); Titanichthys and Dunkleos- teus in North America, Cleveland Shalę (Can 1991); Holonema and Eastmnnosteus in 424 lnte Devonian placoderns.' IVANOV & GINTER the in Australia (Long 199I); and Holonema, Gyroplacosteus and Rhynchodusin the upper part of the Lower Frasnian of the Main Devonian Field. All placodermsmentioned in this paper were characterisedeither by high environ- mental tolerance (Holonema, Gyroplacosteus,Eastmąnosteus, Rhynchodus) or were connected with pelagic carbonate facies: Aspidichthys ingens, Pachyosteus bulla, Titanichthysand Dunkleosteus.Pelagic conditions are confirmed by the absenceof antiarchsand psammosteids,which usually predominateamong the ichtĘofauna of shallow water terrigenousdeposits.

Acknowledgements

Warm thanks are due to Professors Jerzy Dzik (Institute of Palaeobiology, Polish Academy of Sciences, Warsaw) and Grzegorz Racki (University of Silesia, Sosnowiec), who donated their materialfor investigation; to Prof. Michał Szulczewski (Warsaw University), who provided strati- graphic information; and to Dr. Elga Kurik (Instituteof Geology, Tallinn) and Dr. Kim Dennis-Bryan (London), who kindly revised the manuscript.

References

Carr, R. 1991.Reanalysis of /{elntzichthysgouldii (Newberry),an aspinothoracidarthrodire (Placodermi) from the Famennian of northern Ohio, with a review of brachythoracid systematics.-Ztological Journal ofthe Linnean SocieĘ 103,349_390, Czarnocki, J . 1947.Przewodnik XXZjazduPolskiego TowarzystwaGeologicznego w Górach Świętokrzys- kich. - Rocznik Polskiego Towar4tstwaGeologicznego !7,237-299. Denison,R. 1978.Placodermi, In: H.-P, Schultze(ęd,) Handbook of Paleoichthyology2. I28 pp. Gustav Fischer Verlag, Stuttgart-New York. Dennis-Bryan, K. 1987. A new species of eastmanosteidarthrodire (Pisces: Placodermi) from Gogo, Western Australia. - Zoological Journal of the Linnean SocieĘ 90, I_64. Eastman,C.R.lS9Sa.Dentitionof DevonianPtyctodontidae.-AmericanNaturalist32,473488. Eastman,C.R.lS9Sb.Somenewpointindinichthyidosteology.-ArnericanNaturalist32,T4T-768. Eastman,C.R. 1907.Devonic of theNew York formation.- Memoirs of theNew York StateMuseum of Natural History I0,I-235. Gardiner, B.G. & Miles, R.S. 1990. A new genus of eubrachythoracidarthrodire from Gogo, Western Australia. - hological Journal of the Linnean Society 99,159-204. Gońzdro-Ku|czycka,Z.Ig34.Ptyctodontidae górnodewońskiez Gór Świętokrzyskich. - Prace Polskiego In stytutu Geolo gic zne go 3, 1-38. Gorizdro-Ku|czycka, Z, 1950' Dwudyszne ryby dewońskie z Gór Swiętokrzyskich. - Acta Geologica Polonica 1,53-105. Gross, W. I93f. Die Arthrodira Wildungens. - Geologische und Pakiontologische Abhandlungen L9, 5-61. Heintz,A.I93f.Thestructureof Dinichthys,acontributiontoourknowledgeof theArthrodira. - Bashford Dean Memorial Volume- Archaic Fishes 4,II5-224. Ivanov, A.O. 1988. A new arthrodirangenus from the Upper Devonian of the Timan [in Russian].- P aleontolo gić e skij źurnal 2' If3-If5. Ivanov, A.O. I99f . Occurrencesof ptyctodontids(Placodermi) in the Late Devonian of the East European Platform [in Russian]. -Voprosy PaleontologiiL0,I33-I37 . Kulczycki, J. 1956. On the parasphenoid of the .-Acta Palaeontologica Polonica L, 103-11 1. Kulczycki, J. 1957. Upper Devonian fishes from the Holy Cross Mountains. - Acta Palaeontologica Polonicaf,fS5-380. ACTA PALAEONTOLOGTCA POLONTCA (42) (3) 425

Lehman, J.-P. 1956. Les Arthrodires du dóvonien supórieur du Tafilalet (Sud Marocain). - Service GćologiqueMaroc, Notes et Mernoirest29,7-.70. Lelióvre, H., Janvier,P', & Goujet,D. 1981.Les Vórtóbres dóvoniensd'Iran Central.IV: Arthrodireset Ptyctodontes. - Gćobio s 14, 677 _7 09, Lelidvre, H., Goujet, D., & Henn, A. 1990. Un nouveau spócimen d,Holonerna radiatum (Placodermi, Arthrodira) du Dóvonien moyen de la rógion d'oviedo, Espagne.- Bulletin du Musćum national d' Histoire naturelle 12, 53-83. Long, J.A. 1987.Late Devonian fishes from the Gogo Formation,Western Australia _ new discoveńes. - Search18.2O3-f05. Long, J.A. 1991.The long history of Australianfossil fishes. In: P. Vickers-Rich, J.M. Monaghan,R.F. Baird, & T.H. Rich (eds), Palaeontology of Australia, 338428. Monash University Publications Committee,Melbourne. Long, J.A' 1995. A new plourdosteid ańhrodire from the Upper Devonian Gogo Formation of Western Australia. - P alaeontology 3E, 3942. Miles,R.S. l91l.TheHolonematidae(placodermfishes),areviewbasedonnewspecimensofHolonema from the Upper Devonian of Western Australia. - Phibsophical Transactionsof the Royal Society of I-ondon263,I0I-f34. Miles, R.S. & Dennis, K, 1979,A pńmitive eubrachythoracidarthrodire from Gogo, WesternAustralia. - bological Joumal of the Linnean Society 66,3142. obruchev, D. 1933. Holonemidae des russischenDevons. -Travaux de l,Institut Paleo7oologĘueAca- demiedes Sciencesde l'URSS 2,97-116. obruchev, D. (obrućev, D.) 1964.Class Placodermi [in Russian].In: D. obrućev (ed.),osnovy Paleon- tologii I 1, 1I8-I72, Nauka, Moskva. Racki G. 1993.Evolution of the bank to reef complex in the Devonian of the Holy Cross Mountains. - Acta P alaeontolo gic a P olonica 37, 87-I82. Szulczewski M. 1971. Upper Devonian conodonts,stratigraphy and facies developmentin the Holy Cross Mts - Acta GeologicaPolonicaZt, I-If9. Szulczewski, M. 1989. Światowe i regionalne zdarzenia w zapisie stratygraficznym pogranicza franu i famenu Gór Świętokrzyskich' - Przegląd Geologiczny 11, 55l-557. Szulczewski M., Belka 2., & Skompski S. 1996.The drowning of a carbonateplatform: an example from the Devonian- of the Holy Cross Mountains, Poland. - Sedimentary Geology 106, ft49.

Uwagi 9 Późnodewońskich plakodermach z Gór Swiętokrzyskich

ALEXANDER IvANoV i MICHAŁ GINTER

Streszczenie

Celem pracy jest rewizjataksonomicznapóźnodewońskich plakodermów zGór Świę- tokrzyskich, zebranych ptzez Jana Czarnockiego, Juliana Kulczyckiego i Zinudę Gorizdro-Kulczycką. Zbioty te znajdują się obecnie w Muzeum Zieml i w Muzeum PaństwowegoInstytutu Geologicznego w Warszawie.Ponadto opisano kilka nowych okazów, znal.ezionychw ostatnichlatach ptzez I erzegoDz1ka (InstytutPaleobiologii PAN) i Grzegorza Rackiego (Uniwersytet Sląski). Kolekcja starsza zostałapo części opisanaprzez Gońzdro-Kulczyck ą (1934,1 950) i Kulczyckiego(19 56, 19 57 ). okazy badane pochodzą przede wszystkim z Wietrznr. Kadzielni i Psich Górek w Kielcachorazz ostrówki i innych odsłonięćw regionie gałęzicŁ'tm,ataYzez Kowali na południeod Kielc i Płucekw poblizu Łagowa (Fig. 1). Ustalenie wieku okazów ze 4f6 I.ąte Devonian placoderms.. IVr\NoV & GINTER starszychkolekcji (Tabela 1)jest obecnie niezmiernie utrudnione.Wyzej wymienieni aatorzy korzystali, przy określaniupozycji stratygraficznejokazów, z podziatu stra- tygraficznegouzywanego przez Czarnockiego (m.in. w Gorizdro-Kulczycka 1934). Czarnocki stosowałswoisty trójdzielny podział.franu i nie uwzględniałw rozwaŻa- niach stratygraficznychnieizochroniczności granic wydzieleń litostratygraftcznych w dewonie świętokrzyskim,spowodowanej m. in. synsedymentacyjnątektonika blo- kową (Szulczewski 1989). Datowanie okazów nowych przedstawionejest w Tabeli 2. W starszychzbiorach stwierdzonoobecność przedstawicielirodzinPtyctodontidae, Holonematidae,Plourdosteidae, Pholidosteidae, Selenosteidae, Titanichthyidae i Dini- chĘidae. Zilustrowano nie opisaneprzezpoprzedników fragmentyszkie|etuMalero- steus gorizdroe Kulczycki' 1957: puszkę nosową (Fig. 3A-C) i płytkęZanosową (postnasale)(Fig. 3D-G). Wbrew doniesieniom Gorizdro-Kulczyckiej (1950) i Kul- czyckiego (|957) nie potwierdzonoobecności Antiarcha w badanymmateriale. okaz zaliczony przez Kulczyckiego (1957) do Bothriolepis (Antiarcha)jest prawdopodob- nie przedstawicielemBrachythoraci (Arthrodira). Wśródokazów nowych zna|ezionoprzedstawicieli Ptyctodontidae: fragmenĘ pły- tek zębowych(Fig. ŻA_C) i brzusznejśrodkowej (medioventrale; Fig.ZE) Rhynchodus sp. oraz fragment kolca grzbietowego o nieokreślonejprzynalezności gatunkowej i rodzajowej (Fig. 2D). oprócz pĘktodontów lvystępujątam okazy Plourdosteidae i Dinichthyidae, a wśródtych ostatnich duza płytkakarkowa (nuchale),na|eżląca do Dunkleosteus sp. (Fig. 4A) oraz fragmentgrzbietu czaszki wraz z płytkąszczękową dolną (inferognathale)młodocianego osobnika nowego gatunku zrodzajll Eastmano- steus (Fig. aB). Nalezy on prawdopodobniedo tego samegogatunku, co najbardziej kompletny polski okaz Dinichthyidae, opisany przez Kulczyckiego (1957: pl. IV: 1, 2). Gatunkowi temu poświęconebędzie odrębnestudium.