A preliminary note on new ostracoderms from the Lower Devonian (Emsian) of central Poland

A C T A Vol. II

P A L A E O N T O L O G I C A I 95 7

L.BEVERLY TAR LO

P O L O N I C A No•. 2-3

A PRELIMINARY NOTE ON NEW OSTRACODERMS FROM THE LOWER DEVONIAN (EMSIAN) OF CENTRAL POLAND Abstract . - The Emsian sedime n ts of the Holy Cross (Swi ety Krzyz) Mount ains in Centr-al Poland inclu de an essenti ally mar ine seque nce (Lysogory facies) and a con tine ntal placoderm ser ies (K ielce facies). Plac oderm facies occu r with in the marine seq uence. A new richly fossilifer ous dep osit from the continen tal seq uence ha s yielded abundan t remai ns of acanthodians, ar th rodires, the cr ossopte r yg ian Porolepi s, an d heterostraca ns in particular psam mosteids. Thes e Emsian psarnm os teids show similar ities to the Middle Devon ian Balti c forms. They ap pear mor e lik ely to be the foreru nners of the Middle Devonia n Baltic forms than doss

the Sie gen ia n and Emsian Drepana sp is which is a marine- lagoonal type.'

INTRODUCTION

In the su m mer of 1955 the auth or, as gues t of the Paleontologic al Institute of Wars aw Universi t y , collected a number of specimens of ve r tebr a te - beari ng Emsian san ds tone from a quarry north of Dalesz yc e, near Kielce, in Central Poland (see fig. 1). The fauna was recognised by Dr. E. 1. White of the British Museum (Natural History) as including acan th odians, arthrodires, the crossopterygian Porolepis Woodward, and new heterostracans. The fauna is being investigated by Mr. J. Kulczyc ki of th e Institute of Paleozoology , Polish Academy of Sciences, who has gener ously agreed to allow the au thor to describe the Agnatha. A further collection was made in the su m me r of 1956.

The fossils are preserved as natural moulds in a fine grained san dst one. The grains are well rounded, but the interstices are filled with secondary quartz in optical continuity with the original grains, producing a typical quartzite. There is a high pr op or tion of zircon present as well as some feldspar. As a result of subsequent silicification the most delicate ornamenta tion of the fossil remains is perfectly preserved. The plates, spines, scales and oth er skeletal parts have su ffe red little distortion an d can best be examined from casts (for methods of preparation see Rixon & Meade, 1956). Occasionally, small fragm en ts of original bony material are preserved.

Acta Palaeon t olo glca Polo n lca - vol, II/2-3 15

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NEW OSTRACODERMS FROM LOWER DEVONIAN OF CE N T R AL PO L AND 227

GEOLOGY

The Ernsian sedim en ts of the Holy Cross (Swiety Krzyz) Mountains

inclu de an essen t ially marine seq u ence - the Lysogory facies - and

a continental placoderm series - the Kielce facies . Placoderm-bearing beds occur within the marine seq ue n ce, but the correlation of the two series is uncertain . The Lysogory facies contain marine invertebrates (Czarnocki, 1937), whils t the placoderm series in cl u des on ly vertebrate . and plant rem ains .

The su ccess ion, according to Czarnock i (1937), in the Lysogory region is:

Lower Eifeli an : Bukowa Cora Bed s (cul tri jugatu s horizon ) Upper Emsian: Massive join t ed and Scolithu s Sandston e Middle Emsian: Spirijer Sands ton e

Lower Emsian: Barcza Bed s Siegenian hiatus

Upper Gedinn ian : Klono w Beds

Low er Gedinni an: Rzepin Bed s or Mied ziana Cora Con gl om erat e . The su ccession (Czarn ocki, 1937) in the Kiel ce region is:

Low er Eifeli an Dolomit es Shal es with Haliserit es

Bieli n y Con glomerat es (mo ll uscan fau na) Pla cod erm Sands tone

Con glomer at e

Sieg enian an d Gedinni an hia tus.

Gurich (1896) considered the Placod erm Sandston e to be eq uivalent to the Spiri j er Sandston e i. e. Middle Ems ian ; whilst Sobolev (1909) beli eved it to be tr ans gressive an d probably in clu ding par t of the Middle Devonian as well as the upper part of the Lower Devoni an. Late r Cza rnocki (1937) sug geste d that the presence of Machaeracanthus polon icus Gurich an d Ctena can thus sp. in both the Placod e rm Sandstone

an d the Barcza Beds, indica tes that the former is, at least in part, of

Lower Emsian age. Furth ermore , as the fauna of the Bieliny Conglomera tes is sim ila r to that of the Upper Emsian , the Placoderm San ds tone probabl y rep res ents the whol e,of the Emsian .

The ver te brate fauna of the Placod erm Sandston e was described by Gtirich (1896) as including the following gene ra : Psammosteus , Coccost eu s, Heterostius, Bothriolepis, Ctenacanthus an d Machaera- canthus; no description of this fauna has sin ce been made. Czarnocki

22S L. BEVERLYTARLO

(1937, p. 181) wrote that "it is for the grea test pa r t poorly preserved and has no stratigraphical or paleontol ogic al value. It is rather ra r e an d fragmen tary".

The presence of a vas t amo un t of well preserved vertebra t e remains at Dal es zyc e and also at Debska Wola , 18 km south of Kielce (Czarnocki, 1937, p. 181) ena b les a deta iled stu dy of the fauna of one stra t ig r ap h ical horizon to be undertaken . On the basis of su ch a pal eontolo gic al study it may be possib le in the future to est ablish a vertebra te su ccession in the Kielce fac ies; an d, more im portant, to correla te th is wit h the intercal a te d plac od erm beds in the Lysog6r y facies.

PALEONT OLOGY'

Pr eviousl y the only record of an ostracoderm from the region was of Psammosteu s sp. (Giir ich, 1896 , p. 392) from the Pla cod erm Sands ton e at Now a Huta , 12 km nort h-east of Dal es zyc e. The rec en t collections ha ve yielde d at leas t four forms of psam mosteid and two frag ments of a pter aspid. Altho ug h the latter is specifica lly in determi na te, it is of interest bein g the first record of a pteraspid from the Holy Cr oss Mounta ins. Furthermore, with the ex ception of the Middl e Dev on ia n for ms from Spitsbergen (Dinel ey , 1955), it is the only record of a pte raspi d associa te d with the la ter forms of psammost eid .

The st ra tigraphi cal pos iti on occup ied by the Dal es zy ce psammos teid fauna is give n on p. 229.

From the previousl y supposed absen ce of freshwa te r psammosteids in the Emsian , it has been assu med that the Middle Dev onian Balti c fresh wa ter forms must have been derived from the marine-lagoonal Drepanaspis. This conception was put forward by Obruchev (1944, 1947) an d was bas ed primarily on a cons ideration of the branchial plates :

"Evo lution has tak en plac e fr om Drepanaspi s thro ug h Psammolepi s to Psammost eus by the gradual ch ang ing of the bran ch ial plates so tha t they bec ame shorter an d wider. Togeth er wit h th is the pos terior parts of the branchials wer e red uced" (1947, p. 517, trans lated from the Russian). The differences in bran chial plates were cor re lated with the change from a marine-lagoonal to a fluvi atile habit a t (1944, p. 145).

However,when th e branchials of other Middle Devonia n psammosteid genera are ex ami n ed and addi tion al characte rs i. e. ven t ral median plates, are considered, it is eviden t that there is no sim ple rel ationship be tween Drepana spi s, Psammolepis an d Psam m osteus.

1 This note is concerned only with the Agnatha since the Gna th os tomata are to be desc r ib e d by J. Kulczyc k i.

NEW OSTRACODERMS FROM LOWER DEVONIAN OF CE NTR AL POLAND 229

The discovery of a varied psammosteid fauna at th e end of Lower Devonian times, living in a freshwater environment and showing sim ila r ities with the later Middle Devonian forms, renders unlikely the derivation of the Baltic psammosteids from the marine-lagoonal Drepanaspis. It would ra ther indicate tha t the Daleszy ce psammosteids

MAJOR PSAMMOSTEID FAUNAS

Stag es Continental facies Marine facies

BALTIC

- UPPER Psammost eu s Agas si z

DEVONIAN Dyptychosteus Preobrajensky Karetosteus Obruchev Aspidostells Obruchev

BALTIC

Psammosteus Agassi z MIDDLE Psammolepis Agas siz DEVONIAN pycn osteu s Preobraj ensky

Ganos teusRohon Schizosteus Obruch e v YogHnia Obruchev

RHINELAND EMSIAN DALESZYCE FAUNA Drepanaspis Schlu-

tel'

I

LOWER

SIEGENIAN

DEVONIAN Drepanaspis Schlu-

tel'

PODOLIA

WeigeUaspi s Brotzen GEDINNIAN ANGLO-WALES

WeigeUaspi t: Brotzen

I

were the probable forerunners of the later Middle Devonian forms. The ex isten ce of both fresh water and marine psammosteid faunas in Emsian tim es means that a common ancestor must be sought earlier. The psainmosteids in fact must have developed along two independent lines - .a marine repres ented by Drepanaspis and a freshwater by the Daleszyce and Balti c faunas - from at least pre-S ieg enian times.

230 ^{L. BEVERLYTARLO}

The Daleszyc e psammos teids sho w very distin ct ive orn a m en ta t ion.

but there is also considerable variation sho wn in diff eren t parts ~f the car a pace within individuals.

The most common psammost eid , type A, bears clos ely packe d, large rounded tubercles , each crenulated at the bas e, those on the ridge scales being more elong a te . A fragment of a median plate (see pl. I,-fig. 1) sho ws concentric grow th lines an d short radial grooves on the ex terna l su rfa ce probably representing sensory can als . The bran chial plates appear to show some sim ila ri ti es to Pycn osteus. As no ven t ra l med ia n plates have yet been ide n tif ie d, it has not bee n pos s ibl e to assign th is for m to a gen us.

Typ e B is rep resente d by a fragmen t of a median plate (s ee pl. I, fig. 2 for detail) which has tubercl es sim ilar to those of typ e A. They are, howev e r, separa te d from each other by zones of minute tubercle s, similar in some respects to those occu r r ing in Psammolepis undulata Agassiz and in some specime ns of Pycnos teu s tube rculat us (Rohon).

Typ e C is a bran chial plate which has very large, low tu be r cles well separated from each ot he r as in Ganost eu s stellatus Rohon. Howev e r, in ou t li ne the plate ap proa ches Psammol epi s rat he r than Gan osteus.

The las t psam mos teid, type D, has an orname n t of closely packed, small tu be r cles, but as on ly a few sm all frag me n ts have been collected. no suggestion of possible affi nities can be mad e .

It is hoped that fur t her work on the Dal es zy ce psammosteids will rev eal some con ne ct ion with the Middle Devoni an gene ra, whilst a fu r t he r exami na tio n of the Gedinn ian genera Tesseraspi s and Weigeltaspis ma y throw some light on the earlie r history of the grou p.

At present it is possibl e to sta te that the cen t ra l positi on of Drepanasp is in this history is no lon ger as firmly establis he d as it had ap pea re d.

ACKNOWLEDGE MENTS

The author wishes to ex p ress his deep es t gra t it u de for the ass is t a n ce given throughout by Professor R..Kozlowski and Mr. J. Kulczycki of the Institute of Paleozoology, Polish Academy of Scien ces , and by Dr.

E. 1. White, F. R. S. and Mr. H. A. Toombs of the British Museum (Natural History).

Professor W. Gross of Berlin Univer sity provided facilities for ex am in ing the psammosteids in his care ; Professor D. Obruchev of the Academy of Sciences , Moscow, sen t photographs of the psammosteids in his collections, and Dr.Elga Mark of Tallinn - photographs of Pycnosteus.

To all the above persons the author tenders his sincere thanks.

NEW OSTRACODERMS FROM LOWER DEVONIAN OF CE NT R A L PO L A N D 231

Finally the author gra tef u lly acknowled ges 'the gen erous financial assist a nce from the Percy Slad en Memorial Fund of the British Museum (Natural History) an d from the Polish Academy of Sciences, Warsaw , for the field work during the su m me r of 1956.

Depar t men t of Zoolo gy, Universi t y Colleg e Lon d on

Febnwry , 1957

RE F ERENC ES

BROTZ EN F. 1933. We igelta sp is nov. gen .und die Phyloge n ie der pan ze r t ra ge ride n Heteros tra cf. Cb !. Min. etc., B, 648-656.stuttga r t.

CZAR N OCK I J. 1937. Pr zeglad str a t yg r af ii i pal e oge ogr a fii dewonu dolnego Gor Sw ie t okrzy sk ich, (Uberblick der St rat igrap hie und Pala ogeographie des Unterd ev on s im Pol n ischen Mit te lg ebirge). Spraw. P. In st. Geo!. (Bu l!. Ser vo Geo!. Polo), 8, 4, 129-162, German text 163-200. Warsza wa.

DE N ISO N R. H. 1956. A review of the habitat of the ear lies t vertebrates. Fieldia na (Geol. ), 11,357- 457. Ch icago.

DINE L EY D. L. 1955.Som e De v on ian fish remains fr om North Central Ves tsp its be r - gen. Geo i, Mag., 92, 255-2 60. Lond on.

GR OSS W. 1930.Die F'is ch e des mittleren Old Red Stid -Livlands.GeoloPttl ii ont,Abh., N. F., 18, 121-156. Be rlin.

1933.Die F'ischedes ba'ltischen Dev ons.Pala eon to gr aphi ca,79 A, 1-74.Stut tga rt 1942. Die Fische Faunen des haltischen Dev ons und ih re biostratigra p hische Bed eutung. Korrespblo Natur f. -Ver., 64, 373- 436. Rig a.

1950. Die pal a ontol ogi sche un d stra tigraphlsche Bed e u tung der Wirbel tier - faunen des Old Reds und de r ma r ine n altpaliiozoische n Sch ic h te n. Abh. deutsch. Akad. Wi ss., ma th.-n at. Kl. 1949, 1, 1-130. Berlin .

GtJR I CH G. 1896. De s Palaozoicurn im polnischen Mitte lgeb tr ge . Verh. russ. k: min.

Ges., 32, 1-539. St. Petersburg .

KIE LA N Z. 1954. Les Trilo bi tes mes od e v on ie ns de s Monts de Sa in te-Croix (Trylo - bity srodko w o-dew ori s k ie z Gar Sw ie tokrzys k ich ). Pala eon t . Poi., 6, 1-50. WG- szawa,

MAR K E. 1956. On the ge n us Pycnos te us (Psa rn mos te idae , Agna t ha). (In Rus sian).

Ensv. Teadust e Akad. Geoi, Inst. Uurimu sed , I, 74-88, Tallinn.

OBRUCHEV D. 1940. On some psammos teid s from the Len ing rad and Baltic Midd le Devonian. C. R. Acad. Sci . URSS, 28, 766-768. Moscow.

1943a. Yoglinie n. g. late s t pterasp id from the Middle De v onian of the Leningrad district. Ibidem, 41, 41-43.

1943b. A new reconstruction of Drepana spis, Ibidem, 41, 268-271.

1944. An attempt re storation of Psarnmolepis parad ox a, Ibidem, 42, 143-145 1947. On the gen us Psammosteus (He terostraci ). (In Russian) . Ibidem, 5G, 517-520.

PREOBRAJENSKY 1.A. 1911. On some representative s of the fam ily Psammosteidae (In Russian). Sitzber. Naturi, Ges., 19, 21-36. Dorpat.

RIXONA. E.& MEADE M.J. 1956. Cas ting techn ique s.Mus.Jour n., 56,9-13.London. ROHON J. 1901. Beitrage zur Anatomie und Histologie del' 'Psarnmosteiden. Sit z

k. bohm. Ges. Wiss., math.-nat. Cl., 16, 1-31. 'Prague.

232 ^L.BEVERLY TARLO

SAMSONOWICZ J. 1952. Era paleozoicz na w Polsce. In: M. Ks iqzkiewi cz & J. Samsonowicz, Zarys geologii Polski (An ou tli ne of the geology of Polan d: 51-89.Warszawa .

SOBOLEV D. 1909. The Middle De voni an of the Kiel ce-Sandomier z ra nge (In Russ ian). Mat. Geol, Russlands, 24, 41-536. St. Petersbur g.

THAQUA IH R. H. 1903. The Lower Devon ian fishes of Gemiinden. Trans. Roy . Soc. Edin., 40, 723-739. Edinburgh.

- 1905. Supplement to the Low er Dev oni an fishes of Gemund en. Ibidem , 41.

469-475.

WHITE E. 1. 1935. The ostr acoderrn Ptera spis Kner and the relati onships of the agnatho us vertebra tes. Phi l. Trans. Roy. Soc., B, 225, 381-457. Lond on . 1950. The Vertebrate fau nas of the Lower.Old Red Sand stone of the Welsh Borders . Bull. Brit. Mus. (N at. Hist.), Geo!. 1, 51-6 7. Lond on.

1956.Prelim in a ry noteon the range of Pteraspids in West ernEurope. Bull. Inst. Roy. Sci. nat. Belgiq u e, 32, 10, 1-10. Bru x elles.

WILLS L. J. 1935. Rare and new Ostr acodcrrn fishes fro m the Down toni a n of Shropsh ire. Trans. Roy . Soc. Edin., 58, 427-447. Ed inbur gh.

L. BEV ER LY TAR L O

WIAD OM OSCI WS T F;PNE 0 NOWYCH OSTRAK ODER MA CH Z DOLNEGO DEWONU(E MS U) POLSKI SRODKOWEJ

Streszcz enie

Autor rnial sp osobnos c Iat em 1955 i 1956 r. wyeksploa tow ac wraz z mgr J. Kul- czy ckim obf lte szczatki ryb i ostra koderm6w z piaskowca dolno-de won skiego w Daleszyca ch, zach owane w postaci odcis k6w. Bad anie rn ryb zajql sie J. Kulczy c- ki, a opracowa nie ostrakoder rnow zostalo powier zone autorowi .

Na wste pie pod an a jest kr 6tka cha ra k te rys ty ka ztoza ora z st an u zac how ania skamienialoscl. Nastepnie przytoczono zasadnicze dan e dotyczace stra tygrafii dol- nego de won u G6r Sw ie tokr zys kich, gl6w nie na podsta w ie pracy J. Czarnoc kiego (1937). Rep r odu k owana je s t szki cowa mapk a geologiczna G6r Sw iet okrzysk ich wedlug J. Samson owicza (1952).

W wynik u nastepn ych badan zeb ra nego materialu, autor umies zcza faune da les zycka, zgodnie z Czarnockim, w pietrze ems ki m. Cecha charak ter ys tyczna te j fauny ostrak oderm6w jest domina cja w niej psammost eid6w, kt6 re tutaj zyly w sro dowisku slod kow od ny m. Fakt ten jest 0 tyle ciekawy, ze dot ychczas przy- pus zczano (D. Obruczew , 1944 i 1947), ze slodk ow odne srod kowo-dewons kie psam- most eidy wywodza sie z morsko-Iagunowy ch drepana spid6w. Autor skla nia sie ra- czej ku koncep cji, ze psammos teidy reprezentu ja Iin ie rozwoj ow a nie zalezna od linii drep ana spid6w, juz od ep oki przed zigenem. Pierwsza z tych ilini i byla sled- kowod na , a druga - morska.

Pew ne uwagi poswiec one zostaly charak terystyczny m cec h om ornarnentacy j- nym plytek pancerza. Nal ezy przypus zcza c, ze szczeg olo we bad ania zeb ra ne go ma-

NEW OS T R AC O D E R M S FROM LOWER DEVONIAN OF CE N T H A L PO L AN D 233

terialu, bed a ce W tok u, moga rzucic no we swiatlo na stosunek psamm ost eidow daleszyck ic h do rodza j ow sr od kow o- de wo ns ki ch, Juz obecni e jed na k mozna po- wiedziec, ze pr zypuszczenie 0 cen tra lnej pozycji rodza j u Drep a nasp i s w historii tych be zszczek owcow zostaje zach wia ne.

OBJASNIENIA DO PLANSZY

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PL.

Fig.!. Fragment of psammost eid medi an plate. type. A, show ing orn a me n ta tio n.

gr ow t h lines and sens ory can al s; X 2.

Fig . 2. Deta il of ornamentation of type B; X 10.

Both specimens prese rv e d as na tu r a l moulds.

(Ph o t os W. Brac k enbury)

_..CTA PALAEO N T O LOG ICA POLONICA. VOL. II L. B. TARLO. PL. I

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