THE FAMENNIAN ﬁGOLDEN AGEﬂ OF CONODONTS AND AMMONOIDS...

PALAEONTOLOGIA POLONICA — No. 63, 2006

THE FAMENNIAN “GOLDEN AGE”OF CONODONTS AND AMMONOIDS

IN THE POLISH PART OF THE VARISCAN SEA(Fameński „Złoty Wiek” konodontów i amonitów w polskiej części oceanu waryscyjskiego)

by

JERZY DZIK

(WITH 233 TEXT−FIGURES)

W A R S Z A W A 2 0 0 6

I N S T Y T U T P A L E O B I O L O G I I P A N i m . R O M A N A K O Z Ł O W S K I E G O

GUEST EDITOR

GRZEGORZ RACKI

EDITOR

JERZY DZIKCorresponding Member of the Polish Academy of Sciences

ASSISTANT EDITOR

WOJCIECH MAJEWSKI

Palaeontologia Polonica is a monograph series published by the Institute of Paleobiologyof the Polish Academy of Sciences, associated with the quarterly journal ActaPalae on to logica Polonica. Its format, established in 1929 by Roman Kozłowski, remainsvirtually unchanged. Although conservative in form, Palaeontologia Polonica promotesnew research techniques and methodologies of inference in palaeontology. It is especiallydevoted to publishing data which emphasise both morphologic and time dimensionsof the evolution, that is detailed descriptions of fossils and precise stratigraphic co−ordinates.

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Copyright ©by the Institute of Paleobiology of the Polish Academy of Sciences

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Publishedby the Institute of Paleobiology of the Polish Academy of Sciences

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THE FAMENNIAN “GOLDEN AGE”OF CONODONTS AND AMMONOIDS

IN THE POLISH PART OF THE VARISCAN SEA

JERZY DZIK

Dzik, J. 2006. The Famennian “Golden Age” of cono donts and ammonoids in the Pol ish part of the Variscan sea. Palaeontologia Polonica 63, 1–360.

The stratigraphically com plete and ex tremely fossiliferous geo log i cal sec tions in the HolyCross Moun tains and Sudetes, Po land, cover the whole his tory of the Famennian trop i calhigh−di ver sity pe lagic eco sys tem. Ap pa ra tus re con struc tion of 142 cono dont spe cies al −lowed paleobiological in ter pre ta tion of the fau nal suc ces sion. Three fam i lies, nine gen eraand 39 spe cies are newly pro posed. 76 spe cies of goniatites, with one ge nus and five spe ciesnew, and 70 spe cies of clymenias were also iden ti fied. Like in all other equa to rial lo cal i ties,a sig nif i cant (but not cat a strophic) de cline of di ver sity marks the be gin ning of the Famen −nian. The lo cal pe lagic fauna de vel oped mostly as a re sult of suc ces sive re ap pear ances of lin −eages ear lier oc cur ring in the area but tem po rally re moved from it by en vi ron men tal fac tors.Dur ing the whole Famennian, 101 im mi gra tions of cono dont lin eages are doc u mented. In 31 of the lin eages per sist ing in the area a more or less com plete re cord of their phyletic evo lu −tion is rep re sented; they cover about half (46%) of the sum ma rized ranges of all the lin eages. About half of them are suit able for stratophenetic stud ies. The fos sil re cord of the ammo −noids is much more punc tu ated, but it is es ti mated that 110 lin eages was rep re sented there,only 14 of them pos si bly evolv ing phyletically in the area (sin gle case was stratophenetically proven). At the tran si tion be tween goniatites and clymenias, a suc ces sion within the plexusof closely re lated sympatric spe cies is ob served, but the ex act phyletic change is not re corded and prob a bly all the first clymenias are im mi grants from the east. At least two pro foundrebuildings of the fauna within the Famennian are ob served, but only the ter mi nal De vo nianHangenberg event was of truly dra matic na ture. The newly ac quired ev i dence sup ports theearlier notion thate it is more difficult to trace evolution strato pheneti cally in the equatorialregions than in high latitudes.

Key words: Conodonta, De vo nian, ap pa ra tuses, evo lu tion, Holy Cross Moun tains, Po land.

Jerzy Dzik [[email protected]], Instytut Paleobiologii PAN, Twarda 51/55, PL−00−818Warszawa and Instytut Zoologii Uniwersytetu Warszawskiego, Banacha 2, PL−02−079 War −szawa, Po land.

Re ceived 20 No vem ber 2005, ac cepted 3 No vem ber 2006

CONTENTS

In tro duc tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Ac knowl edge ments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Sam pled geo log i cal sec tions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Gołogłowy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Dzikowiec . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Karczówka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Miedzianka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Ostrówka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Kowala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Kadzielnia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Wietrznia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Jabłonna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Ściegnia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Łagów and Płucki . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Cor re la tion and chro nol ogy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Evo lu tion ary da tums . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Cono dont ecostratigraphy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Meth ods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Cono dont el e ments ex trac tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Re con struc tion of cono dont ap pa ra tuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Tax on omy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25In fer ence on evo lu tion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Cono dont ap pa ra tuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Pat terns of molarization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Growth of el e ments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

De scrip tion and clas si fi ca tion of cono donts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Phy lum Chordata Bateson, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Class Conodonta Eichenberg, 1930 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Or der Prioniodontida Dzik, 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Fam ily Icriodontidae Müller and Müller, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Ge nus Latericriodus Müller, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31Ge nus Icriodus Branson et Mehl, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Ge nus Pelekysgnathus Thomas, 1949 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33Ge nus ?Dollymae Hass, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Fam ily Jablonnodontidae fam. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34Ge nus Mitrellataxis Chauff et Price, 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35Ge nus Jablonnodus gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

Or der Ozarkodinida Dzik, 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40Fam ily Prioniodinidae Bassler, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

Ge nus Ligonodina Bassler, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40Ge nus Pluckidina Dzik, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44Ge nus Lagovidina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Ge nus Idioprioniodus Gunnell, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Ge nus Uncadina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50Ge nus Guizhoudella Wang et Wang, 1978. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Fam ily Gondolellidae Lindström, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52Ge nus Branmehla Hass, 1959. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Fam ily Spathognathodontidae Hass, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56Ge nus Synclydognathus Rexroad et Varker, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . 56Ge nus Apatognathus Branson et Mehl, 1934. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Fam ily Francodinidae fam. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Ge nus Vogelgnathus Norby et Rexroad, 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Ge nus Urbanekodina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64Ge nus Francodina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65Ge nus Sweetodina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67Ge nus Planadina gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

Fam ily Polygnathidae Bassler, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Ge nus Pandorinellina Müller et Müller, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

4 JERZY DZIK

Ge nus Mehlina Youngquist, 1945 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74Ge nus Polynodosus Vorontzova, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77Ge nus Ctenopolygnathus Müller et Müller, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . 84Ge nus Immognathus gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85Ge nus Polygnathus Hinde, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88Ge nus Hemilistrona Chauff et Dombrowski, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . 97Ge nus Neopolygnathus Vorontsova in Barskov et al., 1991 . . . . . . . . . . . . . . . . . . . . . . 102Ge nus Lagovignathus gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104

Fam ily Ancyrognathidae fam. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Ge nus Ancyrognathus Branson et Mehl, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Ge nus Polylophodonta Branson et Mehl, 1934. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

Fam ily Palmatolepididae Sweet, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111Ge nus Klapperilepis Dzik, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111Ge nus Tripodellus Sannemann, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122Ge nus Palmatolepis Ulrich et Bassler, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131Ge nus Conditolepis van den Boogard et Kuhry, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . 138

Fam ily Cavusgnathidae Aus tin et Rhodes, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146Ge nus Alternognathus Ziegler et Sandberg, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . 147Ge nus Scaphignathus Helms, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Ge nus Pseudopolygnathus Branson et Mehl, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

Fam ily Elictognathidae Aus tin et Rhodes, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156Ge nus Pinacognathus Branson et Mehl, 1954 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

Fam ily Idiognathodontidae Har ris et Hollingsworth, 1933 . . . . . . . . . . . . . . . . . . . . . . . . 157Ge nus Protognathodus Ziegler, 1969 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157Ge nus Dasbergina Schäfer, 1976 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

Fau nal dy nam ics of the Famennian cono donts in Po land . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165Suc ces sion of the Famennian cono dont fau nas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165Evo lu tion of the Famennian cono donts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Ammonoid conchs and jaws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186Anaptychi and aptychi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186Growth and func tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188

De scrip tion and clas si fi ca tion of ammonoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189Phy lum Mollusca Linné, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190

Class Cephalopoda Cuvier, 1795 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Sub class Ammonoidea Zittel, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190

Or der Goniatitida Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Suborder Tornoceratina Wedekind, 1918. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190

Fam ily Tornoceratidae von Arthaber, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Ge nus Tornoceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191Ge nus Polonoceras Dybczyński, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197Ge nus Armatites Becker, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201Ge nus Protornoceras Dybczyński, 1913. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201Ge nus Pseudoclymenia Frech, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206Ge nus Ostrovkites gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

Fam ily Posttornoceratidae Bogoslovsky, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208Ge nus Gundolficeras Schindewolf, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208Ge nus Posttornoceras Wedekind, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210Ge nus Discoclymenia Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214Ge nus ?Maeneceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215

Fam ily Cheiloceratidae Frech, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217Ge nus Nehdenites Korn in Korn et Ziegler, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . 217Ge nus Cheiloceras Frech, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219

Fam ily Prolobitidae Wedekind, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224Ge nus Raymondiceras Schindewolf, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224Ge nus Prolobites Karpinsky, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226

Fam ily Dimeroceratidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228Ge nus Dimeroceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228

Fam ily Praeglyphioceratidae Ruzhentcev, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232Ge nus Lagowites Bogoslovsky, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232Ge nus Erfoudites Korn, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233

Fam ily Sporadoceratidae Miller et Fur nish, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234

FAMENNIAN CONO DONTS AND AMMONOIDS 5

Ge nus Felisporadoceras Korn, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234Ge nus Sporadoceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235

Fam ily Prionoceratidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240Ge nus Prionoceras Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240Ge nus Balvia Lange, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243Ge nus Acutimitoceras Librovich, 1957 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245

Or der Clymeniida Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246Fam ily Cymaclymeniidae Hyatt, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

Ge nus Genuclymenia Wedekind, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248Ge nus Cymaclymenia Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249

Fam ily Clymeniidae Ed wards, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254Ge nus Trigonoclymenia Schindewolf, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255Ge nus Clymenia Münster, 1834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256Ge nus Aktuboclymenia Bogoslovsky, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257Ge nus Protoxyclymenia Schindewolf, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259Ge nus Kosmoclymenia Schindewolf, 1949. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259

Fam ily Carinoclymeniidae Bogoslovsky, 1975 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266Ge nus Nanoclymenia Korn, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267Ge nus Cteroclymenia Bogoslovsky, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267Ge nus Carinoclymenia Bogoslovsky, 1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269

Fam ily Costaclymeniidae Schindewolf, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269Ge nus Costaclymenia Schindewolf, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269Ge nus Trochoclymenia Schindewolf, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269

Fam ily Gonioclymeniidae Hyatt, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270Ge nus Finiclymenia Price et Korn, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271Ge nus Kalloclymenia Wedekind, 1914 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272Ge nus Gonioclymenia Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275Ge nus Sphenoclymenia Schindewolf, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275

Fam ily Biloclymeniidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279Ge nus Pachyclymenia Schindewolf, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279Ge nus Kiaclymenia Bogoslovsly, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279Ge nus Biloclymenia Schindewolf, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Fam ily Cyrtoclymeniidae Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281Ge nus Cyrtoclymenia Hyatt, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283Ge nus Platyclymenia Hyatt, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Ge nus Nodosoclymenia Czarnocki, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Ge nus Pleuroclymenia Schindewolf, 1934. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289

Fam ily Hexaclymeniidae Lange, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291Ge nus Praeflexiclymenia Czarnocki, 1989. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292Ge nus Stenoclymenia Lange, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293Ge nus Soliclymenia Schindewolf, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295

Fam ily Glatziellidae Schindewolf, 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297Ge nus Rhiphaeoclymenia Bogoslovsky, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297Ge nus Glatziella Renz, 1914 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299

Fam ily Wocklumeriidae Schindewolf, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300Ge nus Kamptoclymenia Schindewolf, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300Ge nus Parawocklumeria Schindewolf, 1926. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301Ge nus Epiwocklumeria Schindewolf, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301Ge nus Wocklumeria Wedekind, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303

Suc ces sion of the famennian ammonoid fau nas in Po land . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304Evo lu tion of the Famennian ammonoids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305

Goniatites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306Clymenias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311

Fau nal dy nam ics of other Famennian pe lagic an i mals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316Con clu sions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320Ref er ences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321Ta bles 1–19 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331In dex of ge neric and spe cific names. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 352

6 JERZY DZIK

INTRODUCTION

The Famennian is among those ep ochs of the Earth his tory, which ex pe ri enced the deep est trans for ma tion of the liv ing world. Not only its bound aries are un usu ally clear−cut but rev o lu tion ary changes oc curred re −peat edly through out its du ra tion (Walliser 1996). The be gin ning of the Famennian cor re sponds to theworld−wide col lapse of reefal coral−stromatoporoid eco sys tems and the fol low ing pro found re duc tion of bi o −log i cal di ver sity at least in its part rep re sented by the fos sil re cord. Af ter ward, the di ver sity of fos sil as sem −blages of pe lagic or gan isms such as ammonoids and cono donts dra mat i cally in creased, to reach lev els un −prec e dented in ear lier or later his tory of these groups. The end of that acme of pe lagic trop i cal fau nas wassim i lar to what hap pened at its be gin ning, re sult ing in a dras tic re duc tion of diversity to just a few lineages ofammonoids and cono donts.

The events de mar cat ing the Famennian are gen er ally be lieved to be of a cat a strophic na ture and are listedamong the great est ex tinc tion events in the his tory of life (Walliser 1996; but see House 2002 and Bambachet al. 2004). Par a dox i cally, there are rea sons to ques tion not only the im por tance of these events but even thevery pos si bil ity to prove their ex is tence at the world−wide scale (Dzik 1995, 2005). One may thus ask: how isit pos si ble that such ex treme points of view emerge from ex am i na tion of es sen tially the same fos sil ma te rial?The an swer is: all this is rooted in dis pa rate meth od ol o gies of in fer ence. The dis tinc tion be tween a lit eralread ing of the fos sil re cord as it stands and a hum ble rec og ni tion of its lim its, with a long dis tance sep a rat ingthe raw data on fos sils from the knowl edge of ac tual trans for ma tions of an cient eco sys tems. From the sec ondpoint of view there is no cor re spon dence be tween the ob served ap pear ance and dis ap pear ance of a lin eage inrock sec tions and its evo lu tion (Dzik 1995). Fur ther more, there is no nec es sary con nec tion be tween thespeciation and evo lu tion rates (Dzik 1999). If this point of view is ac cepted by the reader (how ever un likely it is), the fos sil re cord ap pears to rep re sent a kind of Pla tonic shadow only roughly cor re spond ing to the realsuc ces sion of events. For tu nately for us, how ever sub tle the cor re spon dence is, it may be spe cific enough toal low rea son able in fer ence. Com ing to spe cific points, the dras tic change in di ver sity ob served in the Lau ren −tian sec tions at the Frasnian–Famennian or Famennian–Tournaisian bound aries in di cates just that the trans −for ma tion of the en vi ron ment in the equa to rial re gions of those times (McGhee 1996; Streel et al. 2000;Joachimski and Buggisch 2002; Joachimski et al. 2001, 2004; Bambach et al. 2004; Racki 2005) forced theorganisms earlier living there to migrate elsewhere or to reduce dramatically their area of distribution.

Whether the re moval of pop u la tions from par tic u lar re gions was con nected with their to tal ex tinc tion ornot, can not be in ferred from the avail able ev i dence. The nu mer ous cases of Laz a rus taxa show that this wasnot the case on many oc ca sions. I hardly see any in tel lec tual or prac ti cal profit from the as sump tion that a dis −con ti nu ity in dis tri bu tion of fos sil spe cies in a sec tion marks the ex tinc tion of its lin eage. It is more prof it ableto con sider this to be just a phe nom e non of lat eral shift in dis tri bu tion of eco sys tems, a lo cal (al though wide −spread) re plac ing of one en vi ron ment (to gether with its or gan isms) by an other. This gives a chance to searchfor the place (or refugium) where the pro cess of evo lu tion pos si bly con tin ued in stead of tak ing the risk of cir −cu lar reasoning resulting from basing correlation on the alleged extinction event.

My main goal is thus to sep a rate evo lu tion from eco log i cally con trolled shifts in oc cur rence. This is notan easy task (Dzik 2005) and even if I am succesful in do ing this, the whole ev i dence pre sented here rep re −sents only a small area of the late De vo nian world. To trace lin eages with rea son able con fi dence not only intime but also in geo graph ical space, sev eral sim i larly de signed works have to be com pleted in other re gionsof the World with sim i lar fau nas. One has to hope that we will not wait too long.

In fact, the Famennian is es pe cially suit able for these kind of stud ies. It was an ep och of rel a tively highsea level and pe lagic fau nas of that age are rel a tively wide spread and well sam pled. The ex po sures of the De −vo nian along mar gins of the Variscan orogenic belt in cen tral Eu rope (Fig. 1) are by no means unique in thisre spect. Their spe cial value is only in that these are the clas sic re gions of the paleontological and bio −stratigraphical re search on fos sils of pe lagic Famennian or gan isms. The Sudetes are among the first rec og −nized lo cal i ties of De vo nian ammonoids in the world (von Buch 1839 and nu mer ous pub li ca tions by otherau thors re ferred to in the ammonoid part of this work). Even in the Holy Cross Moun tains, lo cated on the dis −tant east ern tip of the belt, the first De vo nian cono donts were iden ti fied al ready by Gürich (1901). Mono −graphic de scrip tions of the Famennian cono donts from Po land started with Wolska (1967), then with sub se −quent work by Szulczewski (1971), sup ple mented by Nehring (1967), Narkiewicz (1978; Narkiewicz andNarkiewicz 1992), Baliński (1979), and Matyja and Narkiewicz (1995). The ammonoids from the Holy


Cross Moun tains were monographed by Sobolew (1914a; sup ple mented by Dybczyński 1913 and Makowski 1991) and Czarnocki (1989; Pajchlowa et al. 2003). The pres ent pa per is in tended to be an over view of theevolution and dynamics of pelagic faunas in the area.

Il lus trated spec i mens are housed at the In sti tute of Paleobiology of the Pol ish Acad emy of Sci ences inWar saw (ab bre vi ated ZPAL), State Geo log i cal In sti tute in War saw (IG), Mu seum of Nat u ral His tory of theHumboldt Uni ver sity in Berlin (MB), Geo log i cal Mu seum of the Wrocław Uni ver sity (UW), the Nat u ralHis tory Mu seum of the Ukrai nian Acad emy of Sci ences in Lvov (for mer Dzieduszyckis’ Mu seum; MD), and Mu seum of the In sti tute of Ge ol ogy and Pa le on tol ogy of the Tübingen Uni ver sity (UTü).

Ac knowl edge ments. — I am very thank ful to Gilbert Klapper (The Uni ver sity of Iowa), who read themanu script of the cono dont part of this pa per, cor rected nu mer ous er rors and mis takes in the orig i nal ver sionof the text and im proved its Eng lish style. Di eter Korn (Mu seum für Naturkunde, Berlin) and an anon y mousre viewer of fered nu mer ous im prove ments to the ammonoid part, al though accomodation to some of his/herre quests ex ceeded my abil i ties and the pa per re mains de fi cient in many as pects. Grzegorz Racki in tro ducedme to the ge ol ogy of the Holy Cross Moun tains. Cono dont el e ments from sev eral sam ples have been pickedin War saw by Ewa Hara. Most of the pho to graphs of ammonoids were taken by Grażyna Dziewińska. Pub li −ca tion costs were par tially cov ered by the Pol ish Min is try of Sci ence and Informatization (grant No. 2 P04D001 28).

SAMPLED GEOLOGICAL SECTIONS

There are only three ar eas in Po land where the De vo nian rocks can be sam pled in ex po sures: the HolyCross Moun tains in cen tral Po land about 200 km south of War saw, a small area in south ern Po land west ofKraków, and the Sudetes on the south west ern tip of the coun try. Those ex po sures (Fig. 2, 3) are briefly re −

8 JERZY DZIK

Fig. 1. Non−palinspastic palaeogeographic map of cen tral Eu rope in the Famennian (for sources see Dzik 1997, fig. 1). Knownex tent of the lat est De vo nian to ear li est Car bon if er ous lime stone fa cies shown by brick pat tern, cephalopod lime stone as fineshal lower fa cies as coarser; ar eas of in tense sub si dence, where coal bas ins de vel oped in the Car bon if er ous, ver ti cally hatchured;

ex tent of deeper−wa ter (mainly fine clastic) sed i men ta tion shown as grey.

viewed be low start ing with those of the Sudetes, near the south west ern shore of the Variscan sea, on the op −po site side of it in re spect to other regions studied.

GOŁOGŁOWY

The sec tion at Gołogłowy near Kłodzko in the Sudetes is lo cated about 1 km north of the vil lage on theright side and above the aban doned quarry, 40 m upslope the for est road. It was de scribed and il lus trated byHaydukiewicz (1981), who re ported early Tournaisian cono donts from a cal car e ous mudstone near the top of the sec tion. I sam pled the well−bed ded Famennian part of the succesion. Most of sam ples con tained rare andstrongly tec toni cally de formed cono donts. They ap par ently rep re sent a transgressive succesion from a shal −low−wa ter fauna dom i nated by Polygnathus znepolensis and P. extralobatus in the nod u lar lime stone be low(sam ple Goł−11) to an as sem blage rich in Palmatolepis rugosa and Tripodellus gracilis near the top of theDe vo nian bed ded muddy lime stone (sam ple Goł−3). A sim i lar, but more com plete suc ces sion is rep re sentedin the same area, 25 km northward, at Dzikowiec.


Fig. 2. The sam pled sec tions of the early Famennian in the Holy Cross Moun tains, with their rel a tive po si tion shown on a di a −gram matic map of the ex tent of ex po sures of the De vo nian. Po si tion of zonal bound aries and their cor re la tion in di cated by bro −ken lines. Com pos ite sec tion of Wietrznia is as sem bled from tec toni cally disjunct ex po sures; their lo ca tion in di cated on a map of

the quarry.

10 JERZY DZIK

Fig. 3. The sam pled sec tions of the late Famennian in the Sudetes and the Holy Cross Moun tains, rel a tive po si tion of the lat tershown on a di a gram matic map of the ex tent of ex po sures of the De vo nian. Po si tion of zonal bound aries and their cor re la tion in di −

cated by bro ken lines. Lo ca tion of sec tions in the Wapnica quarry in Dzikowiec shown on a map.

DZIKOWIEC

The ge ol ogy of the De vo nian at Dzikowiec has been re cently re viewed in Berkowski (2002) and the Car −bon if er ous by my self (Dzik 1997). The suc ces sion starts there with a con glom er ate, com posed of largeblocks of gab bro with a lime stone crust of prob a bly al gal or i gin, a de tri tal lime stone with crys tal line andmeta mor phic rock peb bles higher up, and the cephalopod Wocklumeria and Gattendorfia Stufen lime stonesnear the top. In the dark grey organodetrital lime stone, rich in cal car e ous tests of foraminifers and al gal de tri −tus, col o nies of var i ous tab u late (Syringopora) and rugose cor als oc cur rep re sent ing a rare case of theFamennian refugium with reefal or gan isms (Berkowski 2002). The only cono dont spe cies en coun tered in the mid dle part of this unit is Polygnathus znepolensis (sam ple Dz−11, 50) sup ple mented up wards by a prio −niodinid (sam ple Dz−51) and a few more spe cies (Dz−52), in clud ing Dasbergina micropunctata, Mehlina,Branmehla, and Tripodellus gra cilis. In a sam ple taken from the base of the up per, nod u lar part of the mainlime stone (Dz−53), a few el e ments of the Polygnathus extralobatus ap pa ra tus have been found, which makesthis part of the suc ces sion sim i lar to that at Gołogłowy. The dark grey nod u lar lime stone be comes more platyup wards and con tains (Dz−16, Dz−62) the shal low−wa ter spe cies P. znepolensis as so ci ated with rare T.gracilis, Palmatolepis rugosa, and non−plat form cono donts. Sam ples from the tran si tional strata be tween thedark grey and red cephalopod lime stone (Dz−69, Dz−72) yielded a more di verse palmatolepidid fauna. Thetran si tion from Dasbergina marburgensis to D. trigonica takes place 0.4 m be low the top of the redcephalopod lime stone of the Wocklumeria Stufe.

The Famenian was sam pled for cono donts near the north ern tip of the quarry (Berkowski’s lo cal ity 3),where the Gattendorfia Lime stone suc ces sion is the most com plete (Dzik 1997) but the base of theWocklumeria Lime stone and the top of the dark grey nod u lar lime stone are there trun cated by a fault. A con −ti nu ity be tween these units can be traced in a rock in the west ern wall of the north ern part of the quarry(Berkowski’s lo cal ity 3). Sam ples were taken also from a rock im me di ately south of the steps lead ing out side the quarry (at the fence of the shoot ing field used for training by local hunters).

A bed−by−bed col lect ing of ammonoids was per formed by Lewowicki (1959) who de scribed in de tailtheir dis tri bu tion. Wocklumeria sphaeroides has not yet been found there. Of spe cial in ter est is the oc cur −rence of Kamptoclymenia endogona in the top most layer of the Famennian, and Kalloclymenia through outthe ex posed cephalopod lime stone up to about 0.5 m be low its top. Glatziella ranges from 1.2 to about 0.2 mbe low the top. Korn et al. (2005) iden ti fied Soliclymenia in the red lime stone 0.4–0.8 be low the top of thesuc ces sion and sug gested that a sig nif i cant part of the lat est Famennian is not rep re sented there. I at temptedalso to col lect ammonoids from these ex po sures but re sults were not es pe cially ex cit ing and most spec i menscome from the scree. A nice fauna with Balvia was col lected from the scree near the ex ploi ta tion cham bers atthe south ern end of the quarry. Most of the Famennian is there hard to ac cess in the ver ti cal rock wall, butamong loose blocks a highly fossiliferous dark grey rel a tively pure lime stone with nu mer ous bi valves is rep −re sented, yield ing the lat est Famennian Protognathodus fauna, otherwise unknown from the quarry.

KARCZÓWKA

In the col lec tion of Dymitr Sobolew housed at the Mu seum of Nat u ral His tory in Kharkov, Ukraine,blocks of cephalopod lime stone from the Karczówka hill south of Kielce are rep re sented. Ev i dently this is the same ma te rial as that re ported by Sobolew (1911). I was al lowed to pro cess for cono donts a small piece ofthis lime stone which yields an as sem blage of a rather un usual com po si tion for the early Famennian of theHoly Cross Moun tains, with or nate sim ple cones of Mitrellataxis. Prob a bly this was a shal low−wa ter en vi −ron ment above the top of a stromatoporoid−coral Frasnian buildup. Such palaeocological con text is typ i calalso for other sec tions of the Famennian in the area.

MIEDZIANKA

A few metres−wide belt of ex po sures of the Famennian ex tends along the south ern slope of the hillMiedzianka near Chęciny (Fig. 4). Ap par ently, the some what marly and thin bed ded rocks of this unit weremore prone of dis lo ca tion than the mas sive mudmound and la goonal de pos its of the un der ly ing Frasnian andGivetian. The cop per ore min er al iza tion within the faults was the rea son for in tense quar ry ing since the 18th

cen tury. The com plex tec ton ics makes stra tig ra phy there dif fi cult to es tab lish and only two sec tions in this


zone have ap peared suit able for study, lo cated on op po site ends of it. In the an cient open mine on the SW ofthe hill rem nants of two min ing gal ler ies are pre served. In tec tonic blocks hang ing above the en trance to theeast ern gal lery and on its east ern wall in dis tinct bed ding is en hanced by weath er ing en abling sam pling forstrati graphic pur poses. Succesion is there reversed, with younger strata downhill.

Fossiliferous strata of the ter mi nal Frasnian, with a brachi o pod co quina with atrypid brachi o pods,ahermatypic cor als, and cepha lo pods are ex posed in front of the en trance to the gal lery on the north ern wallof the mine (men tioned in Szulczewski 1989). Its age was de ter mined by Czarnocki and Samsonowicz(1911) based on find ings of Manticoceras. Si li ceous lithistid sponges and si lici fied goniatite shells oc cur inthe top most part of the co quina, mak ing the suc ces sion sim i lar to that at Kowala (de scribed be low). The most un usual as pect of the cono dont fauna is the com plete lack of Icriodus, so abun dant in the ter mi nal Frasnianstrata else where, and abun dance of Manticolepis rhenana. M. winchelli and “Palmatolepis” linguiformis areas so ci ated but Lagovilepis bogartensis and Klapperilepis ul tima are miss ing. The en vi ron men tal sit u a tion isthus some what sim i lar to that at Kowala but fun da men tally dif fer ent from Płucki (see Dzik 2002). A spec −trum of main cono dont taxa for the suc ces sion was al ready given by Szulczewski (1989). My sam pling re −vealed that there is no con ti nu ity be tween the Frasnian and Famennian here. The co quina is topped with acon glom er ate (sam ple Md−27, Ta ble 3), com posed mostly of peb bles of a light grey lime stone dis sim i lar tothat im me di ately be low in the sec tion. The con glom er ate ma trix yielded an as sem blage of the lat est K. trian −gularis Zone spe cies with a rather in sig nif i cant ad mix ture of the lat est Frasnian spe cies. The lat ter is rathersur pris ing, con sid er ing a rather high pro duc tiv ity of cono dont sam ples from the co quina. Pos si bly, the peb −bles were de rived ei ther from a not so high part of the Frasnian or they rep re sent de stroyed strata de pos ited atthe be gin ning of the Famennian. There was thus a sed i men tary dis con ti nu ity con nected with an ero sion event nearby mark ing a sea level drop in the ear li est Famennian (John son et al. 1985 ).

A red dish organodetrital lime stone ini ti ates the well−bed ded part of the suc ces sion. Beds of a cephalopodlime stone oc curred there once, from which Czarnocki and Samsonowicz (1911) ex tracted Nehdenitesverneuili. A cono dont as sem blage with Klapperilepis crepida has been re cov ered by my self from looseblocks and sam ples from a tec toni cally brecciated zone in the ex po sure. A seem ingly or dered succesion cov −ers the C. marginifera and L. styriacus zones but cono donts show that it is re peated there. The youn gest bedof the se ries near the dis lo ca tion in the mid dle of gal lery is prob a bly of early L. styriacus Zone age, whereasthe south ern block on the hill slope out side the mine ex tends to the P. jugosus Zone. The pat tern of dis lo ca −

12 JERZY DZIK

Fig. 4. Lo ca tion of the Famennian ex po sures at Miedzianka in the Holy Cross Moun tains (A) and places from where sam pleswere taken (B); view from west.

tions is so com plex that the ac tual suc ces sion can be re stored only ten ta tively and a prob a bil ity re mains thatbeds of dif fer ent age are in ter ca lated. As typ i cal for the lo ca tion, sam ples taken from ap par ently the same bed (in di cated with con sec u tive lettering in Tables) may appear to be of different age.

The late Famennian is doc u mented in the aban doned quarry at the SE foot of the hill. This is a marly shalewith cal car e ous con cre tions tec toni cally in serted in the Frasnian mas sive lime stone (Czarnocki and Sam so −no wicz 1911). Among con cre tions are nu clei of clymenias (Cymaclymenia). Czarnocki (1928, 1989) de ter −mined Wocklumeria sphaeroides, Parawocklumeria paradoxa, and sev eral other spe cies di ag nos tic of thelat est Famennian. Ac cord ing to Czarnocki and Sujkowski (1931), Tournaisian black shales with phosphoritecon cre tions and Guerichia once crop ped out above the marly shale of the Famennian.

OSTRÓWKA

Ex po sures of the Famennian on the north ern wall of the Ostrówka quarry are lo cated only 3 km east ofMiedzianka, but their fa cies de vel op ment and strati graphic com plete ness is dra mat i cally dif fer ent, as iden ti −fied al ready by Czarnocki (1928). At pres ent only a few beds of ex tremely con densed Famennian lime stonecan be found top ping the Frasnian (Racki 1993) lime stone with Amphipora, with an ero sional dis con ti nu ity(Szulczewski et al. 1996). In ex treme cases only two beds of the Famennian lime stone are rep re sented, thelower one rep re sent ing the P. trachytera Zone, the up per one D. trigonica Zone (up per velifer and costatuszones; Szulczewski 1978). I sam pled the most com plete sec tion among ac ces si ble ones in front of theTodowa Grząba hill, from where also slabs with clymenoid ammonoids were col lected. The mas sive lime −stone un der ly ing the organodetrital cephalopod lime stone suc ces sion yielded a cono dont as sem blage (sam −ple Ost−0) with Conditolepis marginifera, the dom i nant spe cies be ing Icriodus cornutus. Cono donts of the C. marginifera Zone oc cur even be low, pos si bly the source was a pocket within the lime stone (Szulczewski etal. 1996) al though no such struc tures were dis cern ible in the sam ple. The first bed of black cephalopod lime −stone (sam ple Ost−1) con tains Palmatolepis trachytera with re duced pos te rior pro cess to gether with Lago vi −gnathus styriacus; it is thus sig nif i cantly youn ger than the cono dont−yield ing strata at Łagów−Dule de scribedbe low. The bed im me di ately above (Ost−5) yielded Palmatolepis rugosa and Tripodellus mancus. A thin de −tri tal lime stone layer top ping the suc ces sion yielded Dasbergina trigonica in di cat ing the lat est Famennian.Above, a marly shale with nod u lar lime stone beds and con cre tions fol low, with the Scalio gnathus anchoralisfauna and prob a bly re worked Siphonodella (Szulczewski et al. 1996; Dzik 1997).

Be fore in tense quar ry ing a much more com plete suc ces sion of the Famennian was ex posed in a trenchdug by Jan Czarnocki (1989, p. 23). An other sec tion, only 1.8 m thick has been sam pled for trilobites byOsmólska (1962). Wolska (1967) in her work on cono donts re ferred to Osmólska’s sec tion but it is un likelythat her sam ples truly were taken from it. Bed 2 of Osmólska (1962) is a black bi tu mi nous lime stone ev i −dently cor re spond ing to beds 3 to 5 of Czarnocki (1989), the most fossiliferous clymeniid ho ri zon ac ces si blenow, with Stenoclymenia sandbergeri. Among cono donts from that lime stone (e.g. my sam ple Ost−12) thereis P. trachytera (found in bed 7 by Wolska 1967) and Lagovignathus styriacus (bed 8). Wolska (1967) re −ported C. marginifera from strata 4 to 7, beds 1–3 be ing poor in cono donts. This makes data of Wolska of lit −tle use in cor re lat ing the suc ces sion of clymeniids de scribed by Czarnocki (1989; see also Woroncowa− Marcinowska in Pajchlowa et al. 2003) with the cono dont scheme, de spite ef forts of Szulczewski andŻakowa (1967). I dis solved small pieces of ma trix from sev eral of Czarnocki’s spec i mens of clymeniidshoused at the Mu seum of the State Geo log i cal In sti tute in War saw (ear lier done also by Woroncowa− Marcinowska 2003 and Woroncowa−Marcinowska and Szrek 2004 and an other spec i men from the col lec tion of the Instytut Paleobiologii PAN. This al lows de ter mi na tion of the pos si ble age of some of the beds. Sam pletaken from ammonoid spec i men ZPAL AmVII/185 (Ost−185 on Ta ble 17) con tains Tripodellus gonio cly me −niae, from IG 284.II.265 and IG 284.II.293 (Ost 265 and Ost−293) Palmatolepis rugosa has been ex tracted.

KOWALA

The stud ied ma te rial co mes from the quarry of the ce ment plant Nowiny II lo cated im me di ately south ofthe vil lage Kowala, in prox im ity to the rail road sec tion stud ied by Szulczewski (1971). This is the most com −plete and thick est sec tion of the Famennian in the Holy Cross Moun tains. Cono donts and ammonoids fromboth its Frasnian and Tournaisian parts have been al ready de scribed by my self (Dzik 1997, 2002). Cono donts


oc cur through out the whole sec tion but sam ples are usu ally of low pro duc tiv ity and spec i mens are cov eredwith clay. Be cause of lit tle use of this ma te rial to ap pa ra tus re con struc tion I col lected rel a tively few sam ples.The sec tion has been stud ied stratigraphically also by Berkowski (2002).

The base of the Famennian is lo cated within a unit of chert−bear ing lime stone with a wavy bed ding, and isnot dis cern ible lithologically (Racki and Baliński 1998; Dzik 2002; Racki et al. 2002; Bond and Zatoń 2003).5.5 m above, the lime stone grad u ally be comes lam i nated and marly with bed ding planes cov ered by largethalloid al gae of pos si bly codiacean af fin i ties (Racki et al. 2002, fig. 3b), Concavicaris crus ta ceans andammonoid anaptychi. In sam ple Ko−155 from the base of this unit Klapperilepis ul tima is al ready as so ci atedwith Tripodellus clarki (or Klapperilepis schuelkei) and a few other spe cies of Klapperilepis. 13 m above thebase the first Conditolepis prima was en coun tered, and K. rhomboidea about 24 m from the base. Faults cut thesec tion in this part and it re mains dif fi cult to es ti mate how much of it is miss ing or du pli cated, but a sam pletaken about 2 m above the fault con tains still ap prox i mately the same fauna with K. rhomboidea.

An unlabelled wooden box with fossiliferous marls sim i lar to those from Kowala has been trans ferred tome by Zofia Kielan−Jaworowska, col lected by her in 1946 for the late Jan Czarnocki. Among fos sils there is a poorly pre served spec i men pos si bly rep re sent ing Cheiloceras lagoviense and the nautiloid Gonatocyrtocerascf. guerichi. If truly col lected at Kowala, it prob a bly co mes from near the fault, as in di cated by cono donts(sam ple Ko?−168; Ta ble 8).

A suc ces sion of about 93 m of marl with con cre tions and lime stone in ter ca la tions con tin ues from abovethe fault to the top of the Famennian and of fers an un usu ally com plete suc ces sion of cono donts andammonoids. Conditolepis quadrantinodosa is pres ent from at least 80 m be low the top (sam ple Ko−161); C.marginifera about 7 m higher (Ko−163). In the black shale 63–65 m be low the top of the sec tion, py ritic nu −clei of Protornoceras and as so ci ated ammonoids oc cur; the fauna was known ear lier only as re worked inQua ter nary clay at Sieklucki’s brickpit in Kielce (Dybczyński 1913, Sobolew 1912a, 1914a, b; House 1970).The as so ci ated cono dont as sem blage in cludes Conditolepis distorta and Palmatolepis ampla.

The lat ter spe cies is re placed (sam ple Ko−18) by Palmatolepis trachytera 40 m be low the top and about 4m be low a very char ac ter is tic in ter ca la tion of a card board−like black shale with Guerichia, punc tured but notbreak ing when hit by a ham mer. This li thol ogy marks the Platyclymenia annulata event (see Bond and Zatoń 2003). Lime stone nod ules im me di ately above the shale con tain very well pre served ju ve nile cono dont spec i −mens (sam ple Ko−8a).

An other mem ber of the P. trachytera lin eage, P. rugosa, emerges 26 m be low the top of the Famennian(sam ple Ko−181), as so ci ated there with L. styriacus. 4 m higher, Pseudopolygnathus jugosus (sam pleKo−131, Ko−194) ap pears with in creased con tri bu tion of lime stone in ter ca la tions first. These higher Famen −nian strata, ex posed now in the north ern wall of the quarry, were iden ti fied by Czarnocki (1933) in trenchesdug at this place. His sec tion ranged from the green ish nod u lar marly lime stone in ter ca lat ing with shales con −tain ing an ammonoid as sem blage with Clymenia. Calcitic skel e tons of ben thic or gan isms are rep re sentedthere by rhabdomesid bryozoans and platyceratid gastropods.

A black shale in ter ca la tion splits a 1.5 m thick nod u lar green ish lime stone above the more marly lay ersinto halves. The lime stone is topped with a thicker bed where Dasbergina trigonica first appears (sam pleKo−123). This part of the sec tion shows thus a rather low sed i men ta tion rate as com pared with that rep re sent −ing the K. triangularis to P. trachytera zones.

6.2 m of marl with lime stone nod ules fol low then with al ter na tion of ol ive green and red lay ers. This unitap par ently cor re sponds to the most fossiliferous part of trenches yield ing the di verse and well−pre servedclymeniid as sem blage with Kalloclymenia de scribed by Czarnocki (1989). Un for tu nately, in the trench sam −pled by my self (Dzik 1997) only nu clei of ammonoid conchs de void of shells were found. All spec i mens col −lected from these beds be long to stratigraphically undiagnostic spe cies of Prionoceras and Sporadoceras.

Above the last red nod u lar bed a 3.4 m thick succesion of ol ive shale and marls fol lows with rare nod u larlime stone in ter ca la tions and two beds of light green ish clay (pos si bly ben ton ite). Wocklumeria sphaeroidesand Parawocklumeria paradoxa oc cur there. Two beds of nod u lar lime stone, each about 0.3 m thick yieldedthe last Famennian−type cono donts, Tripodellus gracilis, and Branmehla suprema (sam ple Ko−75; Dzik1997). Above is an other black, burn ing shale 1.1 m thick (re ported al ready by Czarnocki 1933, weath ered inthe trench stud ied by my self but now ex posed in the quarry; Filipiak and Racki 2005) and 1 m thick bed oftuffite. The tuffite yielded the Acutimitoceras fauna of the ter mi nal Famennian, which is tax o nom i cally im −pov er ished but rich in spec i mens. Sim i larly im pov er ished cono dont Protognathodus as sem blage was en −

14 JERZY DZIK

coun tered in a lam i nated lime stone in ter ca la tion 2 m above the tuffite and 2 m be low the first oc cur rence ofdi ag nos tic Tournaisian cono donts (Dzik 1997). The Kowala sec tion ap pears thus prob a bly the most com plete and fossiliferous among known succesions across the Devonian–Carboniferous boundary.

A bore hole has been drilled in the vi cin ity of the quarry, de scribed by Romanek and Rup (1990) andŻakowa and Radlicz (1990). Cono donts from the core were stud ied by Nehring−Lefeld (1990). The Fras −nian–Fammenian bound ary was in cor rectly de ter mined as co in cid ing with the top of mas sive lime stone. Thelime stone strata above has yielded at depth 237.7 to 236.7 m spec i mens of Belodella (Nehring−Lefeld 1990,pl. 4: 12–13), the lin eage ter mi nat ing its oc cur rence with the end of the Frasnian of the Holy Cross Moun −tains. Ap par ently, sim i larly as in the out crop (Racki and Baliński 1998; Dzik 2002), the bound ary is lo catedclose to the top of the somewhat silicified limestone.

KADZIELNIA

The cephalopod lime stone at Kadzielnia has been de scribed palaeontologically for the first time by Gürich(1896), and Sobolew (1912a) de ter mined its Famennian age. Wolska (1967) and Szulczewski (1971) sam pledit for cono donts iden ti fy ing the crepida to quadrantinodosa zones. Makowski (in Szulczewski 1971, p. 67) iden −ti fied Tornoceras sublentiforme (his T. acutum) among cepha lo pods of the lime stone. Nautiloids from this bedwere de scribed by my self (Dzik 1984). The Cheiloceras lime stone rests at Kadzielnia im me di ately on the mas −sive reefal Frasnian lime stone and its basal bed con tains a shal low−wa ter cono dont as sem blage with dom i nantIcriodus cornutus and Polygnathus praecursor. The palmatolepidids emerge in great num ber in the main bed of the cephalopod lime stone with ran domly dis trib uted large conchs of nautiloids and goniatites. The pres ence ofad vanced Klapperilepis ter mini and Conditolepis prima in di cates the late K. crepida Zone.

A sig nif i cant fau nal change is ex pressed in the up per part of the over ly ing bed of pure lime stone (sam pleKa−3). Many spe cies ter mi nate their oc cur rence (in clud ing Klapperilepis crepida), whereas K. rhomboidea,Conditolepis glabra, C. falcata, and C. klapperi emerge. The change is so abrupt that a sed i men tary dis con ti −nu ity with sig nif i cant gap in de po si tion seems likely. These spe cies con tinue to the end of the sam pled lime −stone suc ces sion. Above is a marly unit with nod u lar lime stone in ter ca la tions yield ing blind trilobitesDianops and poorly preserved cephalopods.

WIETRZNIA

The aban doned Wietrznia quarry in Kielce (now a na ture sanc tu ary) is among the Holy Cross Moun tainssec tions of the Frasnian most pro duc tive in cono donts (re viewed in Dzik 2002). The Famennian ex po suresthere are less suit able for strati graphic stud ies be cause of in tense tec ton ics. Be hind the north ern wall of thequarry there is a dis lo ca tion zone with al most ver ti cally dip ping Famennian marls of var i ous age in con tactwith units of the Frasnian, dif fer ent in each ex po sure. Szulczewski (1989) in ter preted the tec ton ics assynsedimentary. Al though this seems pos si ble as a re sult of break ing and dif fer en tial set tling of the un der ly −ing Frasnian reefal or mud mound body, the ev i dence for this re mains weak. More likely, the Famennian pe −lagic strata in var i ous out crops rep re sent tec toni cally shuffled different parts of the same lithologic column.

Among cono dont sam ples avail able to my self, there is no ma te rial rep re sent ing the ear li est Famennian. Thegeo log i cally old est one is Wtr−34 taken from a tec tonic brec cia im me di ately be low the large block left in themid dle of the quarry. The cono dont as sem blage from there in cludes Tripodellus schuelkei with a wide an gu larplat form. Prob a bly the same brec cia is ex posed in the wall of the quarry nearby (block A? of Szulczewski1989). It is topped with bed ded lime stone con tain ing T. clarki (Wtr−27) as so ci ated with the first Conditolepis.The lime stone suc ces sion with Klapperilepis robusta (Wtr−26) ends there with a sed i men tary dis con ti nu ity pen −e trated with bur rows. A suc ces sion of marls with nod u lar lime stone in ter ca la tions starts from there. Early K.ter mini in the basalmost layer (Wtr−25) in di cates some time gap at the dis con ti nu ity.

The first Famennian bed top ping the Frasnian suc ces sion pub lished in Dzik (2002) is prob a bly of some −what youn ger age, as sug gested by ad vanced T. variabilis. The next por tion of the Famennian has been sam −pled in the dis lo cated block in the mid dle of the quarry, where a se ries of lam i nated dark lime stone and cal −car e ous shale with pyritized radio lar ians and phosphatized car a paces and ap pend ages of the pe lagic crus ta −cean Concavicaris oc curs. In a sam ple taken at the base of the block (Wtr−32), K. crepida has been iden ti fied; in a sam ple from lime stone in ter ca la tion near the top of the ex posed suc ces sion (Wtr−33), Conditolepis prima


has been en coun tered. Nod ules from the same unit brecciated in the wall east of the block have yieldedpoorly pre served goniatites.

JABŁONNA

This is per haps the most in tensely stud ied sec tion of the Famennian in the Holy Cross Moun tains. Un for −tu nately, it is also prob a bly among the most tec toni cally dis turbed. A trench has been dug there in 1949 byCzarnocki (1989, fig. 8) and ammonoids have been col lected bed−by−bed. Some of the clymeniids have beende scribed by Czarnocki (1989) and trilobites by Osmólska (1962). She de scribed the sec tion and cor rectlyde ter mined the age of bed 9 as cor re spond ing to the Cheiloceras Stufe but her re port of Clymenia (Cyma −clymenia in Czarnocki 1989) in beds 14 and 18, the lat ter sug gested to rep re sent the Wocklumeria Stufe, re −mains a mys tery. The ma te rial of un pre pared goniatites and clymeniids from the trench has been trans ferredto my self in 1972 by Halszka Osmólska and cono donts have been ex tracted from the ma trix. They in di catethe K. crepida Zone ex tend ing from beds (my cono dont sam ples J−) 3 to 16, ma te rial from beds 18–19 is notrep re sented, the P. trachytera Zone is pres ent in beds 20 to 24 (where also the first clymeniids are rep re −sented). The L. styriacus Zone is pres ent in bed 26, and the P. jugosus Zone in bed 27. Higher strata are notrep re sented among cono dont sam ples, but bed 33 yielded Kosmoclymenia and Epiwocklumeria. As shownon the pub lished sketch (Czarnocki 1989) much of the suc ces sion is rep re sented by rub ble and this may bethe rea son why bed 23, con tain ing C. marginifera, does not fit in stratigraphically.

In 1962 an other trench has been dug in ap prox i mately the same place and cono donts from there were de −scribed by Wolska (1967). Un for tu nately, the strata ex posed in the new trench have not been cor re lated withthose from the old one. The first Conditolepis prima (Palmatolepis glabra glabra of Wolska 1967) has beenen coun tered in bed 15 (at tri bu tion of spec i men on her pl. 7:12 to bed 9 is prob a bly a mis take), so the strataabove prob a bly cor re spond to bed 16 of Czarnocki. I sam pled the re mains of the trench in 1980 and the rep re −sen ta tion of the first oc cur rence of C. prima in con tin u ous suc ces sion has been con firmed (sam ple J−46).Synclydognathus ancestralis, the spe cies known else where only from sam ple Wtr−32 at Wietrznia and pre −sum ably of very re stricted oc cur rence, has been found in sam ple J−45a of my new se ries and in sam ple frombed 16, which seems to pro vide a rather firm ground for cor re la tion. In fact this fits well also the dis tri bu tionof C. tenuipunctata in both series.

Higher sam ples of Wolska were taken from the rub ble and their spa tial re la tion ship in the sec tion can notbe es tab lished. They cor re spond to the part of Czarnocki’s trench above the dis lo ca tion. Bed 27 of Wolska(1967) cor re sponds to bed 20 of Czarnocki (1989). It ap pears thus that a sig nif i cant part of the midFamennian is not rep re sented in the trenches.

A fun da men tally dif fer ent pic ture is pre sented by the core of bore hole Jabłonna IG 1 (Żakowa et al.1983). Al though highly in com plete and punc tu ated by tec tonic brec cias in its lower part, it shows a rel a tivelycom plete suc ces sion of the mid Famennian. The sam pled sec tion starts above a brec cia, higher than intrenches, as K. ter mini is al ready rep re sented there (depth 75.0 m), which thus cor re sponds prob a bly to bed 3of Czarnocki (1989). The first C. prima oc curs at depth 74.0 m, im me di ately above an other brec cia. The firstoc cur rence of C. falcata at depth 71.5 sug gests a cor re spon dence to bed 19 of Wolska, al ready within thebrec cia in Czarnocki’s sec tion. In fact, most of the core cor re sponds to this hi a tus, with K. rhomboidea and C. marginifera oc cur ring up to 64.5 m. Ev i dently there is a tec tonic dis con ti nu ity above, corresponding to theupper unit in Czarnocki’s trench.

The first ap pear ance of P. jugosus at depth 63.5–63.7 sug gests a cor re spon dence to bed 27. Dasberginatrigonica has been found at depth 62.9 by Żakowa et al. (1983), which shows a suc ces sion closely sim i lar tothat at Dzikowiec. Ap par ently bed 33 of Czarnocki cor re sponds to this youn gest Famennian lime stone. Ablack shale with lime stone nod ules and tuffite at depth 61.7–61.8 prob a bly rep re sent the ter mi nal De vo nianHangenberg event, in close anal ogy with the Kowala succession.

ŚCIEGNIA

This is the lo cal ity “Wzdół Plebański” of Kościelniakowska (1967). The sam pled sec tion is along thecreek be hind the al lot ment No. 16. This is a rather mo not o nous suc ces sion of nod u lar or grey−greenwavy−bed ded marly lime stone with a spec tac u lar in ter ca la tion of black clay of the Platyclymenia annulata

16 JERZY DZIK

Event in the mid dle. Cono dont sam ples are gen er ally of rather low pro duc tiv ity. The old est of them(Wzd−10) con tains Conditolepis distorta and nu mer ous Alternognathus spec i mens with a wide plat form. Inprox im ity to the black shale (sam ple Wzd−7) the first Dasbergina ap pears, and above it Lagovignathusgranulosus has been found. 1.2 m above the black shale there is a tec tonic dis con ti nu ity sur face but prob a blynot much of the sec tion is miss ing as L. granulosus oc curs still above it (sam ple Wzd−9). A sig nif i cant re −build ing of the assemblage took place be tween this level and the top most ex posed bed (Wzd−13) withPalmatolepis rugosa. A sig nif i cant dis lo ca tion sep a rates this part of the sec tion from that to the north, wherethere is a small ex po sure of an early Famennian nod u lar lime stone within dark clays. The cono dont as sem −blage con tains Conditolepis prima and C. gilberti.

ŁAGÓW AND PŁUCKI

The old est unit of the Famennian in the Płucki−Łagów sec tion is the Lower Łagów Beds (dolne warstwyłagowskie), a rhyth mic in ter ca la tion of grey nod u lar marly lime stone and shales. The Beds were iden ti fied by Sobolew (1912b) and Czarnocki (1989) in Dule, where they are in tec tonic con tact with the over ly ing lime −stone strata of the Up per Łagów Beds along a fault. Such a rock crops out near the mouth of the ra vine onboth sides and the most com plete ex po sure is in the back yard of the house num ber 73 on the Słupecka street.Lithologically iden ti cal strata crop out along the road to Nowa Słupia and are in con ti nu ity with the Płuckisec tion. Cono dont sam ples show their early Famennian age and the only lithic dis tinc tion at which the baseof the Lower Łagów Beds could be de fined is the top of the dark lime stone and marl suc ces sion con tain ingthe Up per Kellwasserkalk ho ri zon, ter mi nat ing 4.0 m above it. The basal stra tum (sam ple Pł−40) con tains thefirst Klapperilepis delicatula.

The Słupecka 73 ex po sure is poorly fossiliferous, ex cept for the cephalopod lime stone beds, and cono −donts ex tracted from the nod u lar lime stone at the base con tain a cono dont as sem blage with Conditolepisfalcata. More than 4.0 m of the suc ces sion are ex posed there be low a cephalopod lime stone bed with largeMaeneceras, 20 cm thick. It is sep a rated from the last bed of the nod u lar lime stone with a sed i men tary dis −con ti nu ity and cov ered with a ferruginous stromatolite, mark ing an other dis con ti nu ity sur face. The lime stone succesion above, with five beds sep a rated by shale in ter ca la tions of thick ness di min ish ing up wards, maycon ve niently be con sid ered the basal part of the Up per Łagów Beds (górne warstwy łagowskie), to getherwith the cephalopod lime stone bed. The bed im me di ately un der ly ing the cephalopod lime stone con tains C.marginifera and the same cono dont as sem blage oc curs through out the strata ex posed above. Palmatolepisampla ap pears for the first time in the bed immediately above the ferrugineous stromatolite.

This means that the Cheiloceras as sem blage, which is rich in fos sils, de scribed by Sobolew (1914) andknown to me only from loose blocks col lected from the scree, must have been de rived from lat eral equiv a −lents of the strata ex posed at Słupecka 73, prob a bly from fossiliferous lime stone lenses.

A few spec i mens of the large nautiloid Mecynoceras trans ferred to me by Andrzej Piotrowski (Dzik1985) were claimed to be col lected on the left slope of the val ley. I have not been able to con firm this lo ca −tion. The lime stone ma trix (sam ple Ł−Mec; Ta ble 3) con tains Klapperilepis circularis and K. quadrantinodo −solobata, be ing thus older than any part of the Dule sec tion. A lense of cephalopod lime stone of this age wasonce ex ploited in a trench at Janczyce (Makowski 1991), about 20 km to east, in the same fa cies zone of theHoly Cross Moun tains. An al most com plete suc ces sion of the Famennian has been traced in the bore holeJanczyce 1 by Matyja and Narkiewicz (1992, 1995).

Gürich (1896, 1901) was the first who iden ti fied fos sils from the “Sacculus−Bank” on the right slope ofthe Niwa val ley in Łagów, and de ter mined their geo log i cal age. Sobolew (1911) pub lished a list of fos sils,mostly from no lon ger ex ist ing ex po sures of a fossiliferous lime stone of the Up per Łagów Beds on the leftslope of the val ley. His more de tailed strati graphic and palaeontological work on the right side of the val ley(Sobolew 1912b), in the slope ex po sure of ra vine Dule, re mains the main source of in for ma tion on this clas −sic lo cal ity. An in ter pre tive sketch of the wall was published in Czarnocki (1989).

As a re sult of ac tiv ity of fos sil col lec tors at Dule, the ammonoid−bear ing lime stone lenses were al readycom pletely ex ploited when in 1974 I at tempted to make a pre cise sam pling in a few trenches dug across theex po sure. The deep est ly ing bed reached by the trench is a mas sive synsedimentary brecciated lime stonewith peb bles of grey marly lime stone iden ti cal with that oc cur ring in the Lower Łagów Beds. In a sam ple ofsuch peb ble (Ł−39; prob a bly con tam i nated with the ma trix) Conditolepis glabra has been found. This sup −


ports its age older than the ma trix (Ł−38), where only the more ad vanced mem ber of the same lin eage, C.distorta, oc curs. The top most part of the bed is rich in py rite and con tains phosphorite peb bles com posedmostly of lar val gas tro pod shells. Orig i nally aragonitic mol lus can shells are fre quently re placed with py rite,offernig sur pris ingly pre cise rep li cas of their orig i nal mor phol ogy (Dzik 1994). The tornoceratid goniatitePseudoclymenia has been also found in this part of the bed. The over ly ing 2 m of a dark bed ded lime stonewith marly shale in ter ca la tion con tains sev eral spe cies of the cheiloceratids, with the dom i nant Dimeroceraspolonicum. In this zone prob a bly the most fossiliferous lens of the “Sacculus−Bank” of Gürich (1896) was lo −cated, ac ces si ble to col lect ing un til 1971. No clymeniid has been found in this unit by my self and I guess thatthose re ported by Sobolew (1914) and Czarnocki (1989) al leg edly from this stra tum were col lected in thenorthern part of the exposure from a lens incorrectly correlated with it.

At the base of the black mudstone cov er ing the fossiliferous lime stone bed, nu mer ous small lime stonecon cre tions (sam ple Ł−4) yielded P. trachytera and L. granulosus. Sam ples from beds of a black lime stoneabove were bar ren of cono donts, but al most cer tainly they cor re spond to blocks of the black lime stone withthe heterocoral Oligophylloides (Ł−40) and clymeniids oc cur ring in the scree. They all con tain P. trachytera.Higher up in the sec tion only un de ter min able flat tened clymeniids oc cur in the shale with in ter ca la tions ofblack lime stone with the abun dant bi valve Guerichia (well ex posed behind the allotment Słupecka 59).

CORRELATION AND CHRONOLOGY

I be lieve that in sert ing units of chronostratigraphy or biochronology (re cently re viewed by Lindsay 2003,but see Zalasiewicz et al. 2004) be tween those of geo chron ol ogy and lithostratigraphy is of lit tle pur pose andre sults in un nec es sary com pli ca tion of ter mi nol ogy (Dzik 1995). While rea son ing on age cor re la tion basedon paleontological ev i dence, it seems enough to sep a rate clearly data and hy poth e ses on evo lu tion ary events(unique but of low res o lu tion) from faunistic or floristic ex pres sions of en vi ron men tal changes (po ten tially of high res o lu tion but repeatable in principle).

The geo chron ol ogy of the Famennian was orig i nally based on ammonoid biostratigraphic sub di vi sionsas sumed to re flect evo lu tion. There are rea sons to be lieve that truly the ammonoids were the fast est evolv ingor gan isms of the De vo nian and eas i est to de ter mine tax o nom i cally (also ow ing to their low pop u la tion vari −abil ity). How ever, ex cept for a few clas si cal places in cen tral Eu rope, Rus sia, Aus tra lia, and north ern Af rica,they are rel a tively rare fos sils. It seems thus more prac ti cal and re li able to base sub di vi sion of the Famennianon the evo lu tion of cono donts even if they do not en able equally sub tle sub di vi sions. This ob vi ously re quiresa bi o log i cal ap proach to their tax on omy (that is, study of their pop u la tion vari abil ity) and all avail able mor −pho log i cal ev i dence has to be used (that is, data on the com po si tion of their whole ap pa ra tuses). This at ti tudeis fol lowed here and, al though ammonoids are also used to de ter mine chro no log i cal succesion of events, thecono dont−based zonation is the background standard to which all other ways of inference are supplementary.

Be low a re view of the evo lu tion ary events that can be used for in ter con ti nen tal cor re la tion of the cen tralEu ro pean stan dard are re viewed in chro no log i cal or der, start ing from the be gin ning of the Famennian. Toavoid mis un der stand ing, I re fer to zones intented to be chro no log i cal units (“chrons”) based on evo lu tion aryevents us ing full tax o nomic names of the in dex taxa in ital ics. Early, mid, or late units within them do nothave for mally de fined bound aries and are used in lower case. In names of biostratigraphic or ecostratigraphic zones only the spe cies rank name is used in roman letters, without the genus name.

EVOLUTIONARY DATUMS

The clas sic Famennian cono dont biostratigraphic scheme was pro posed by Ziegler (1962, 1971). Sincethat time sev eral im prove ments were in tro duced both by its au thor and other stu dents (sum ma rized byZiegler and Sandberg 1984, 1990, and Schülke 1999). All of them fol lowed the same ba sic way of in fer ence,that is bas ing the cor re la tion on the ac tual ranges of morphologies in the rock sec tions. De spite de clared ref −er ence to the phyletic evo lu tion of par tic u lar cono dont lin eages, the ver ti cal con cept of chronospecies or even a plain typology was ap plied. An ex pres sion of this meth od olog i cal at ti tude is the “Famennian” con cept ofsub spe cies, as nei ther a geo graphic race nor a seg ment of evo lu tion ary tran si tion. Some of such un der stood

18 JERZY DZIK

sub spe cies are equiv a lent to reg u lar bi o log i cal spe cies (e.g., “Palmatolepis glabra acuta”) in the mean ing ap −plied by stu dents of cono dont ap pa ra tuses from other geo log i cal pe ri ods, oth ers (e.g., “Palmatolepis qua −dran tinodosa inflexa”) seem to be just morphotypes within pop u la tion vari abil ity of a spe cies. I in sist on ap −ply ing the populationl at ti tude to the fos sil ma te rial and on de fin ing the bound aries be tween chronospecieshor i zon tally. The global cor re la tion stan dard (less pre cise, but reliable) could then truly be based on thephyletic evolution of the most suitable conodont lineages.

Most of the tra di tional zonal sub di vi sions of fered by Ziegler (1962) can eas ily serve this pur pose af ter ami nor re def i ni tion, but those in tro duced by Ziegler and Sandberg (1984) are more dif fi cult to ap ply. Themain ob sta cle is that the evo lu tion is re li ably doc u mented in very few trop i cal cono dont lin eages used in theFamennian biostratigraphy. Even in cases when this is po ten tially pos si ble, a non−bi o log i cal ap proach to tax −on omy oblit er ates the ob served evo lu tion ary change. The three youn ger of the zonal units of Ziegler andSandberg (1984) are de fi cient in this as pect. Thus, the zonal spe cies “Palmatolepis postera” is prob a bly onlya mar ginal mor phol ogy in the pop u la tion vari abil ity of P. schindewolfi. Al though the or i gin of P. schinde −wolfi is trace able in the fos sil re cord, its great pop u la tion vari abil ity makes iden ti fi ca tion of tran si tionalstages rather un re li able in small sam ples and sev eral other cono dont lin eages in the same time span seemmore prac ti cal as in dex fos sils. The tax on omy and chro no log i cal bound aries of “Palmatolepis expansa”(claimed to be a mem ber of the Tripodellus gonioclymeniae lin eage) is un cer tain and many of its oc cur rences re ported in the lit er a ture are prob a bly based on tax o nomic mis iden ti fi ca tions. The or i gin of the youn gest spe −cies, “Siphonodella” praesulcata re mains ob scure and its iden ti fi ca tion is usually uncertain because ofco−occurrence of homeomorphic species and lack of knowledge of the apparatus composition.

There fore I pro pose to come back to the orig i nal zonal sub di vi sion of the Famennian by Ziegler (1962)with only un avoid able cor rec tions re sult ing from the prog ress in un der stand ing of the tax on omy and evo lu −tion of the in dex spe cies. All but the last zonal units (or rather da tums) used here are based on the in dex spe −cies cho sen by Ziegler (1962), nomenclatorially cor rected. I am skep ti cal re gard ing re li abil ity for in ter con ti −nen tal cor re la tion of any ear lier pro posed time units of lower rank. Mod i fied def i ni tions of the evo lu tion aryzones are given be low with a brief re view of other prob a ble evo lu tion ary changes within their time spans ofpossible correlative value.

Klapperilepis triangularis Zone. — There is no evo lu tion ary change, which could be used to de fine thebase of the Famennian as cur rently un der stood (that is at the end of the Up per Kellwasserkalk en vi ron men talevent cor re spond ing to the base of triangularis zone in the Montagne Noire; Klapper et al. 1993; House et al.2000) but just an eco log i cally con trolled dis ap pear ance of a few warm−wa ter pe lagic spe cies [in Schülke’s(1999, p. 10) words, fol lowed by a “mass oc cur rence of Palmatolepis triangularis”]. The sub di vi sion of thelat est Frasnian zone pro posed by Gi rard et al. (2005) does not ap ply to the Holy Cross Moun tains as in thePłucki sec tion “Palmatolepis” linguiformis con tin ues to the end of the Up per Kellwasser ho ri zon to getherwith all its cono dont fauna, whereas in the Kowala sec tion Klapperilepis ul tima is miss ing in the lat estFrasnian (Dzik 2002).

The change from K. ul tima to K. triangularis took place some what later (Dzik 2002). This means that theear li est part of the Famennian, de fined on the en vi ron men tal change, has to be placed within the zonal unitbased on the evo lu tion, which starts in the lat est Frasnian and con tin ues to the ear li est Famennian (I pro posed Ancyrodella curvata as prob a bly the most re li able marker of this kind for the base of the ter mi nal part of theFrasnian; Dzik 2002).

Ziegler (1962) sub di vided his triangularis Zone in three parts. The mid dle unit was de fined on the first ap −pear ance of Klapperilepis delicatula and Tripodellus clarki (as un der stood in the pres ent work). Both theselin eages ap peared in the Holy Cross Moun tain by mi gra tion from elewhere (Dzik 2002) at ap prox i mately thesame time as the evo lu tion ary change from Klapperilepis ul tima to K. triangularis. If ap plied to the lo cal,cen tral Eu ro pean biostratigraphy, this first unit cor re sponds thus to prob a bly a very brief time be tween theend of the Frasnian and the base of the evo lu tion ary K. triangularis Zone. The change to K. proto −rhomboidea, as pro posed by Schülke (1999), moves the bound ary be tween the units a lit tle but the or i gin ofthe new in dex spe cies is equally cryp tic and dif fi cult to trace. The up per unit of Ziegler’s triangularis Zone isde fined on the ap pear ance of Conditolepis tenuipunctata. This is prob a bly a true evo lu tion ary event, in ef fect of nar row ing the plat form of P1 el e ment and wid en ing the plat form of P2 el e ments of the an ces tralConditolepis lobicornis, but the ex act course of evo lu tion re mains to be dem on strated. Any way, this may be


a use ful da tum for the global cor re la tion, better than the or i gin of K. robusta proposed by Schülke (1999), theancestry of which remains unknown.

The change from Klapperilepis to Palmatolepis (Dzik 2005) may ap pear use ful for cor re la tion in the mid −dle of the Zone. Near the end of the zone, the triramous P2 el e ments orig i nated in the lin eage of Tripodellusbut tran si tional pop u la tions are poly mor phic and co−oc cur with more con ser va tive lin eages of the same ge nus which makes their ap pli ca tion to dat ing rather un re li able. The tran si tion from Klapperilepis proto rhombo −idea to K. rhomboidea is of even less im por tance be cause of rar ity of these spe cies and their great pop u la tionvari abil ity.

Klapperilepis crepida Zone. — The zonal in dex spe cies orig i nated from K. robusta, as shown by Schülke (1999). Both chronospecies are vari able and tran si tional pop u la tions con tain modal morphologies of both ofthem. The sig nif i cant over lap in their ranges as in ter preted by that au thor is thus prob a bly a re sult of ap pli ca −tion of the ver ti cal spe cies con cept. Schülke (1999) pro posed to de fine the base of the zone on the or i gin of“Palmatolepis abnormis sandbergi”. This is a good evo lu tion ary da tum, as long as M el e ments are used toiden tify the (sub)spe cies. Un for tu nately, the most com mon P1 el e ments are in di vid u ally hardly dis tin guish −able from those of the an ces tral K. triangularis. The tran si tion took place much ear lier than the first ap pear −ance of K. robusta, thus well within the K. triangularis Zone in its traditional meaning.

In the Holy Cross Moun tains, sec tions K. robusta changes into K. ter mini with a grad ual in crease in theheight of denticle rows. This trans for ma tion is of cor re la tive value and Ziegler (1962) used it to de fine thebound ary be tween the lower and mid dle parts of the Zone. It re mains un clear whether the dif fer ent time ofap pear ance of K. crepida and K. ter mini is re lated to their evo lu tion, re sults of mi gra tions or an ar ti fact of tax −on omy. An other fast evolv ing lin eage of the zone is that of Conditolepis tenuipunctata ® C. glabra, the ba −sis of Ziegler’s up per part of the Zone. This is a phyletic change that has to be care fully rec og nized. Ap par −ently, Schülke (1999) ap plied there the ver ti cal spe cies con cept, which re sulted in a chro no log i cal over lap ofsuc ces sive spe cies.

Ziegler (1962) in tro duced the rhomboidea Zone lo cated be tween crepida and quadrantinodosa zonesbased on the ap pear ance of Klapperilepis rhomboidea. This spe cies is clearly a suc ces sor of K. proto −rhomboidea. The evo lu tion ary change was slow and dif fi cult to trace be cause of a wide pop u la tion vari abil −ity and morphologic sim plic ity of P1 el e ments. The ap pear ance of K. rhomboidea in the Holy Cross Moun −tains is an eco log i cal event of ques tion able cor re la tive value. Prob a bly its old est oc cur rence is at Kadzielnia(sam ple Ka−3) where it co−oc curs with the late K. ter mini. Im Łagów, Kowala and Miedzianka the spe ciesco−oc curs with C. quadrantinodosa and dis ap pears be fore its change into C. marginifera.

Conditolepis quadrantinodosa Zone. — The highly vari able in dex spe cies C. quadrantinodosa orig i −nated from C. klapperi, but the change has not been traced in any sec tion. Some time af ter this event, C.inflexoidea orig i nated allopatrically in re spect to C. quadrantinodosa and then re turned to the area of oc cur −rence of the C. quadrantinodosa ® C. marginifera lin eage. This prob a bly en forced both sympatric pop u la −tions to nar row their vari abil ity (char ac ter dis place ment). C. inflexoidea vari abil ity was re duced mostly to the morphotype with oval plat form and nar row crest along its an te rior lobe. In fact, a re mark able re build ing ofcono dont as sem blages is con nected with the en trance of C. klapperi to the Kadzielnia suc ces sion. Per haps itwould be rea son able to de fine the base of the Zone at this event, but the or i gin of C. klapperi re mains cryp tic.

Conditolepis marginifera Zone. — Ziegler (1962) used the dis ap pear ance of C. inflexoidea as the ba sisfor dis tin guish ing his two parts of the C. quadrantinodosa Zone. This is clearly an eco log i cal phe nom e nonand not a re li able ba sis for time cor re la tion. The change from C. quadrantinodosa to C. marginifera is not somuch con nected with in tro duc tion of any evo lu tion ary nov elty but rather with de crease in pop u la tion vari −abil ity. Any way, these spe cies oc cur usu ally in large num ber of spec i mens and their tax o nom i cal iden ti fi ca −tion seems rather easy. Ziegler and Sandberg (1984) pro posed the or i gin of their sub spe cies “Palmatolepismarginifera utahensis” as the marker for the base the up per unit but the bi o log i cal meaning of this taxonremains to be settled.

Per haps the or i gin of Palmatolepis ampla from P. perlobata is also of po ten tial cor re la tive value withinthis zone but these lin eages ap par ently split allopatrically.

Palmatolepis trachytera Zone. — The zone, with its lower bound ary de fined at the or i gin of the nom i nalspe cies was in tro duced by Ziegler and Sandberg (1984). The tran si tion from P. ampla is still in ad e quately

20 JERZY DZIK

doc u mented but very likely. The old est pop u la tion of P. trachytera share the pres ence of sharp pos te riorlobes with its an ces tor. In the orig i nal mean ing, this was the bound ary be tween the lower and mid dle part ofZiegler’s (1962) velifera Zone, based on the ap pear ance of Scaphignathus velifer (trans ferred to marginiferaZone as its up per most unit by Ziegler and Sandberg 1984). This lin eage of shal low−wa ter cono donts is un dera strong en vi ron men tal con trol in its dis tri bu tion and was rightly aban doned as the ba sis for global time cor −re la tion. Un for tu nately, this re fers also to the ap pear ance of Dasbergina granulosa, the ba sis for rec og ni tionof the uppert part of the velifera (or the whole trachytera) Zone.

The or i gin of Lagovignathus granulosus from L. lagoviensis is of po ten tial cor re la tive value in the youn −ger part of the zone.

Lagovignathus styriacus Zone. — The change from Lagovignathus granulosus to the nom i nal spe ciesde fines the base of the zone, as in tro duced by Ziegler (1962). Ziegler and Sandberg (1984) re named it as thepostera Zone de fined on the or i gin of “Palmatolepis perlobata postera” which seems to be a morphotype ofP. schindewolfi. The or i gin of L. styriacus seems to be a much more re li able ba sis for the age correlation.

Of po ten tial cor re la tive value is also the as so ci ated Palmatolepis rugosa. As shown by the mor phol ogy of the P2 el e ment, this is a mem ber of the P. trachytera lin eage but in sec tions stud ied by my self there is al waysa gap in their dis tri bu tion, thus the tran si tion can not be demonstrated.

Ziegler and Sandberg (1984) pro posed the or i gin of T. mancus as the lower limit of their up per posteraZone but this is a rare spe cies of un clear re la tion ship to both the an ces tral T. gracilis and its pos si ble suc ces −sor T. gonioclymeniae. Al though of po ten tial cor re la tive value this event re mains to be tested.

Pseudopolygnathus jugosus Zone. — The up per part of Ziegler’s styriaca Zone was de fined on the ap −pear ance of P. jugosus. The or i gin of icrion with three rows of denticles in this Pseudopolygnathus (or per −haps rather Dasbergina) lin eage seems to be rather well doc u mented. The unit cor re sponds roughly to the tra −di tional costatus Zone to gether with Ziegler and Sandberg’s (1984) lower expansa Zone. The tax o nomic sta −tus of “Palmatolepis expansa” pro posed to de fine the lower bound ary of the zone, re mains to be clar i fied. Asmen tioned al ready above, no such spe cies has been en coun tered in the Holy Cross Moun tains or the Sudetes.I see also no pos si bil ity to trace in the avail able ma te rial the evo lu tion ary or i gin of “Siphonodella”praesulcata, defining its upper limit.

Ziegler’s costatus Zone was de fined on the ap pear ance of Pseudopolygnathus ziegleri, as un der stood inthis work. Un like P. jugosus, the or i gin of this spe cies re mains cryp tic and its oc cur rence, sim i lar to otherFamennian and Tournaisian spe cies of the ge nus, is rather cha otic, ev i dently as a re sult of a high eco log i calsen si tiv ity. This re fers as well to P. aculeatus, cho sen by Ziegler and Sandberg (1984) to de fine their mid dleexpansa Zone. Even the change from P. ziegleri to P. ultimus, of po ten tial cor re la tive value (de fin ing the up −per expansa Zone), may be mis lead ing. At Kowala the an ces tral spe cies re ap peared af ter some time of oc cur −rence of P. ultimus.

Tripodellus gonioclymeniae, used by Ziegler (1962) to de fine the mid dle and up per parts of the costatusZone re mains of cryp tic or i gin, al though its or i gin from T. mancus is likely. The lin eage orig i nated prob a blyallopatrically from P. gracilis but the al leg edly tran si tional role of “Palmatolepis expansa” (Ziegler andSandberg 1984) is far from proven.

Dasbergina trigonica Zone. — The or i gin of the nom i nal spe cies of this newly pro posed zone fromDasbergina marburgensis is shown by a tran si tion se ries in the up per part of the clymeniid lime stone atDzikowiec in the Sudetes. The zone roughly cor re sponds to Ziegler and Sandberg’s (1984) praesulcata Zone. The top most part of the unit in cludes the Protognathodus kockeli fauna. The lat ter im mi grated to the equa to −rial zone to gether with an im pov er ished cold−wa ter ammonoid fauna near the end of the Famennian (if de −fined at the end of the Hangenberg black shale en vi ron men tal event). The ex act time and place of or i gin of P.kockeli re mains to be de ter mined and the three units within the praesulcata Zone, de fined on the dis ap pear −ance of T. gonioclymeniae and en trance of P. kockeli, are of clear ecostratigraphic nature.

CONODONT ECOSTRATIGRAPHY

Like any other ap pli ca tion of evo lu tion ary changes to age cor re la tion, the cono dont zonation of theFamennian al lows only a crude ap prox i ma tion to chro no log i cal re la tion ships among fossiliferous strata. It isthus tempt ing to re fine the strati graphic res o lu tion by us ing short−term en vi ron men tal changes of pos si bly


wide ex tent. The eas i est and prob a bly most re li able tech nique of such a cor re la tion is the palynological meth −od ol ogy of com par ing changes in per cent age con tri bu tion of eco log i cally sen si tive taxa, the pro ce dure fre −quently used in connecion with conodont biofacies analysis.

Such “palynological” logs based on the Famennian cono donts (Figs 5, 6) ap pear prom is ing for thewithin−re gion cor re la tion but much more re search is needed to ap proach truly use ful re sults. It has ap pearedthat much more dense sam pling would be nec es sary than done by my self for the ap pa ra tus study. Only onesuc ces sion is sam pled well enough in this re spect, that is the up per part of the Kowala sec tion (Fig. 6). Arather clear pat tern of al ter na tion of pre sum ably shal low− (and cold?−) wa ter fau nas dom i nated by Pseudo −polygnathus and more open−sea palmatolepidid−rich as sem blages is vis i ble in the log. Three cy cles coverthere the P. trigonicus Zone and an other three the P. jugosus Zone. In the co eval part of the Dzikowiec sec −tion these cy cles are not so clearly rec og niz able, mostly be cause of a rel a tively deeper−wa ter en vi ron ment.Un like Kowala, im me di ately af ter the transgressive event near the be gin ning of the P. jugosus Zone a rel a −tively sta ble en vi ron ment was es tab lished there.

Ex cept for the trench at Kowala, the sam pling is too sparse to show con vinc ingly such a pat tern, but somelarge−scale cy cles are shown by the dis tri bu tion of the open−sea palmatolepidids and shal low−wa ter icrio −dontids. The dis tri bu tion of Conditolepis falcata, the dom i nant spe cies in the Holy Cross Moun tains lateFamennian, seems mean ing ful. There are three peaks in its dis tri bu tion sep a rated by the dom i nance ofIcriodus (at Miedzianka), polygnathids (Łagów) or both taxa (at Kowala). Two such peaks cor re spond to theC. marginifera Zone, the P. trachytera Zone cor re sponds to a prob a ble shallowing, and the third peak cov ersat least the lower part of the L. styriacus Zone. More dense sam pling would prob a bly re veal a smaller scalecyclicity within each of those events, as sug gested by the re cord at Łagów, where the late C. marginiferaZone deep en ing seems to be split into two events and a sim i lar cyclicity as in Kowala may have oc curred in

22 JERZY DZIK

Fig. 5. Logs of per cent con tri bu tion of cono dont el e ments rep re sen ta tive of the main lin eages for sam ples from the earlyFamennian of the Holy Cross Moun tains. Rel a tive po si tion of sam ples stan dard ized in ref er ence to pro posed chronologic sub di −

vi sions.

the P. trachytera Zone. The com plex tec ton ics makes sam pling dif fi cult at this site, and it is even less re li able in this re spect at Miedzianka.

The K. crepida Zone is char ac ter ized by a rather sta ble en vi ron ment, ex cept for its transgressive base. InK. triangularis Zone and the ear li est Famennian strata de pos ited be fore this Zone, sev eral peaks in abun −dance of Icriodus are rec og niz able but in com plete sec tions do not al low use of their po ten tial cor re la tivevalue. This may be of po ten tial value in es tab lish ing a high res o lu tion chro nol ogy of the Late De vo nian, as asim i lar rythmicity seems to be ex pressed also in the Chi nese sec tions (Wang and Ziegler 2002).

Un for tu nately, the rel a tively com plete re cord is avail able only for rather deep−wa ter en vi ron ments of theFamennian of Po land. Shal low−wa ter succesions are known from ex po sures only in the Cra cow re gion(Baliński 1995) al though they have been reached by nu mer ous drillholes in Silesia (Narkiewicz 1978; Belka1998), Pomerania (Matyja 1993), and the Lublin re gion (Matyja and Żbikowska 1985). The cores yieldedcono dont fau nas with many spe cies un known in the area sam pled by my self. These in clude bi zarre Cteno −polygnathus rarus (Baliński, 1995), Omolonognathus transformis Gagiev, 1979 (Baliński 1995), Mashkoviasilesiensis Belka, 1998, and Icriodus chojnicensis Matyja, 1972. Po ten tial oc cur rence of sim i lar as sem blagesin the Holy Cross Moun tains is sug gested by the iso lated sam ple from Karczówka con tain ing or nate new spe −


Fig. 6. Logs of per cent con tri bu tion of cono dont el e ments rep re sen ta tive of the main lin eages for sam ples from the earlyFamennian of the Sudetes and Holy Cross Moun tains. Rel a tive po si tion of sam ples stan dard ized in ref er ence to pro posed

chronologic sub di vi sions.

cies of Mitrellataxis. The same spe cies have been iden ti fied in the sin gle pro duc tive sam ple from co evalstrata at the Żbik ra vine near Dębnik col lected by my self.

METHODS

Ammonoid conchs and other macrofossils on which this study is based, have been ex tracted from the rock and il lus trated us ing stan dard, widely used tech niques and equip ment, which do not need to be de scribed inde tail. A some what more so phis ti cated method has been ap plied to chem i cal ex trac tion of microfossils.These were mostly phos phatic cono dont el e ments or fish teeth and scales, but also sec ond arily phosphatizedor pyritized ammonoid conchs. Be cause the treat ment of sam ples may in flu ence re sults of ex trac tion, theprocedure is presented below.

CONODONT ELEMENT EXTRACTION

The cono dont sam ples were dis solved in ace tic (pure lime stone) or for mic (marls, dolomitized lime −stones) ac ids. Al ways there was less acid added than nec es sary to dis solve the car bon ate and this pro videdsome buff er ing. The res i due was passed through a 1 mm sieve to re move acid−re sis tant flakes of the shale and un dis solved lime stone pieces. The large frac tion was screened af ter be ing dried for ex tremely large spec i −mens. The re main ing frac tion was de canted, in stead of sieved, to avoid loss of very small el e ments andbreak age of ex tremely frag ile ones. Res i dues rich in clay, de vel op ing mud cracks af ter be ing dried, werewashed again. Dry res i dues with high con tent of non−phos phatic de bris were sep a rated in the elec tro mag netic min er al og i cal sep a ra tor, af ter com ing again through a 1 mm sieve to re move ag gre gates block ing the fun nelof the sep a ra tor. The an gle and am per age of sep a ra tion were cho sen sep a rately for each se ries of sim i lar sam −ples. It was avoided to pro ceed with sep a ra tion too fast or to re move much more than half of the low−mag −netic frac tion of the res i due in one pass. Com monly the sep a ra tion was re peated a few times. Only in case ofsilica−rich samples from the Sudetes were heavy liquids used in separation.

The spec i mens are kept loose in plas tic slides (Franke Zellen) in stead of be ing glued to the pa per, as isthe Amer i can prac tice. The rea son for this is that glu ing of a very large num ber of spec i mens is time con −sum ing and makes mov ing and seg re ga tion of spec i mens (nec es sary to ar range them in ap pa ra tus sets, asex plained be low) dif fi cult. It may also be de struc tive to frag ile spec i mens. All to gether 235,031 spec i menshave been tax o nom i cally iden ti fied and counted. Some sam ples were dis solved for ad di tional ma te rial atlate stages of prep a ra tion of the work and some mi nor in con sis ten cies in data pre sented on Ta bles 1–19may re sult from this.

Mea sure ments (pub lished in Dzik 2002 and 2005) have been done not on ac tual spec i mens but on theircon tour draw ings made with a mi cro scope draw ing ap pa ra tus. For the mea sure ments, the spec i mens ofpalmatolepidids were first ar ranged in rows. Their most sta ble ori en ta tion is with the denticle row (ca rina)ver ti cal but the draw ing ap pa ra tus shows the view from the right oc u lar, that is slightly oblique. Po ten tially itwould be pos si ble to make spec i mens ap pear ver ti cal in such view but the dis tor tion does not seem sig nif i −cant as long as all spec i mens were drawn un der the same an gle. To as sure this, the pos te rior lobe of all spec i −mens, both dextral and sinistral, was oriented towards the right while drawn.

RECONSTRUCTION OF CONODONT APPARATUSES

The most ef fi cient way to re store cono dont ap pa ra tuses at the pres ent stage of knowl edge is by in vent ingwork ing hy poth e ses on their com po si tion to fal sify and ver ify them while col lect ing new sam ples (Dzik1991). All the el e ments in each sam ple have to be sep a rated into mor pho log i cal classes to match them intosets of par tic u lar spe cies. It is help ful to con sider the shape, ro bust ness of denticulation and col or ation of el e −ments at this stage of rea son ing. In this study, the el e ments within a large Franke cell were ar ranged in pileswith a hair mounted on a holder. Sets rep re sent ing the icriodontids, prioniodinids, polygnathids, andpalmatolepidids were then trans ferred to sep a rate cells and the sep a ra tion was con tin ued un til se ries of all theel e ment types for each spe cies were ar ranged. Their various combinations represent hypotheses to be tested.

24 JERZY DZIK

In prin ci ple, the whole se ries of el e ment types of a spe cies has to be rep re sented in a large sam ple. Each of the iden ti fied mor pho log i cal classes can thus be at trib uted to a multielement spe cies. First, the al ready known ap pa ra tuses were iden ti fied and re moved. Pro vi sional group ing of re sid ual el e ments was at tempted to fit thebetter known ap pa ra tuses. This was then tested by re peat ing the same pro ce dure with suc ces sive sam ples.Usu ally such con cepts are soon fal si fied (ob vi ously, this re quires that sam ples are sig nif i cantly dif fer ent inspe cies com po si tion, be ing taken from var i ous ho ri zons and kinds of the rock). The re main ing, not re jectedhy poth e ses on the ap pa ra tus com po si tion are fur ther tested by look ing for a pos si bly close rel a tive (pref er a −bly be long ing to the same evolv ing lin eage) with mor pho log i cally sim i lar el e ments in par tic u lar ap pa ra tus lo −ca tions. The whole phylo gen etic tree has to be log i cally consistent in its every aspect, including continuity ina homologous series of elements between species.

The main ob sta cle in re con struct ing the Late De vo nian cono dont ap pa ra tuses is the de fi ciency ofnon−plat form el e ments in sam ples (their un bal anc ing). As a re sult only the ap pa ra tuses of nu mer i cally dom i −nant spe cies are re li ably known at pres ent and many re con struc tions of Famennian apparatuses remainincomplete.

A spe cial as pect of ap pa ra tus re con struc tions is that phylo gen eti cally re lated sympatric spe cies sharesome of the el e ment types. More pre cisely, their pop u la tion vari abil ity may ap pear wider than the dif fer encebe tween spe cies. In such cases the spec i mens not at trib ut able to par tic u lar spe cies have to be counted to −gether for the whole spe cies group (as given in the Ta bles 1–19) or sep a rated ar bi trarily (as in cluded in logsof per cent con tri bu tion).

TAXONOMY

Not only the ap pa ra tus study but also tax o nomic de ci sions have been taken sep a rately for each sam ple.The pop u la tion ap proach to the spe cies−level tax on omy was used. Al ready while as sem bling el e ments intoap pa ra tuses, the spec i mens within a sam ple were sep a rated into morphologic classes on the ba sis of theunimodal dis tri bu tion of morphologic char ac ters, more or less objectively.

No no men cla tor ial tax o nomic con sid er ations were un der taken un til a rea son able sta bil ity of the hy poth e −sis on the ap pa ra tus com po si tion was reached by re peated tests. Spe cies have been de fined on the ba sis of themost di ag nos tic char ac ters within the ap pa ra tus, ir re spec tive of whether the char ac ter is tic el e ment is easy tore cover (bears a plat form) or not. Only af ter the ap pa ra tus struc ture of a spe cies is es tab lished, can sin glespec i mens be safely iden ti fied tax o nom i cally with pre ci sion ex press ing their di ag nos tic value. This meansthat it is waste of time to at tempt spe cies iden ti fi ca tion sep a rately for each el e ment. The mor phol ogy alone isusu ally not enough to de ter mine its spe cies iden tity. It may even hap pen that this is done de spite the mor phol −ogy, if a spec i men ap pears to be an end mem ber of a con tin u ous spec trum of pop u la tion vari abil ity. In stead,its co−oc cur rence in the same sam ple with taxonomically identifiable specimens is relied on (for detaileddiscussion of methodology see Dzik 1991).

INFERENCE ON EVOLUTION

The method of chronophyletics (Dzik 1995, 2005) has been used to study the course of evo lu tion. Data on spe cies de rived from par tic u lar sam ples were ar ranged ac cord ing to their po si tion in time and to themorphologic dif fer ences. This is enough to en hance pos si ble chronomorphoclines rep re sented by suc ces −sions of pop u la tions, more or less con tin u ous in their mor pho log i cal gra da tion. The fos sil re cord of someFamennian cono donts ap pears vir tu ally com plete, en abling ap pli ca tion of stratophenetics to doc u ment theirevo lu tion bio met ri cally (Dzik 2005). This kind of re search has not been widely ap plied only be cause of tech −ni cal lim i ta tions: there are too many spe cies to be iden ti fied and de scribed to make application of thetime−consuming biometrics practical.

Hy poth e ses on the course of evo lu tion pro duced with ap pli ca tion of chronophyletics are falsifiable byretrodiction (Dzik 1995, 2005) and this is ba si cally the same kind of rea son ing as that used in test ing the ap −pa ra tus re con struc tions.


CONODONT APPARATUSES

The most strik ing as pect of a cono dont oral ap pa ra tus is its sub di vi sion into morphologic (and func tional)units, rep re sented an te ri orly by the sharp, in ci sor−like el e ments of the M se ries (usu ally only a sin gle pair),the comb−like el e ments of the S se ries in the mid dle (usu ally 9 el e ments), and the ro bust, usu ally molarized Pse ries at the pos te rior end (usu ally two pairs). The no ta tion sys tem for the el e ments within se ries fol lows here the pro posal of Purnell et al. (2000). Their po si tion and ori en ta tion in the an i mal body is rel a tively well rec −og nized, al though only a frac tion of taxa is known from ar tic u lated nat u ral as sem blages of el e ments andsome mi nor dif fer ences in views re main. Descriptives for ori en ta tion of all el e ments are based on their se rialhomology with the plat form se ries el e ments, not on the ac tual ori en ta tion in the ap pa ra tus (Fig. 7). The ap pa −ra tus is a highly di ver si fied struc ture, of fer ing a lot of information on mechanisms of its morphogenesis,possible function, and evolution.

PATTERNS OF MOLARIZATION

A morphologic gra di ent is usu ally ex pressed in cono dont ap pa ra tuses, the most ro bust and the sharp est el −e ments be ing those lo cated at the op po site ends of the ap pa ra tus. In nu mer ous cono dont lin eages of var i ousage the occlusal sur face of the pos te ri orly lo cated el e ments of the ap pa ra tus in creased their work ing sur face,ap par ently in con nec tion with their grind ing func tion (e.g., Sweet 1988). This is an as pect of molarization,anal o gous to that in mam ma lian evo lu tion. In rare cases the evo lu tion ary ex pan sion of a ro bust ap pear ancereached as far as the anteriormost el e ments of the ap pa ra tus (for in stance in the Si lu rian Johnognathus). Thepat terns of molarization var ied, but three ways to reach the needed ef fect were usu ally cho sen by naturalselection: development of a platform, icrion or peniculus.

Plat form. — This is the most com mon kind of molarization. The plat form is a thick en ing of the el e mentwall im me di ately above the mar gin of the basal cone. This may be con nected with a nar row ing of the basal

26 JERZY DZIK

MMM

SSS

SSS

SSS

SSS

SSS

PPP

PPP

111

222

333

444

000

111

222

anterior

posterior

right(dextral)

left(sinistral)

Fig. 7. A. Ap pa ra tus of the best known palmatolepidid cono dont, the lat est Frasnian Lagovignathus bogartensis (Stauffer, 1938). Re con struc tion based on the clus ter de scribed by Lange (1968; with ori en ta tion of S 0 re versed and S1 ar bi trarily cho sen, slightlymod i fied af ter Dzik 1991), seen from its ven tral side. Descriptives for el e ments’ ori en ta tion used in the text given. B–D. Re gen −er a tion and patho log i cal mal for ma tion of ramiform polygnathid cono dont el e ments S3–4 from the C. quadrantinodosa Zone atMiedzianka (sam ple Md−2) with regerating pro cesses and two spec i mens dis placed and uni fied dur ing life (D); spec i mens ZPAL

cXVI/2126, 2128, and 2127.

cav ity and fi nally its in ver sion (as is the case with the polygnathid and palmatolepidid ap pa ra tuses). The mar −gin of the plat form may as well re main nar row and its growth then fol lows an ex pan sion of the deep basalcav ity (as is the case with the Or do vi cian balognathids). This dif fer ence does not af fect the shape of theocclusal sur face (as well ex em pli fied by the Car bon if er ous Cavusgnathus and Mestognathus). Nev er the less,the plat form of an el e ment with a nar row basal cav ity is much more mas sive and its de vel op ment re quiredmuch more ex pense of cal cium phos phate. This is prob a bly why the cold−wa ter cono donts gen er ally de vel −oped plat form el e ments with a thin wall whereas in trop i cal en vi ron ments, where access to calcium isunlimited, the conodonts developed massive elements.

Icrion. — This is a way to molarize al ter na tive to the plat form, al though it may de velop as the ter mi nalstage in the evo lu tion of a plat form. The molarized area is formed by tips of denticles of the blade fol lowedby par al lel rows of ac ces sory denticles of the same height (Dzik 1991). Icrion is usu ally of a fusiform shapeand its denticles tend to be ar ranged in trans verse ridges. Al most al ways this kind of molarization char ac ter −ize el e ments with a deep basal cav ity and thin walls, typ i cal of rel a tively cold−wa ter en vi ron ments (the mostcom mon be ing the icriodontids and idiognathodontids). Rarely it de vel oped also in elements with invertedbasal cavity.

Peniculus. — This is a mod i fi ca tion of the icrion with very nu mer ous, nee dle−like denticles form ing akind of brush (Lindström and Ziegler 1965). This struc ture de vel oped in its typ i cal form in three groups ofpolygnathid cono donts: the Givetian to Frasnian Skeletognathus lin eage, the Frasnian ancyrodellids, andFamennian rel a tives of Mehlina. It re mains to be de ter mined whether it orig i nated only once in a cladecom posed by those cono donts or it is just an ef fect of a reg u la tory mech a nism con trol ling the el e mentgrowth. Usuallly the peniculus was a ju ve nile fea ture and changed into a reg u lar plat form af ter the basalcav ity in verted. De spite the sug ges tion by Lindström and Ziegler (1965), this does not seem to be a case ofpa thol ogy.


Fig. 8. A. Di a gram matic pre sen ta tion of the model of cono dont el e ment growth and morphogenesis with as sump tion that thephos phatic tis sue was an in ter nal skel e ton un der a keratinous work ing sur face (mod i fied af ter Dzik 2000). B. Im prints ofameloblasts re stricted to the most in tensely grow ing part of the P1 el e ment, pre sum ably the base ment mem brane did not de velopthere, be ing re stricted to the cen tral part of the plat form; Polygnathus semicostatus Branson et Mehl, 1934 from the P. trachyteraZone at Łagów (sam ple Ł−27; spec i men ZPAL CVI/2570; see also Fig. 62P). C. Im prints of ameloblasts cov er ing whole plat −form of the P1 el e ments el e ment, with ap par ent pro lif er a tion cen ters at tips of denticles; Klapperilepis ul tima (Ziegler, 1958)

from the pre−K. triangularis Zone strata at Płucki (sam ple Pł−36; spec i men ZPAL CVI/1509; see also Fig. 79E).

GROWTH OF ELEMENTS

The cono dont el e ments grew by add ing layer by layer of cal cium phos phate on its occlusal (work ing) sur −face. This is a close anal ogy (per haps even homology) to the enamel of ver te brate teeth and scales. It seemsthus rea son able to ex pect that mech a nisms con trol ling the histogeny of el e ments were sim i lar to those reg u −lat ing de vel op ment of ver te brate teeth. The main dif fer ence is that cono dont el e ments did not erupt. There isa va ri ety of opin ions how they acted me chan i cally while being secreted (reviewed in Dzik 2000).

Morphogenesis. — Any cono dont el e ment, ir re spec tive of its fi nal size and shape, started its de vel op −ment as a min ute phos phatic cone (Fig. 8A). One may guess that for ma tion of the cusp of the cone was con −trolled by sim i lar fac tors as the de vel op ment of enamel−built cusps of Re cent ver te brate teeth, that is in ef fectof for ma tion of an “enamel knot” in the se cre tive ep i the lium above (Dzik 2000). With the sub se quent growthof early cono donts (Mitrellataxis and Jablonnodus may rep re sent this ar chaic pat tern in the Famennian), theel e ment in creased the thick ness of its wall and height of the cusp, chang ing its shape only a lit tle. In more de −rived cono donts, el e ments ex panded ba sally into pro cesses or a plat form un til enough space for a morpho −genetic field emerged to al low for ma tion of ad di tional coni form units (denticles or tu ber cles). The num ber ofdenticles is thus more or less strictly cor re lated with the el e ment size (Dzik and Trammer 1980). This as pectof the el e ment growth rad i cally in flu ences their mor phol ogy and to avoid mis lead ing dif fer ences and sim i lar −i ties, the specimens are illustrated here in the same magnification, whenever possible.

More or less prom i nent lon gi tu di nal ridges or na ment ing el e ments prob a bly de vel oped un der morpho −genetic con trol of a sim i lar kind as the denticles. Apart from them, the el e ment sur face may be smooth or or −na mented with a re tic u late pat tern. By anal ogy with Re cent fish scales, it may be spec u lated that the smoothsur face was se creted un der a rel a tively well−de vel oped base ment mem brane of the se cre tory ep i the lium,whereas in fast−grow ing re gions, where the mem brane was miss ing, par tic u lar se cre tory cells (ameloblasts)di rectly con tacted with the phos phatic tis sue, leav ing im prints on its sur face (Dzik 2000). Re tic u late or na −men ta tion char ac ter izes usu ally tips of denticles and mar gins of the plat form (Fig. 8B), but in some cono −donts (for in stance the palmatolepidids) it cov ers the whole el e ment sur face. Al though ini ti a tion of tu ber clesor denticles was con nected with proliferiation of ameloblasts (Dzik 2000), their size did not change dur ingsub se quent growth (von Bit ter and Norby 1994). In rare cases (Playfordia, Guizhoudella), the elementsurface is covered with minute tubercles.

Sinuosa larva. — In early mem bers of plat form cono dont lin eages, ju ve nile P1 el e ments with con i calbasal cav ity de vel oped an in cip i ent plat form above the basal cone, some times at some dis tance from it. Thissep a ra tion of struc tures was grad u ally oblit er ated as an ef fect of the ex pan sion and thick en ing of the plat −form. At the end of growth the con i cal basal cav ity is rep re sented only by an in dis tinct pit. In some lin eages,how ever, the in verted cav ity of the masive plat form re placed the cone rather abruptly and even in ma ture el e −ments a prom i nent basal cone is rec og niz able as a dis crete unit. Such a dras tic change in the on tog eny ap par −ently rep re sents a pro found func tional and phys i o log i cal trans for ma tion. Changes of this kind have to be re −ferred to as a meta mor pho sis; the early stage marked by so distinct a discontinuity is a larva (Fig. 9).

In Neopolygnathus this lar val stage lasted 12–14 days, as in di cated by the growth in cre ments (as sum ingthese are daily) and then the in verted basal cav ity de vel oped be low the dor sally lo cated in cip i ent plat form.The plat form and its basal cav ity ex panded in all di rec tions, fi nally en clos ing the con i cal unit (Fig. 9). In thelin eage of Tripodellus a grad ual evo lu tion in the ex tent of the lar val stage can be traced. In early mem bers ofthe lin eage, there is no rec og niz able con i cal el e ment stage in tax o nom i cally iden ti fi able ju ve nile spec i mens.It de vel oped in pop u la tions from the C. marginifera Zone, be ing rec og niz able only on one side of the P1 el e −ment. Such spec i mens are tra di tion ally re ferred to as Palmatolepis sinuosa, so I pro pose the name “sinuosalarva” for this de vel op men tal stage. These cono donts, clas si fied here in T. schleizius, meta mor phosed af ter 5to 15 days. In later pop u la tions from the P. trachytera Zone the change oc curred af ter 25–35 days(Tripodellus gracilis). In even more ad vanced Tripodellus gonioclymeniae, which ap par ently orig i nated as are sult of a short en ing of the postlarval growth and ear lier mat u ra tion (neoteny), the con i cal cav ity ex isted un −til the end of the el e ment growth. An im pres sive case of par al lel ism in this re spect is the de vel op ment in ap −prox i mately the same time of a strongly asym met ric lar val basal cone in the com pletely un re lated poly −gnathid Lagovignathus? dissimilis. The sinuosa larva also in this case expanded almost to the end of theelement growth.

28 JERZY DZIK

It can only be spec u lated why the sinuosa larva de vel oped in these lin eages. In the case of the poly −gnathids, ju ve nile spec i mens may pos si bly have de pended on yolk or were bred in a kind of pouch. Al ter na −tively, per haps they fed on a kind of food dif fer ent from that used by adults, not need ing the grind ing ac tionof the plat form se ries el e ments. The lat ter ex pla na tion does not re fer to Tripodellus, as no change of theocclusal sur face cor re sponds there to modifications of the base.

Ter mi na tion of growth. — Al though cono dont el e ments in most sam ples show great dif fer ences in sizeand no doubt they func tioned while grow ing, prob a bly the growth of most of them ter mi nated un der ge neticcon trol. This is not so ap par ent in the case of el e ments grow ing al most iso met ri cally, but even then a con cen −tra tion of growth lines may indicate its termination.

The growth in cre ments (Fig. 9), fre quently well vis i ble within the basal cav ity (or on the sur face of the in −verted basal cav ity), show a sur pris ingly reg u lar and rhyth mic dis tri bu tion. In el e ments of me dium size theycount 60–70. If these are daily in cre ments, cono donts reached their ma tu rity ap prox i mately af ter two to threemonths. As noted above, the change from a coni form to in verted basal cav ity in Neopolygnathus occurredafter two weeks.

The in creased den sity of in cre ments is not the only ex pres sion of ceased growth. Much more im pres siveand con vinc ing is the change in the cur va ture of pro cesses. The tip of prom i nently denticulated el e ments isfre quently re curved, de vel op ing a barb−like form, with the ter mi nal denticle point ing in the same di rec tion asthe pro cess. Such an el e ment could grow fur ther only in the height of its denticles, with the length of pro −cesses re main ing con stant. This phe nom e non is com mon in so many un re lated taxa, that one may sug gestthat the on tog eny of all conodonts was controlled in a similar way.

Re gen er a tion. — A res to ra tion of bro ken cusps and denticles to their orig i nal shape is a com mon phe −nom e non in cono dont el e ments, al though vari ably dis trib uted among sam ples and spe cies. This is proof thatthese skel e tal struc tures were in volved in a me chan i cal ac tion and per ma nently cov ered with se cre tory softtis sue. Less com mon is the re gen er a tion of whole pro cesses. It ap pears that at the be gin ning the mech a nismof re gen er a tion of pro cesses was sim i lar to that of sin gle denticles. The newly formed bud was mor pho log i −cally sim ple and some what ir reg u lar in ap pear ance (Fig. 8). How ever, a reg u lar pro cess sim i lar to its ju ve nile


Fig. 9. “Sinuosa” larva of the Famennian cono donts and evo lu tion of tim ing of its meta mor pho sis. Bases of P1 el e ments with pre −sum ably daily in cre ments are shown. A. Alternognathus beulensis Ziegler et Sandberg, 1984 from the P. trachytera Zone atŚciegnia (Wzdół Plebański, sam ple Wzd−10; ZPAL CVI/2371, see also Fig. 110B). B. Neopolygnathus communis (Branson etMehl, 1934) from the P. trachytera Zone at Kowala (sam ple Ko−15; spec i men ZPAL cXVI/2510, see also Fig. 70G).C. Tripodellus schleizius (Helms, 1963) from the P. trachytera Zone at Kowala (sam ple Ko−8; spec i men ZPAL cXVI/2932) with rel a tively late meta mor pho sis. D. T. minutus (Branson et Mehl, 1934) from slightly older strata of the same zone and lo cal ity(sam ple Ko−9a; spec i men ZPAL cXVI/2942) with ear lier oc cur ring meta mor pho sis. E. Ancyrognathus sp. n. from the K. crepida

Zone at Jabłonna (sam ple J−65; spec i men ZPAL cXVI/2577; see also Fig. 76D).

coun ter part emerged sub se quently. Ap par ently the whole set of morphogenetic ma chin ery re es tab lished,with com plex and reg u larly dis trib uted Anlagen of denticles. As the basal struc ture of the pro cess was ob vi −ously de stroyed prior to re gen er a tion, the only source of morphogenetic fac tors was the se cre tive tis sueabove. The mech a nism of the morphogentic con trol di rected by a vas cu lar net work above the el e ment waspro posed by my self and it seems un avoid able to ac cept the pres ence of a truly mechanically resistant,perhaps keratinous cup arming the internal mineral skeleton of conodont elements (Dzik 2000).

DESCRIPTION AND CLASSIFICATION OF CONODONTS

Be cause of the large num ber of taxa de scribed in this pa per for mal syn on ymy lists are not given and onlythe most im por tant syn onyms are com mented on. The strati graphic and geo graphic dis tri bu tion data re feronly to Po land and to type spec i mens and spe cies. For most of the ear lier named Famennian spe cies such data can be ex tracted from pub li ca tions in re spect to plat form P1 el e ments but it might be mis lead ing to trans ferthem me chan i cally to ap pa ra tus taxa. They have to be con firmed by ap pa ra tus stud ies, un til now per formedto a very lim ited ex tent (e.g., Klapper and Philip 1971; van den Boogaard and Kuhry 1979; Metzger 1994;Schülke 1999).

Emended di ag no ses, re fer ring to ap pa ra tus com po si tion wher ever pos si ble, are given for gen era and spe −cies (and some fam i lies). The char ac ters listed in a di ag no sis are in tended to dif fer en ti ate taxa at the samerank and only those in cluded in the same taxon of higher rank. The di ag no ses are hi er ar chi cal in the sensethat char ac ters shared by all mem bers of the higher rank taxon (plesiomorphies) are ig nored in char ac ter iz inglower rank taxa. These char ac ters may serve to di ag nose the higher rank taxon only if they are, or their com −bi na tion is, unique to the taxon. To dis tin guish spe cies homoemorphic in some as pects, so com mon amongcono donts, it it thus nec es sary to con sider the whole hi er ar chi cal set of di ag no ses, up to the tax o nomic rankuni fy ing such forms similar to each other in result of convergent evolution.

Phylum CHORDATA Bateson, 1886Class CONODONTA Eichenberg, 1930

Order Prioniodontida Dzik, 1976Fam ily Icriodontidae Müller et Müller, 1957

Di ag no sis. — P2 el e ments of the ap pa ra tus usu ally with icrion on the ex ter nal pro cess, el e ments of the Sse ries more or less re duced, sim pli fied mor pho log i cally and bear ing pro cesses with unmineralized bases ofdenticles.

Re marks. — The icriodontids have their roots in the Or do vi cian and the orig i nal as pect of their ap pa ra tus is a triramous P1 el e ment with icrion on its prob a bly ven tral (sic!) pro cess and a set of ramiform com plex el e −ments sim i lar to other prioniodontids but with a ten dency to re duce denticulation in some of them or sep a ratepar tic u lar denticles on the pro cesses that be came less and less min er al ized (e.g., Dzik 1994). At the be gin −ning of the De vo nian this change was al ready ad vanced, the an te rior pro cess in P1 el e ment be ing rep re sentedonly by an ex pan sion of the base, but the dor sal pro cess still pre served denticulation (Serpagli 1982). The De −vo nian lin eage of Latericriodus with this kind of ap pa ra tus con tin ued un til the mid Givetian rep re sentedthere by its type spe cies (Bultynck 2003). A fur ther re duc tion of the dor sal pro cess in the late Emsian re sulted in the emer gence of the lin eage of Icriodus, with only one pro cess rep re sented in the P1 el e ment (Klapper and Ziegler 1967). This branch con tin ued to the mid Famennian, in some parts of its oc cur rence rep re sented bysev eral sep a rate lin eages. Mean while ad di tional pro cesses with icrion de vel oped in off shoots of Lateri −criodus or the icrion was re placed with a sin gle row denticles in oth ers. As a re sult of the disapearance of theicrion in the late Frasnian, the lin eage of Pelekysgnathus orig i nated. Prob a bly from this lin eage emergedcono donts lack ing the P se ries el e ments at all and, in re place ment, chang ing their coni form el e ments of therest of the ap pa ra tus into ro bust struc tures, even tu ally de vel op ing a kind of icrion (Fungulodus). Rare spec i −mens from the Famennian of the Holy Cross Moun tains seem to doc u ment the evo lu tion from Latericriodusto the late Famennian Antognathus, but also the early evo lu tion of Pelekysgnathus and ap pa ra tuses com posed ex clu sively of “sim ple cones” (Fig. 121).

30 JERZY DZIK

Genus Latericriodus Müller, 1962

Type spe cies: Icriodus latericrescens Branson et Mehl, 1938.

Di ag no sis. — Fusiform icrion on the ven tral pro cess, well denticulated dor sal pro cess with a ten dency tode velop icrion on it and to re duce com pletely the ru di men tary, undenticulated an te rior process.

Re marks. — The ear lier de scribed late Famennian icriodontid spe cies with ad di tional denticulated pro −cesses, at trib uted to Antognathus (Sandberg and Ziegler 1979) dif fer from those of the early De vo nianLatericriodus mostly in their ir reg u lar shape and denticulation and in bend ing of the pos te rior pro cess in the op −po site di rec tion (ven tral, not dor sal). This may not be enough to sub stan ti ate a ge neric level dis tinc tion, as thespec i mens de scribed be low are mor pho log i cally and stratigraphically tran si tional. Un ques tion able Lateri −criodus is un known above the Givetian varcus Zone but its oc cur rence is gen er ally disjunct (Bultynck 2003)and there are re ports on its oc cur rence in the ear li est Frasnian (Sandberg et al. 1994, Aboussalam 2003).

Latericriodus sp.(Figs 10A and 121)

Ma te rial. — One spec i men.Re marks. — The early Famennian P1 el e ment from Jabłonna seems tran si tion be tween those of the lat est

Givetian Latericriodus latericrescens (Branson et Mehl, 1938) (Klapper in Ziegler 1975) and the lateFamennian Antognathus mowitzaensis (Sandberg and Ziegler, 1979) (Sandberg and Dreesen 1984), and thisnicely cor re sponds to its geo log i cal age. Latericriodus (or Antognathus) rarus sp. n. from Ostrówka andKowala may be an other con nect ing link in this long−last ing but mostly cryp tic lineage.

Oc cur rence. — The K. triangularis Zone at Jabłonna.


Fig. 10. Prim i tive Famennian icriodontids from the Holy Cross Moun tains in occlusal (A, B), K, I–L1, and pos te rior views.A. Latericriodus sp.; left P1 el e ment from the Klapperilepis triangularis Zone at Jabłonna (sam ple J−53a); spec i men ZPAL cXVI/1705. B, C. Latericriodus (or Antognathus) rarus sp. n.; left P1 (holotype) and P2 el e ments from the L. styriacus Zone atOstrówka (sam ple Ost−5); spec i mens ZPAL cXVI/1797–1798. D–U. Icriodus alternatus Branson et Mehl, 1934 from the ear li est Famennian at Płucki (D–K, sam ple Pł−36) and the K. triangularis Zone at Jabłonna (L–U, sam ple J−53a); P1 (C, I–K, M–O),P2 (E, P), and S (F–H, P–U) el e ments ZPAL cXVI/1528, 1526, 1530, 1531, 1532, 1604, 1527, 1529, 1606, 1605, 1613, 1614,1607, 1609, 1611, 1612, 1610, and 1608. Note patho log i cally multicusped S el e ments (G, H) and dif fer ence in height of the cusp

be tween ju ve nile P1 el e ments of dif fer ent age (cf. D ver sus N).

Latericriodus (or Antognathus) rarus sp. n.(Figs 10B, C and 121)

Holotype: P1 el e ment ZPAL cXVI /1797 (Fig. 10B).Type ho ri zon and lo cal ity: Sam ple Ost−5, late Famennian L. styriacus Zone at Ostrówka, Holy Cross Moun tains.Der i va tion of name: Re fer ring to rar ity of this spe cies in the Holy Cross Moun tains.

Ma te rial. — Four spec i mens.Di ag no sis. — Icrion of P1 el e ment with ad di tional denticle lat eral to the cusp.Re marks. — The spec i mens from Ostrówka and Kowala may rep re sent the first stage in de vel op ment of

an ad di tional pro cess char ac ter iz ing spe cies of Antognathus (see Sandberg and Dreesen 1984). Tak ing the P1el e ment alone, it seems likely that the pro cess de vel oped de novo and is not ho mol o gous with the dif fer entlyori ented pro cess in Latericriodus sp. How ever, the as so ci a tion of a sur pris ingly prim i tive, denticulated P2 el −e ment of seem ingly Early De vo nian mor phol ogy sug gests that this is rather a kind of late Devonian “livingfossil”.

Oc cur rence. — The late L. styriacus Zone at Kowala and Ostrówka.

Genus Icriodus Branson et Mehl, 1938Type spe cies: I. expansus Branson et Mehl, 1934 from the Givetian Mineola Lime stone near Big Springs, Mis souri

(Klapper in Ziegler 1975).

Icriodus alternatus Branson et Mehl, 1934(Figs 10D–U and 121)

Type ho ri zon and lo cal ity: Prob a bly Saverton Shale near Mon roe City, Mis souri, early Famennian (Klapper in Ziegler1975).

Ma te rial. — 10,161 spec i mens.Di ag no sis. — In icrion of P1 el e ments denticles of the mid dle row weaker than those of lat eral rows and

al ter nat ing with them; cusp trans verse to the base.Re marks. — In the ear li est Famennian sam ple Pł−36 (Fig. 10D–K) P1 el e ments of Icriodus are highly

vari able. Most of them show a fusiform icrion with small tu ber cles of the mid dle row. In nar row spec i mensthey tend to dis ap pear. The basal cav ity is gen er ally large and wide, al though in rare spec i mens it may be nar −rower, sim i lar to geo log i cally youn ger pop u la tions of the lin eage. Ju ve niles show small and straight cusp, ini −tially as so ci ated with a sin gle ven tral denticle that usu ally merges the cen tral row but fre quently it is lat er allydis placed. With growth, the cusp be came prom i nent and some what in clined, which re sults in de vel op ment of a ridge along its occlusal sur face; the re sult ing ap pear ance re stores to some de gree the sep a ra tion of the cuspfrom the icrion typ i cal of ju ve niles. In pro file view, the icrion is vir tu ally straight, only slightly con vex insome spec i mens. Con tri bu tion of such spec i mens to the some what youn ger sam ple J−53a (Fig. 10D–K)seems slightly higher. As so ci ated non−plat form el e ments range in shape from tri an gu lar to nee dle−like, withno ap par ent mor pho log i cal classes (pos si ble ex cep tions are the wid est and most ro bust spec i mens, prob a blyrep re sent ing P2 el e ments; Fig. 121). All show sharp edges and rel a tively flat ap pear ance but in sam pleWtr−27 sev eral ro bust spec i mens with subquadrate or subtriangular sec tion of the base oc cur. They dif ferfrom el e ments of much youn ger Mitrellataxis circularis in their widely expanding basal cone.

Oc cur rence. — From the base of the Famennian to the early K. crepida Zone in all stud ied lo cal i ties. The spe cies oc curs also in the Dębnik area (Baliński 1995).

Icriodus cornutus Sannemann, 1955(Figs 11 and 121)

Type ho ri zon and lo cal ity: Black lime stone with Nehdenites verneuili at Breitengrund, Frankenwald (Ziegler 1975).

Ma te rial. — 2,104 spec i mens.Di ag no sis. — Cusp of P1 el e ment in clined dor sally, nar row icrion of P1 el e ments with ir reg u larly dis trib −

uted denticles of the mid dle row, which are some times slightly higher than those of lat eral rows; in ma turespec i mens they merge to pro duce ir reg u lar transverse ridges.

Oc cur rence. — At Jabłonna, the lin eage of Icriodus re ap pears with this spe cies af ter a gap in oc cur rence, when Pelekygnathus was the dom i nant icriodontid; in Kadzielnia there seems to be a con ti nu ity be tween I.alternatus and I. cornutus within the K. crepida Zone. Tran si tional sam ples show a great pop u la tion vari abil −

32 JERZY DZIK

ity, with spec i mens of morphologies sim i lar to holotypes of I. alternatus, I. cornutus, and I. costatus oc cur −ring to gether. In sam ples J−65 and Ka−14 ro bust P1 spec i mens sim i lar to the holotype of Icriodus costatus(Thomas, 1949) from the late Famennian Ma ple Hill Shale at Burlington, Iowa (Klapper in Ziegler 1975) areas so ci ated with equally ro bust non−plat form el e ments or na mented with lon gi tu di nal striae. Ro bust P1 el e −ments of this spe cies some what re sem ble those of the Car bon if er ous Eotaphrus and per haps this may be their more appropriate generic affiliation.

Rare spec i mens from the P. trachytera Zone are prob a bly re worked. Icriodus ap par ently did not sur viveC. marginifera in the Holy Cross Moun tains area, but I. chojnicensis Matyja, 1972 from Pomerania and theLublin area (Matyja 1972, 1993; Matyja and Żbikowska 1974) may be a con tin u a tion of the lin eage in mar −ginal parts of the Variscan basin.

Genus Pelekysgnathus Thomas, 1949

Type spe cies: P. inclinatus Thomas, 1949 from the late Famennian Ma ple Mill Shale in Washigton County, Iowa.

Di ag no sis. — Icriodontids with P1 el e ments bear ing only the ven tral pro cess (as in Icriodus) with a sin gle row of denticles.

Pelekysgnathus planus Sannemann, 1955(Figs 12A–Q and 121)

Type ho ri zon and lo cal ity: Black lime stone with Nehdenites verneuili at Breitengrund, Frankenwald (Sannemann 1955a).

Ma te rial. — 1,113 spec i mens.Di ag no sis. — The blade of P1 el e ments straight in pro file view or gently con vex, with the cusp not sep a −

rated from other denticles or slightly higher but al most straight.Re marks. — The early form of Pelekysgnathus planus from the ear li est Famennian shows a lin ear pro file

of the blade with a weak denticulation and the cusp not sep a rated from other denticles, sim i larly as in theFrasnian (Dzik 2002). In sam ples of some what youn ger age, for in stance J−53a (Fig. 12A), along with suchspec i mens a new mor phol ogy emerged, with a dom i nant cusp and sharp, sep a rate denticles. The blade re −mains short, how ever, which only par tially re sults from gen er ally small size of el e ments. In later (typ i cal)


Fig. 11. Mid Famennian icriodontid Icriodus cornutus Sannemann, 1955 from Kadzielnia in the Holy Cross Moun tains.A–C. An ces tral pop u la tion from the late K. triangularis (sam ple Ka−16) with al most ver ti cal cusp, tran si tional from I. alternatus; P1 (A, B) and P2 (C) el e ments ZPAL cXVI /1646, 1648, and1647. D–G. Ad vanced pop u la tions with prom i nent and strongly inclined cusp from the K. crepida Zone (D, E, sam ple Ka−2; F, G, Ka−14); P1 (D–F) and P2 (G) el e ments ZPAL cXVI /1650,

1649, 1644, and 1643, re spec tively. Note striation of coni form el e ments some what re sem bling that in Mitrellataxis.

forms with a gently in clined cusp, the pro file of the blade is con vex, sim i lar as in the holotype (Sannemann1955a). Ma ture ro bust el e ments show a weak denticulation. As so ci ated non−plat form el e ments (e.g., J−03)rep re sent two sep a rate classes of tri an gu lar el e ments (prob a bly P2) wider than in Icriodus alternatus andmore robust needle−like elements.

Oc cur rence. — Con tin ues from the lat est Frasnian to the C. marginifera Zone (Ł−13); spec i mens fromthe P. jugosus Zone at Ostrówka are prob a bly re worked.

Genus ?Dollymae Hass, 1959Type spe cies: D. sagittula Hass, 1959 from the late Tournaisian Chappel Lime stone of Texas.

Di ag no sis. — Dor sal pro cess in P1 el e ments with a ingle row of denticles and the basal cav ity ex pandedon both sides up to de vel op ing denticulated processes.

Dollymae? guizhouensis Wang et Wang, 1978(Figs 12R and 121)

Type ho ri zon and lo cal ity: Late Famennian of Daihua For ma tion at Daihua, Guizhou, China; as so ci ated with Wocklumeriaand Bispathodus (Wang and Wang 1978).

Ma te rial. — Six spec i mens.Di ag no sis. — Undenticulated, tranversely wid ened basal cone in P1 el e ments, ad di tional denticulation

ven trally of the cusp.Re marks. — The in cip i ent denticulation be low the cusp makes this very rare lat est Famennian spe cies un −

like other spe cies of Pelekysgnathus. As no ticed al ready by Over (1992), the lat eral ex pan sion of the base sug −gests that this may be the be gin ning of the lin eage rep re sented in the Tournaisian by Dollymae bouckaertiGroessens, 1977 and Eotaphrus bultyncki (Groessens, 1977), sub se quently di ver si fied into elab o rate Carboni −ferous forms.

Oc cur rence. — The D. trigonica Zone at Kowala and Dzikowiec.

Family Jablonnodontidae fam. n.Di ag no sis. — Ap pa ra tus com posed of only coni form el e ments mim ick ing those of the Or do vi cian

protopanderodontids.

34 JERZY DZIK

Fig. 12. Icriodontids with out icrion from the Famennian of the Holy Cross Moun tains and Sudetes. A–Q. Pelekysgnathus planusSannemann, 1955 from the early Famennian at Jabłonna (A, H, sam ple J−53a; B–G, I, J, sam ple J−5,) and Karczówka (K–Q);P1 (A–E, K–M) and P2 (F, N ) and S (G–J, O–Q) el e ments ZPAL cXVI /1615, 1618, 1620, 1619, 1621, 1622, 1623, 1617, 1624,1616, 1628, 1630, 1629, and 1631–1634, re spec tively. R. Dollymae? guizhouensis Wang et Wang, 1978 from the late Famennian

at Dzikowiec (sam ple Dz−48); P1 el e ment ZPAL cXVI /1645.

Re marks. — In ter pre ta tion of taxomic af fin i ties of the Famennian “sim ple cones” is a dif fi cult task. Thegeo log i cally youn gest cono dont be ing un ques tion ably a con tin u a tion of the early Pa leo zoic coni form cono −donts is Belodella, the panderodontid which did not sur vive to the Famennian. The last oc cur rences of thatperi−reefal spe cies are lat est Frasnian in age (e. g., Dzik 2002). The seem ingly coni form cono dont el e mentsof the co eval icriodontids are mostly iso lated denticles of el e ments with oth er wise unmineralized pro cesses.At least three lin eages of the icriodontids per sisted into the Famennian. Their coni form denticles are of vir tu −ally the same mor phol ogy as those of geo log i cally older spe cies, be ing as so ci ated with icrion−bear ing P1 el e −ments. Yet, “sim ple cones” oc cur in abun dance also in ho ri zons and whole sec tions of the Famennian whereicriodontid P1 el e ments are miss ing. Some of them are very sim i lar to the Or do vi cian protopandero dontids or distacodontids. The Famennian Mitrellataxis of Chauff and Price (1980; growth in cre ments within the basalcav ity show that this is a cono dont, not a fish scale, as sug gested for the re lated Fungulodus by Wang andTurner 1985) is a homeomorph of the early Or do vi cian Clavo hamulus. El e ments in dis tin guish able fromthose of the Or do vi cian Drepanoistodus have been de scribed as parts of the ap pa ra tus of FamennianPelekysgnathus (Sandberg and Dreesen 1984). In the Holy Cross Moun tains such el e ments oc cur abun dantly in some sam ples (Ta ble 1). No re cur rent as so ci a tion with the P1 el e ments of Pelekysgnathus or Icriodus hasbeen no ticed, and wher ever those are rep re sented in a rea son able num ber, they are as so ci ated with flat andwide tri an gu lar el e ments like those from older strata (e.g., Dzik 2002).

It is tempt ing to as sume that these are cases of fur ther re duc tion in min er al iza tion of the icriodontid oralap pa ra tus. Per haps in some cases this ac tu ally took place. Nev er the less, some other as so ci a tions of coni formel e ments show a re cur rent mor pho log i cal dif fer en ti a tion closely re sem bling that in the early Palaeozoicprotopanderodontid cono donts. Al though not all el e ment types rep re sent truly dis crete classes and mor pho −log i cal tran si tions are more com mon than in un doubted protopanderodontids, the sim i lar ity is strik ing andex tends even to de vel op ment of geniculation in some el e ments, mim ick ing the Or do vi cian distacodontids(Sandberg and Dreesen 1984). A pos si bil ity of such an af fin ity can not be ex cluded, as much of theFamennian cono dont fauna is of cryp tic or i gin. It emerged as an ef fect of ex ten sive mi gra tion events and thepres ent knowl edge of the Si lu rian and Devonian simple cones is still very limited outside restricted regions of Europe.

The al ter na tive, that this is a case of sec ond ary in tro duc tion of reg u lar ity to ear lier rather in dif fer entmorphologic vari abil ity of the icriodontid denticles is sup ported by the rel a tively late emer gence of thesecono donts and the tre men dous time gap sep a rat ing them from the last in dis put able protopanderodontids (endof the Or do vi cian). If these were sec ond arily sim pli fied icriodontids, then all the plat form se ries lo ca tionswere either lost, or reduced to the cusp.

There are thus two pos si ble so lu tions to the prob lem of the Famennian “sim ple−cones”: (1) these aredenticles of el e ments from an icriodontid lin eage with com pletely re duced P el e ments or (2) these are sur −vi vors of the Or do vi cian protopanderodontids, that is a Laz a rus lin eage not rec og nized pre vi ously from the whole Si lu rian and most of the De vo nian. Un til this is sue is set tled, the ma te rial of sim ple cones not be −long ing to Icriodus and Pelekysgnathus is here de scribed pro vi sion ally in terms of coni form ap pa ra tus ter −mi nol ogy.

Genus Mitrellataxis Chauff et Price, 1980

Type spe cies: M. chevronella Chauff et Price, 1980 from the Ma ple Mill Shale at Kalona Clay Pit, Iowa; Scaphignathusvelifer Zone.

Di ag no sis. — Ro bust coni form el e ments with shal low or in verted basal cav i ties of cir cu lar outline.Re marks. — The Famennian “sim ple cones” are of three kinds: some are or na mented with prom i nent

lon gi tu di nal ribs (typ i cal Mitrellataxis), oth ers show a vir tu ally smooth sur face (Jablonnodus), yet oth ersde vel oped ir reg u lar tuberculation with ameloblasts im prints, thus de vel op ing a kind of icrion (Fun −gulodus–Conchodontus; Over 1992). A fine striation oc curs also in ro bust icriodontid denticles (Over1992) al though there seems to be a morphologic gap be tween them and Mitrellataxis. A sim i lar gap sep a −rates smooth icriodontid denticles and el e ments of M. circularis, re sem bling rather denticles of ro bustprioniodinids. Mitrellataxis conoidalis may be the con nect ing link be tween ro bust coni form el e ments ofIcriodus and more de rived spe cies of its ge nus; this is the sole rea son why it is clas si fied among thePrioniodontida.


Mitrellataxis conoidalis sp. n.(Figs 13V–X and 121)

Holotype: Spec i men ZPAL cXVI/1627 (Fig. 13V).Type ho ri zon and lo cal ity: Bed 5, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of the name: Re fer ring to conoidal shape of el e ments.

Ma te rial. — 34 spec i mens.Di ag no sis. — All el e ments with a sharp con i cal cusp or na mented by lon gi tu di nal ribs; rel a tively deep

basal cavity.Re marks. — From M. chevronella and M. ornata, the new spe cies dif fers in hav ing only con i cal el e −

ments, from M.? asymmetrica and spe cies of Jablonnodus in pres ence of lon gi tu di nal ribs.Oc cur rence. — The late K. triangularis (J−37) to the K. crepida zones at Kadzielnia, Jabłonna, and

Wietrznia.

Mitrellataxis ornata sp. n.(Figs 13Y–GG and 121)

Holotype: Spec i men ZPAL cXVI/1640 (Fig. 13FF).Type ho ri zon and lo cal ity: Sam ple of cephalopod lime stone taken by Dymitr Sobolew, early Famennian late K. triangularis

Zone at Karczówka, Holy Cross Moun tains.

36 JERZY DZIK

Fig. 13. Pos si ble Famennian icriodontids with com pletely re duced plat form se ries in the ap pa ra tus. A–K. Mitrellataxis?asymmetrica sp. n. from the L. styriacus Zone at Ostrówka (sam ple Ost−12); spec i mens ZPAL cXVI /1575, 1567, 1576, 1573(holotype, D), 1574, 1572, 1571, 1569, 1568, 1570, 1566, re spec tively. L–T. Mitrellataxis circularis (Wang et Wang, 1978)from the same sam ple and from Kowala (T, sam ple Ko−8), spec i mens ZPAL cXVI /1578–1580, 1584, 1585, 1583, 1581, 1582,and 1577. U. Mitrellataxis coronella Chauff and Price, 1980 from the P. jugosus Zone at Kowala (sam ple Ko−63); spec i menZPAL cXVI /2334; V, X. Mitrellataxis conoidalis sp. n. from the K. crepida Zone at Jabłonna (bed 5); spec i mens ZPAL cXVI/1627 (holotype, V), 1625 and 1626. Y–GG. Mitrellataxis ornata sp. n. from the late K. triangularis Zone at Karczówka, (Y–FF) and early Famennian at the Żbik ra vine in Dębnik near Cra cow; spec i mens ZPAL cXVI/1636, 1635, 1637, 1642, 1641, 1638,

1639, 1640, and 2963, re spec tively (holotype, FF).

Der i va tion of the name: Re fer ring to or nate rib bing of the cusp.

Ma te rial. — 35 spec i mens.Di ag no sis. — Most spec i mens with blunt apex and ovoid ap pear ance of the cusp, rare con i cal el e ments;

all bear sharp lon gi tu di nal ribs merg ing in a chev ron pat tern on the ex ter nal sur face of the cusp or pass ingcon tin u ously across the tip.

Re marks. — From the mor pho log i cally clos est M. chevronella dif fers in a more reg u lar and less ro bustap pear ance of cusps and usu ally flat or con cave basal cav ity. In the type sam ple of M. ornata sharply con i calel e ments also oc cur. Whether they be long to the same ap pa ra tus or rep re sent sympatric oc cur rence of M.conoidalis re quires testing with more material.

Oc cur rence. — The type lo cal ity and early Famennian of Żbik ra vine in Dębnik near Cra cow.

Mitrellataxis? asymmetrica sp. n.(Fig. 13A–K)

Holotype: Spec i men ZPAL cXVI/1673 (Fig. 13D).

Type ho ri zon and lo cal ity: Sam ple Ost−12, late Famennian L. styriacus Zone at Ostrówka, Holy Cross Moun tains.

Der i va tion of the name: Re fer ring to the asym me try of most el e ments.

Ma te rial. — 52 spec i mens.Di ag no sis. — Sharp con i cal cusp with len tic u lar sec tion, smooth sur face and sharp edges.Re marks. — From typ i cal spe cies of Mitrellataxis, the new spe cies dif fers in a smooth sur face of the

cusp, from M. circularis in sharp−edged cusp and gen er ally nar row base. The idea that the Famennian “sim −ple cones” are iso lated denticles of sec ond arily sim pli fied icriodontids fits es pe cially this spe cies. The el e −ments show lin early ex tend ing bases, quite long in some spec i mens, and they may be eas ily visu al ised as be −ing dis trib uted along a non−minerized process.

El e ments of M.? asymmetrica re sem ble most the coni form el e ments of Icriodus (e.g., that on Fig. 10U)differring only in that no in cip i ent denticles or angu la tions de velop near the bese, as is usual in at least someel e ments of Icriodus. This is thus ei ther a case of homeomorphy or the spe cies is the end−mem ber of anicriodontid lin eage with re duced P1 el e ments. Wang (1989) il lus trated a sim i lar el e ment from the Poly −gnathus perbonus Zone of the Emsian un der the name Drepanodina subcircularis Wang, 1981. This maymean that the lineage was long−lasting.

Oc cur rence. — Sparsely oc cur ring in many sam ples rang ing from the P. trachytera Zone at Kowala andJabłonna to the L. styriacus Zone at Ściegnia and Ostrówka.

Mitrellataxis circularis (Wang et Wang, 1978)(Figs 13L–T and 121)

Type ho ri zon and lo cal ity: Late Famennian of Daihua For ma tion at Daihua, Guizhou, China; as so ci ated with Wocklumeriaand Pseudopolygnathus.

Ma te rial. — 3,295 spec i mens.Di ag no sis. — Cusp with oval cross sec tion, el e ments vari able in shape but not form ing dis crete morpho −

logic classes, shal low basal cav ity usu ally filled with tissue.Re marks. — De spite gen eral sim i lar ity of smooth el e ments, re la tion ship of this spe cies to Jablonnodus is

not likely be cause they do not group in any re cur rent morphologic classes. They rather re sem ble in col ourand shape cusps and denticles of as so ci ated ro bust prioniodinids. It can not be ex cluded that this is ac tu ally aprioniodinid with uncalcified bases of pro cesses. That these were truly denticles of uncalcified pro cesses oficriodontids or prioniodinids is sup ported by the lin ear elon ga tion of the basal body, with a lon gi tu di nal fur −row (Fig. 13T).

The only il lus trated spec i men of Drepanodus circularis Wang et Wang, 1978 has a short cusp, whichmakes it sim i lar to M. conoidalis sp. n., so the tax o nomic iden ti fi ca tion pro posed here re quires con fir ma tionby the Chi nese topotype ma te rial.

Oc cur rence. — Ranges from the late K. triangularis Zone to the top of the Famennian in al most all lo −cal i ties.


Mitrellataxis coronella Chauff et Price, 1980(Figs 13U and 121)

Type ho ri zon and lo cal ity: Un named basal shale mem ber of the Sul phur Springs For ma tion, prob a bly lower costatus Zone(Chauff and Price 1980).

Ma te rial. — Sin gle spec i men from sam ple Ko−116 at Kowala, P. jugosus Zone.Di ag no sis. — Waistlike con stric tion near the base, small low cusp in the mid dle of flat occlusal sur face,

rel a tively deep basal cavity.Re marks. — The only spec i men found has the occlusal sur face par tially cov ered with clay but the con i −

cal tip in the cen tre and ribs around the mar gin are clearly rec og niz able.

Genus Jablonnodus gen. n.Type spe cies: J. oistodiformis sp. n. from the Famennian of the Holy Cross Moun tains, Po land.

Di ag no sis. — Ten dency to de velop geniculate “oistodontiform” and sym met ri cal “erectiform” el e mentsin the ap pa ra tus, homeomorphic with the Or do vi cian Distacodontidae; weak morphologic dis par ity be tweenprob a ble P and S series.

Re marks. — Mound (1968) and Wang and Wang (1978) il lus trated sev eral such el e ments un der the ge −neric name Drepanodina. Un for tu nately, the holotype of Drepanodina lachrymosa Mound, 1968 from thelate Famennian Up per Wabamun Group of Al berta, the type spe cies of the ge nus, is prob a bly a denticle ofPelekysgnathus, as sug gested by the ap pear ance of its base.

Jablonnodus oistodiformis sp. n.(Figs 14 and 121)

Holotype: M el e ment ZPAL cVI/1557 (Fig. 14M).Type ho ri zon and lo cal ity: Bed 24, mid Famennian P. trachytera Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: Re fer ring to some sim i lar ity of one el e ment in the ap pa ra tus to that of the Or do vi cian ge nus Oistodus.

Ma te rial. — 934 spec i mens.

38 JERZY DZIK

Fig. 14. Sim ple−cone Jablonnodus oistodiformis sp. n. from the early P. trachytera Zone at Jabłonna (bed 24; A–N) and the L.styriacus Zone at Ostrówka (O–U, sam ple Ost−12) in the Holy Cross Moun tains; all in pos te rior view. A, B, O, P. Right and leftP1? el e ments; spec i mens ZPAL cXVI/1545, 1546, 1560, and 1564, re spec tively. C, D, Q. Right and left P2 (?) el e ments 1546,1548, and 1559. E, F, R, S. S0? el e ments 1549, 1550, 1562 and 1561. G–L, T. Right and left S1–4? el e ments 1551–1557 and 1563.

M, N, U. Right and left M? el e ments 1557, 1558, and 1565.

Di ag no sis. — Pos si ble M el e ments al most geniculate, cusp of most el e ments with sharp edges and baseswith oval con tour.

Re marks. — El e ments of the spe cies are char ac ter is tic in their Drepanoistodus−like ap pear ance and are easyto sep a rate from other as so ci ated sim ple−cones. It is not eco log i cally as so ci ated with spe cies of Mitrel lataxis with smooth el e ments and sam ples rich in the lat ter do not con tain J. oistodiformis. Iden ti fi ca tion of dis crete el e menttypes of the ap pa ra tus was pos si ble ow ing to their abun dance in sam ples Ost−12 and J−24 (Ta ble 17).

El e ments ten ta tively iden ti fied as be long ing to the plat form se ries have a strongly bent cusp. They formtwo classes dif fer ing in out line of the base (Fig. 121), those with rounded tri an gu lar out line are pro vi sion allypro posed to be P1 el e ments, base out line of pos si ble P2 el e ments is cir cu lar. El e ments with a rather erect cuspshow the sym me try tran si tion se ries from the strictly sym met ri cal prob a ble S0 el e ments through those withthe base flat tened from one side (S1–2) to those hav ing flat ten ing on both sides (S3–4). A sep a rate class is rep −re sented by prob a ble M el e ments with the whole cusp strongly inclined to the base and very asymmetric.

From other spe cies of the ge nus, J. oistodiformis dif fers in the shape of pos si ble M el e ments and rel a tively dis tinct classes of the el e ment types (at least P, S, and M). Sandberg and Dreesen (1984, pl. 3: 10, 11, 23, 24)il lus trated even more ad vanced, fully geniculate el e ments as so ci ated with equally de rived other el e mentsfrom the late Famennian (expansa Zone) of Col o rado and Eng land. In each of those lo cal i ties they co−oc curwith dif fer ent icriodontids. The lin eage seems thus to have been evolving rather fast.

Oc cur rence. — Prob a bly the late C. marginifera Zone at Jabłonna, Miedzianka and Łagów, cer tainly theP. trachytera Zone at Jabłonna, Kowala, Łagów and Ostrówka, pos si bly con tin ued to the early L. styriacusZone at Jabłonna and Ostrówka.

Jablonnodus erectus sp. n.(Figs 15 and 121)

Holotype: P1 el e ment ZPAL cVI/1587 (Fig. 15B).


Fig. 15. Sim ple−cone Jablonnodus erec tus sp. n. from the late Lagovignathus styriacus Zone at Ostrówka (A–L, N–Q, sam pleOst−265) and the early P. jugosus Zone at Jabłonna (M, bed 27) in the Holy Cross Moun tains; in pos te rior, ven tral (I1) and me dial(L2 and M2) views. A, B. Left P1? el e ments; spec i mens ZPAL cXVI /1586−1587, re spec tively. C, D. Right P2 (?) el e ments 1588,1589. E, F, H–L. Right and left S2–4? el e ments 1591, 1593, 1596, 1600, 1590, 1592, and 1597. G, M. S0? el e ments 1602 and 1601.

N–O. Right S1? el e ments 1598, 1599. P, Q. Left M? el e ments 1595 and 1594.

Type ho ri zon and lo cal ity: Sam ple Ost−284.II.265, late Famennian L. styriacus Zone at Ostrówka, Holy Cross Moun tains.Der i va tion of name: Re fer ring to erect cusp in the sym met ri cal el e ment of the ap pa ra tus.

Ma te rial. — 937 spec i mens.Di ag no sis. — Pos si ble M el e ments with erect cusp, bases of most el e ments with ir reg u larly eye drop con −

tour.Re marks. — Sam ple Ost−265 of fers enough ma te rial to al low iden ti fi ca tion of dis crete el e ment types and

their homology with those of the stratigraphically older J. oistodiformis. Prob a ble P1 el e ments bear widesharp ridges and have a char ac ter is tic out line of the base, acutely nar row ing at ends and swol len in the mid dle but with one side strongly de pressed. Such a de pres sion is pres ent in the out line of the oth er wise roundedbase of rather gracile prob a ble P2 el e ments (Fig. 121). El e ments of the sym me try tran si tion se ries are sim i larto those of J. oistodiformis ex cept for be ing some what more robust; probable M elements are less inclined.

From the other spe cies of the ge nus, J. erec tus dif fers in the shape of pos si ble M el e ments and the widecusp of prob a ble P2 el e ments. Their basal cav ity is rel a tively shal low and wide. P el e ments show also a char −ac ter is tic ex ter nal extension of the base.

Oc cur rence. — Pos si bly the top of the K. crepida Zone at Jabłonna, the C. marginifera and P. trachyteraZone at Łagów, the L. styriacus Zone at Ostrówka, Miedzianka, and Ściegnia; the P. jugosus Zone at Kowala and Ostrówka.

Order Ozarkodinida Dzik, 1976Family Prioniodinidae Bassler, 1925

Ge nus Ligonodina Bassler, 1925Type spe cies: L. pectinata Bassler, 1925 from the mid Frasnian of New York (Hud dle 1968).

Di ag no sis. — The anteriormost el e ment of the ap pa ra tus (M) with a wide and flat cusp.

40 JERZY DZIK

Fig. 16. Spe cif i cally un de ter mined early Famennian prioniodinid Ligonodina sp. from the K. triangularis Zone at Jabłonna(sam ple J−58) in the Holy Cross Moun tains. P1 (A), P2 (B), S0 (D), S1 (C), S2 (G?), S3–4 (E, F), and M (H) el e ments; spec i mens

ZPAL cXVI/11728, 1729, 1731, 1730, 1733, 1734, 1732, and 1735, re spec tively.

Ligonodina pectinata Bassler, 1925(Figs 16?, 17A, B, and 122)

Type ho ri zon and lo cal ity: Rhinestreet Shale Mem ber of the West Falls For ma tion at Weyer, New York (Hud dle 1968).

Ma te rial. — 419 spec i mens, some of doubt ful spe cific af fin ity.Di ag no sis. — Prom i nently denticulated dor sal pro cess and large tri an gu lar cusp of the M el e ment, rel a −

tively shal low basal cav i ties of all elements.Re marks. — Iden ti fi ca tion of this spe cies in the ear li est Famennian is based on rather weak ev i dence:

one ju ve nile M el e ment from sam ple Pł−40 (Fig. 17B), which may be an ex treme va ri ety of Cteno −polygnathus and a ro bust but teratologically de formed spec i men from sam ple Pł−1. This re fers also to thegen er al ized prioniodinids from the K. triangularis Zone at Jabłonna (Fig. 16). In youn ger Famennian stratafrom other lo cal i ties much more de rived prioniodinids are rep re sented. Un for tu nately, other el e ments in theap pa ra tus are usu ally of lit tle di ag nos tic value and even if sev eral names are avail able for the Famennianprioniodinids, among them Ligonodina delicata Branson et Mehl, 1934, with the type oc cur rence in theSaverton Shale at Sees Creek, Mis souri (Mound 1968 in tro duced a new name for it be cause of homonymy atthat time, but there is no lon ger need for it). From the same rea son sev eral oc cur rences of the species areidentified only tentatively.

In the Frasnian pop u la tions of the spe cies there is some un cer tainty re gard ing dis tinc tion be tween S1 andP1 el e ments, both show ing a wide pop u la tion vari abil ity. Po si tion in the ap pa ra tus of the el e ments with shortand strongly twisted ven tral pro cess and rel a tively high cusp (Dzik 2002, figs 7A and 8A, B) re mains con tro −ver sial. In the lat est Frasnian sam ples un doubted P1 el e ments have been iden ti fied (Dzik 2002, fig. 8C), withro bust denticles, a high cusp, and arched dor sal pro cess. In the Famennian their homo logues show a lessprom i nent denticulation but more blade−like ap pear ance. This may be a case of directional evolution.

In the basalmost Famennian sam ple at Miedzianka (Md−27) wide M el e ments closely sim i lar to those ofL. albidens sp. n. co−oc cur with ro bust S3–4 el e ments re sem bling some what those of Idioprioniodus un −cadino ides sp. n. This may be a pop u la tion tran si tional be tween L. pectinata and L. albidens but the ma te rial(con glom er ate!) may con tain re worked Frasnian specimens as well.


Fig. 17. Early Famennian suc ces sors of the main prioniodinid lin eage of Ligonodina from the Holy Cross Moun tains. A, B. Ligo −nodina pectinata Bassler, 1925 from the ear li est Famennian at Płucki (sam ple Pł−40); S4 (A) and M (B) el e ments ZPALcXVI/1712, 1713. C–T. Ligonodina albidens sp. n. from the late K. triangularis and K. crepida zones at Jabłonna (bed 7, C; J−6,D–G; J−45a, H–N) and Wietrznia (M–T, sam ple Wtr−21); P1 (H and P), P2 (C and I), S0 (J and R), S1 (K?), S2–4 (D, F, L, O, N, Q,and S), and M (E, G, M, and T) el e ments; spec i mens ZPAL cXVI/1722, 1725, 1726, 1724, 1727, 1736–1739, 1742, 1743

(holotype, M), 1740, 1741, 1750, 1753, 1751, 1752, and 1754, re spec tively.

Oc cur rence. — Only the ear li est Famennian at Płucki, pos si bly also the late K. triangularis Zone atJabłonna and Miedzianka.

Ligonodina albidens sp. n.(Figs 17C–T and 122)

Holotype: Spec i men ZPAL cXVI/1743 (Fig. 17M).Type ho ri zon and lo cal ity: Sam ple J−45a, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of the name: The name re fers to albid col or ation of denticles in this spe cies.

Ma te rial. — 417 spec i mens.Di ag no sis. — Short tri an gu lar pro cess of M el e ment, ex ter nal pro cesses of the sym me try tran si tion el e −

ments few in number.Re marks. — The wide cusp of M el e ments in di cates re la tion ship of the spe cies to the Ligonodina main lin −

eage but re main ing el e ments of the ap pa ra tus are highly de rived mor pho log i cally. This spe cies is rather eas ilyrec og niz able ow ing to sparsely dis trib uted prom i nent denticles of adult el e ments and short ex ter nal pro cessesof S el e ments, only sin gle denticles rep re sent ing them in the S0 el e ment. In this re spect and gen eral ap pear anceof non−M el e ments, the spe cies re sem bles the geo log i cally youn ger Lagovidina obliqua sp. n. but re la tion shipbe tween these spe cies re mains un clear and may be a re sult of homeomorphy. The spe cies oc curs both in thedeeper−wa ter area of Jabłonna and shal lower en vi ron ment of Wietrznia lo cated above the Frasnian mudmound.

Oc cur rence. — The late K. triangularis and K. crepida zones at Jabłonna, Wietrznia, Kadzielnia andMiedzianka.

Ligonodina multidens sp. n.(Figs 18 and 122)

Holotype: Spec i men ZPAL cXVI/1659 (Fig. 18I).Type ho ri zon and lo cal ity: Sam ple Md−1, early Famennian late K. triangularis Zone at Miedzianka, Holy Cross Moun tains.Der i va tion of the name: Re fer ring to rel a tively nu mer ous denticles on M el e ment.

Ma te rial. — 25 spec i mens.Di ag no sis. — Straight and rel a tively long, al most trans versely ori ented pro cess of M el e ment with

denticles of rel a tively uni form height.

42 JERZY DZIK

Fig. 18. Gen er al ized early Famennian prioniodinid Ligonodina multidens sp. n. from Kadzielnia (A, F, and H, sam ple Ka−2;B–E, and G, sam ple Ka−3) and Miedzianka (I, sam ple Md−1) in the Holy Cross Moun tains. P1 (A), P2 (B), S0 (C), S1 (D), S2 (E),

S3–4 (F), and M (G–I) el e ments; spec i mens ZPAL cXVI/1651–1656, 1658, 1657, and 1759 (holotype, I), re spec tively.

Re marks. — The ap pa ra tus of the spe cies is rather gen er al ized in ap pear ance and may be a suc ces sor of a Frasnian lin eage. Avail able ma te rial is rather lim ited and in some cases it is dif fi cult to de cide to which of as −so ci ated prioniodinid spe cies par tic u lar el e ments be long. How ever, the mor phol ogy of M el e ments seemsdis tinc tive enough to sub stan ti ate sep a ra tion at the spe cies level; its wide and trans versely ori ented processsupports the generic affiliation.

Oc cur rence. — The late K. triangularis to K. crepida zones at Miedzianka and Kadzielnia.

Ligonodina latibasalis sp. n.(Figs 19 and 122)

Holotype: Spec i men ZPAL cXVI/1749 (Fig. 19F).Type ho ri zon and lo cal ity: Sam ple J−45a, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of the name: The name re fers to wid ened bases of most el e ments.

Ma te rial. — 153 spec i mens.Di ag no sis. — Re duced dor sal pro cess of M el e ment with a few low denticles, widely gap ing basal cav i −

ties of P2 el e ment and short ened pro cesses of S el e ments with few large denticles.Re marks. — The mor pho log i cally sim pli fied ap pear ance of el e ments makes ge neric af fil i a tion of this

prob a bly paedomorphic prioniodinid rather uncertain. Oc cur rence. — The K. crepida Zone at Jabłonna, Wietrznia and prob a bly Łagów.

Ligonodina sudetica sp. n.(Figs 20 and 122)

Holotype: Spec i men ZPAL cXVI/1887 (Fig. 20I).Type ho ri zon and lo cal ity: Sam ple Dz−8, late Famennian P. jugosus Zone at Dzikowiec, the Sudetes.Der i va tion of the name: Re fer ring to the Sudetes Moun tains, where the ma te rial was found.

Ma te rial. — 257 spec i mens.Di ag no sis. — High arched el e ments with flat sharp denticles, M el e ments with tri an gu larly nar rowed

base, undenticulated, pointed ex ter nal pro cess and vari able length and ori en ta tion of in ter nal pro cess.Re marks. — Be cause of the un usu ally wide range of vari abil ity it is not cer tain if truly only one

prioniodinid spe cies is rep re sented in the late Famennian of Dzikowiec. How ever, there seems to be a com −plete gra da tion be tween ex treme morphologies of the most char ac ter is tic M el e ments (Fig. 20I–L). Also thege neric af fil i a tion of the spe cies is un cer tain and it may as well be long to Idioprioniodus. In fact, some of itsM el e ments re sem ble those of Ligonodina pectinata, oth ers Idioprioniodus ruptus sp. n. The most char ac ter −is tic as pect of most el e ments of L. sudetica is elon ga tion of their cusps and sharp denticles and rel a tivelyshort pro cesses.

Oc cur rence. — The P. jugosus and D. trigonica zones at Dzikowiec and pos si bly Kowala.


Fig. 19. Sim pli fied early Famennian prioniodinid Ligonodina latibasalis sp. n. from Jabłonna (A–F, sam ple J−45a) andWietrznia (G, sam ple Wtr−32) in the Holy Cross Moun tains. P1 (A), P2 (B), S0 (C), S1–2 (D), S3–4 (E), and M (F, G) el e ments;

spec i mens ZPAL cXVI/1744–1749 (holotype, F), and 1755, re spec tively.

Genus Pluckidina Dzik, 2002Type spe cies: Pluckidina lagoviensis Dzik, 2002 from the late Frasnian of the Holy Cross Moun tains, Po land.

Di ag no sis. — El e ments of the ap pa ra tus with a ten dency to an gu lar lat eral bend ing, S3 and S4 el e mentsmor pho log i cally dis tinct, with long and straight outer processes.

Pluckidina lipperti (Bischoff, 1956)(Figs 21 and 123)

Type pop u la tion: Up per Kellwasserkalk of Schmidt’s quarry near Braunau.

Ma te rial. — 183 spec i mens.Re marks. — The ear li est Famennian Pluckidina is of a rather gen er al ized mor phol ogy (Fig. 21A–F). It

may be dis tinct tax o nom i cally from the lat est Frasnian type pop u la tion of the spe cies but with so few spec i −mens avail able it is dif fi cult to iden tify pos si ble dif fer ences. The ma te rial seems to fit within the range of itsvari abil ity. The pop u la tion from the K. crepida Zone at Jabłonna (Fig. 21G–N) shows a some what more ro −bust denticulation. Whether this re flects an evo lu tion ary change or only ecophenotypic adaptation remains to be elucidated.

Co eval, but usu ally allopatric, spe cies of Ligonodina dif fer from those of Pluckidina mostly in the shapeof the high arched S1 el e ments (also M el e ments) and gen er ally more del i cate denticulation of S el e ments.The an gu lar appearence of S1 el e ments is an in di ca tion of the prox im ity to early gondolellids.

Oc cur rence. — From the be gin ning of Famennian to the early K. triangularis Zone at Płucki and the lateK. triangularis to K. crepida zones at Jabłonna.

Pluckidina purnelli sp. n.(Figs 22 and 123)

Holotype: Spec i men ZPAL cXVI/1681 (Fig. 22G).Type ho ri zon and lo cal ity: Bed 24, mid Famennian P. trachytera Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: In rec og ni tion of Mark Purnell’s re search on prioniodinid ap pa ra tuses.

44 JERZY DZIK

Fig. 20. Prob lem atic late Famennian prioniodinid Ligonodina sudetica sp. n. from Dzikowiec, the Sudetes (A, D, H, and L, sam −ple Dz−9; B, C, F, and I, sam ple Dz−8; E, G, J, and K, sam ple Dz−10). P1 (A), P2 (B), S0 (C, D), S1 (E), S2 (F, G), S3–4 (H), andM (I–L) el e ments; spec i mens ZPAL cXVI/1877, 1886, 1885, 1878, 1880, 1884, 1881, 1879, 1887 (holotype, I), 1883, and 1876,

re spec tively.

Ma te rial. — 83 spec i mens.Di ag no sis. — Straight, al most trans versely ori ented in ner pro cess of M el e ment with prom i nent

denticles, rel a tively long ex ter nal pro cess of S3–4 el e ments ex tend ing al most along the same line as thecusp.

Re marks. — This is a rather gen er al ized prioniodinid but the shape of the M el e ment makes it dif fer entfrom all other Famennian prioniodinids oc cur ring in the re gion. Also the del i cate and mostly al ter nat ingdenticulation of el e ments is rather un usual for the prioniodinids en abling rather easy sep a ra tion of its el e −ments from those of as so ci ated mem bers of the fam ily. The spe cies has a rather short range of oc cur rence inthe Holy Cross Moun tains but prob a bly was rep re sented in the whole area, as it has been found at Jabłonnaand Ostrówka, which are relatively distant from each other.

Oc cur rence. — The P. trachytera Zone at Jabłonna and the L. styriacus to P. jugosus zones at Ostrówka.

Genus Lagovidina gen. n.Type spe cies: Lagovidina obliqua sp. n.

Di ag no sis. — In verted basal cav ity in all el e ments, M el e ment with a short dor sal pro cess and stronglybent cusp, prob a ble P2 with the dor sal pro cess an gu larly bent close to the base and a ten dency to de velop anad di tional pro cess near the bending.


Fig. 21. Early Famennian suc ces sors of the prioniodinid lin eage of Pluckidina lipperti (Bis choff, 1956) from the be gin ning ofthe K. triangularis Zone at Płucki (sam ples Pł−36, A, B, and F) some what youn ger strata at the same lo cal ity (sam ple Pł−1, D,E) and tran si tion from the K. triangularis to K. crepida Zone at Jabłonna (G, I, and M, sam ple J−53; J, sam ple J−60; N, bed 6;H, K, L, bed 7) in the Holy Cross Moun tains. P1 (A, G), P2 (B, H and J), S0 (F), S1 (E and L), S2 (D and I), S3–4 (K), and M (M,N) el e ments; spec i mens ZPAL cXVI/1706, 1707, 1711, 1709, 1708, 1710, 1714, 1715, 1717–1719, 1716, 1720 and 1721,

respectively.

Lagovidina obliqua sp. n.(Figs 23 and 122)

Holotype: Spec i men ZPAL cXVI/1674 (Fig. 23R).Type ho ri zon and lo cal ity: Sam ple Ł−28, early Famennian C. quadrantinodosa Zone at Łagów, Holy Cross Moun tains.Der i va tion of name: Re fer ring to the skewed base of M el e ments.

Ma te rial. — 625 spec i mens.Di ag no sis. — M el e ment with tri an gu lar out line and straight base, P se ries with short pro cesses and a

high cusp, P1 el e ment with the in ner pro cess bi fur cat ing, P2 el e ment strongly bent and twisted, some times de −vel op ing ad di tional anterior process.

Re marks. — The lin eage emerged at Kadzielnia to gether with Klapperilepis crepida and Tripodelluslobus and con tin ued af ter the dis ap pear ance of these spe cies. This old est pop u la tion shows a lat er ally bentdor sal pro cess of the P2 el e ment with out ad di tional branch and a slight angulation at the base of M el e ments.In a prob a bly slightly youn ger sam ple from Miedzianka (Md−1), still con tain ing K. crepida, the dor sal pro −cess of the P2 el e ment is al ready bi fur cated but the cusp is rel a tively small in most spec i mens. Prob a bly this is

46 JERZY DZIK

Fig. 22. Late Famennian prioniodinid Pluckidina purnelli sp. n. from the P. trachytera Zone at Jabłonna (A–G, bed 24) and the L. styriacus Zone at Ostrówka (H–N, sam ple Ost−12) in the Holy Cross Moun tains. P1 (A and H), P2 (B and I), S0 (C and J),S1 (D and K), S2 (E and L), S3–4 (F and M), and M (G and N) el e ments; spec i mens ZPAL cXVI/1675–1888 (holotype, G),

respectively.

a case of a grad ual evo lu tion but the ma te rial is not abun dant enough to dem on strate this re li ably and to sub −stan ti ate erec tion of more chronospecies. More over, the co−oc cur rence of L. multidens sp. n., with some of its el e ments some what sim i lar to those of early mem bers of Lagovidina, makes their sep a ra tion not com pletelysure. Phylo gen etic re la tion ship of L. multidens and Ligonodona albidens to the new spe cies is un clear.A puz zling as pect of the ap pa ra tus struc ture of the lat ter spe cies is that it also con tains prob a ble P2 el e ment


Fig. 23. De rived prioniodinid Lagovidina obliqua sp. n. from the late K. triangularis Zone at Jabłonna (F, sam ple J−53a), theK. crepida Zone at Kadzielnia (A and G, sam ple Ka−7; B, C, and H, sam ple Ka−6; D, E, I, and J, sam ple Ka−3) and Miedzianka(sam ple Md−1, K–N), and the C. quadrantinodosa Zone at Łagów (O–S, sam ple Ł−28). P1 (A, D, F, K, and P), P2 (B, E, L, and O),S1–2 (N), S3–4 (C, M, and Q; note that the lat ter is a clus ter of two el e ments), and M (G–J, R) el e ments; spec i mens ZPALcXVI/1907, 1663, 1664, 1660, 1661, 1723, 1906, 1665, 1662, 1905, 1666, 1667, 1669, 1668, 1671, 1670, 1673, 1674 (holotype,

R), and 1672, re spec tively.

with a high cusp and in ner pro cess which tends to bi fur cate. Whether this is a re sult of par al lel evo lu tion orshared in her ited char ac ter, re mains to be clar i fied.

Oc cur rence. — Early form in the K. crepida Zone at Kadzielnia, typ i cal form in the C. quadrantinodosaand the early C. marginifera zones at Łagów and Miedzianka.

Genus Idioprioniodus Gunnell, 1933

Type spe cies: Idioprioniodus typus Gunnell, 1933 from the Stephanian (Mis sou rian) Kan sas City Group.

Di ag no sis. — M el e ment with strongly ab oral ly bent an te rior pro cess.

Idioprioniodus ruptus sp. n.(Figs 24 and 123)

Holotype: Spec i men ZPAL cXVI/1695 (Fig. 24G).Type ho ri zon and lo cal ity: Bed 24, mid Famennian P. trachytera Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: Re fer ring to the “rup tured” ap pear ance of the ex ter nal pro cess in a ramiform el e ment.

Ma te rial. — 905 spec i mens, some of ques tion able af fin i ties.

48 JERZY DZIK

Fig. 24. Pos si ble an ces tor of Car bon if er ous prioniodinids, Idioprioniodus ruptus sp. n. from the P. trachytera Zone at Jabłonna(A–G, bed 24) and the L. styriacus Zone at Miedzianka (M ,N, sam ple Md−14), and Ostrówka (H–L, O, P, sam ple Ost−12). P1 (Aand H), P2 (B and I), S0 (C and J), S1 (D and K), S2 (E and L), S3–4 (F, N, and O), and M (G, M, and P) el e ments; spec i mens ZPAL

cXVI/1695 (holotype, G), 1698–1702, 1697, 1696, 1703, and 1704, re spec tively.

Di ag no sis. — At least some large S3–4 el e ments show an gu lar twist ing of the ex ter nal pro cess, oth er wisegen er al ized mor phol ogy of the apparatus.

Re marks. — Most of the col lected Idioprioniodus spec i mens are ju ve nile. Only sam ple Md−14 fromMiedzianka yielded frag men tary but ma ture el e ments that show the unique mor phol ogy of S3–4 el e ments.The or i gin of the spe cies is cryp tic; some what sim i lar forms are known among Frasnian Pluckidina spe cies(see Dzik 2002).

Oc cur rence. — The C. marginifera to D. trigonica zones at Kowala, Miedzianka and Ostrówka.

Idioprioniodus uncadinoides sp. n.(Figs 25A–H and 123)

Holotype: Spec i men ZPAL cXVI/1890 (Fig. 25F).

Type ho ri zon and lo cal ity: Bed 27, late Famennian early P. jugosus Zone at Jabłonna, Holy Cross Moun tains.

Der i va tion of name: Re fer ring to the sim i lar ity of some el e ments to Uncadina gen. n.

Ma te rial. — 23 spec i mens.Di ag no sis. — Ro bust S3–4 el e ments with short, prom i nently denticulated ex ter nal pro cess and wide in ner

pro cess bear ing disproportionally small, in clined denticles.Re marks. — The ap pa ra tus re con struc tion of this spe cies is poorly sup ported de spite the char ac ter is tic

ap pear ance of its di ag nos tic S3–4 el e ments. Such el e ments in sam ple Wtr−25, from the K. triangularis Zone at Wietrznia, are as so ci ated with the set of other el e ments of Idioprioniodus. It has to be noted, how ever, thatthis sam ple sig nif i cantly dif fers in age from those with the most typ i cal spec i mens and the ro bust ap pear anceof the in ner pro cess may be in this case a “gerontic” fea ture. A some what sim i lar ap pear ance is ex hib ited bythe me dial pro cess of the S0 el e ment of Trichognathus tumida of Branson and Mehl (1934, pl. 16: 28). Somesup port to so wide a gap in strati graphic oc cur rence is of fered by a find ing of a ju ve nile spec i men with a sim i −lar as pect from co eval strata at Jabłonna (sam ple J−45a, Fig. 25G), in co−oc cur rence with Uncadina unca.This spec i men re sem bles also Guizhoudella.


Fig. 25. Ab er rant prioniodinid Uncadina and its poorly known pos si ble less de rived rel a tive. A–H. Idioprioniodus uncadinoidessp. n. from the K. triangularis Zone at Wietrznia (un cer tain tax o nomic iden tity; A, D, E, and H, sam ple Wtr−25) and Jabłonna (G,sam ple J−45a), and the P. jugosus Zone of the same lo cal ity (B, C, F, bed 27). P1 (A?), P2 (B), S0 (C), S3–4 (D, F, and G), and M(E and H) el e ments; spec i mens ZPAL cXVI/1893, 1888, 1889, 1894, 1895, 1890 (holotype, F), 1891, and 1896, re spec tively.I–L. Uncadina unca (Bis choff, 1956) from the K. triangularis Zone at Jabłonna (sam ple J−45a). S el e ments, spec i mens ZPAL

cXVI/1897, 1998, 1900, 1899, re spec tively.

Oc cur rence. — The K. triangularis to K. crepida zones tran si tion at Wietrznia and the P. jugosus Zone at Jabłonna and Kowala.

Genus Uncadina gen. n.

Type spe cies: Hindeodella unca Bis choff, 1956 from the Early Famennian at the Bicken quarry in the Rhenish Slate Moun −tains.

Di ag no sis. — Only one type of asym met ric el e ment with short, prom i nently denticulated ex ter nal pro −cess, strong cusp and ro bust in ner pro cess with disproportionally del i cate denticulation.

Re marks. — De spite a great num ber of col lected el e ments only one type of them has been iden ti fied. Itseems un likely that any other el e ment kind was as so ci ated with them. All are more or less asym met ric and re −sem ble prioniodinid S3–4 el e ments, es pe cially those of Idioprioniodus uncadinoides sp. n. This is the only ba −sis for clas si fi ca tion of the ge nus among prioniodinids.

Uncadina unca (Bischoff, 1956)(Figs 25I–L and 123)

Type ho ri zon and lo cal ity: Klapperilepis crepida Zone at the Bicken quarry in the Rhenish Slate Moun tains.

Ma te rial. — 724 spec i mens.Di ag no sis. — As for the ge nus.Oc cur rence. — The K. crepida Zone at Jabłonna and Wietrznia.

Genus Guizhoudella Wang et Wang, 1978

Type spe cies: Guizhoudella triangularis Wang et Wang, 1978 from the Late Famennian of Daihua For ma tion at Daihua,Guizhou, China.

Di ag no sis. — Ramiform el e ments of the ap pa ra tus with sur face gran u lar in places; denticles fused intowide, denticulated blades.

Re marks. — Most el e ments of the ap pa ra tus of Guizhoudella dinodontoides sp. n. are homeomorphicwith those of Dinodus leptus Coo per, 1939. In the Tournaisian sam ples of the lat ter spe cies del i cateramiform S and M el e ments co−oc cur with closely sim i lar P2 el e ments (Elictognathus) that have been sug −gested to be long to the Siphonodella ap pa ra tus (Sweet 1988). The only spe cies of “Siphonodella” whichcould be matched with these sets of non−P1 el e ments in the ma te rial stud ied by my self (Dzik 1997) is “S.”lobata. The Famennian spe cies is so sim i lar to that from the Tournaisian that its in de pend ent or i gin is dif fi −cult to ac cept. The prob lem thus emerges how the “Elictognathus” and “Siphonodella” el e ments orig i −nated, if they truly be longed to the same ap pa ra tus. It has to be noted that in Dinodus no tuberculation ofGuizhoudella (or Playfordia, see Dzik 2002) type oc curs. In stead, the el e ments show dis tinct im prints ofameloblasts.

“Elsonella” rhenana Lindström et Ziegler, 1965 from the lat est Givetian hermanni−cristatus Zone (Lind −ström and Ziegler 1965) dif fers from G. dinodontoides sp. n. only in a low−an gle ar range ment of pro cesses inthe prob a ble S1 el e ment. This sug gests that the strati graphic dis tri bu tion of the Famennian Guizhoudella spe −cies re flects their evo lu tion, in which the ap pa ra tus be came grad u ally dis or ga nized, with in creased vari abil ity and ir reg u lar ap pear ance of el e ments. It would then de part from the Dinodus−like mor phol ogy. The long−last −ing Guizhoudella spe cies lin eage ap pears to be cryptic for most of its range.

Guizhoudella triangularis Wang et Wang, 1978(Figs 26 and 124)

Type ho ri zon and lo cal ity: Sam ple ACE 366 with Tripodellus gonioclymeniae, Daihua For ma tion at Daihua, Guizhou,China (Wang and Wang, 1978).

Ma te rial. — 28 spec i mens.Di ag no sis. — El e ments with thin walls of the very deep basal cone, strongly curved cusps; sharp, nu mer −

ous denticles.Re marks. — The holotype of the type spe cies rep re sents an S1 el e ment. S3–4 el e ments from the same

sam ple were named Belodella bilinearis by Wang and Wang (1978). This is so bi zarre a cono dont that iden ti −fi ca tion of its el e ments does not cause any dif fi cul ties, even if they are very rare in sam ples. Some spec i mens

50 JERZY DZIK

re sem ble P el e ments of typ i cal prioniodinids (Fig. 26A) but vari abil ity is so wide that rec og ni tion of dis creteel e ment types may not be possible.

Oc cur rence. — The L. styriacus to D. trigonica zones at Jabłonna, Kowala, Ostrówka, and Dzikowiec.

Guizhoudella dinodontoides sp. n.(Figs 27 and 124)

Holotype: Spec i men ZPAL cXVI/1933 (Fig. 27E).Type ho ri zon and lo cal ity: Sam ple J−45a, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: Re fer ring to the sim i lar ity of el e ments to Dinodus.

Ma te rial. — 12 spec i mens.Di ag no sis. — Rel a tively small basal cone and wide, flat denticles; pro cesses of ramiform el e ments with a

low plat form form ing a rib bon−like struc ture along their basal parts.Re marks. — Sim i lar to the type spe cies, the new spe cies is also long rang ing and ex tremely vari able, but

spe cies dis tinc tion is rather cer tain. Only ju ve niles lack ing char ac ter is tic denticulation and rib bon−like plat −form (e.g., Fig. 27Q, R) may be mis iden ti fied. Much less cer tain is iden ti fi ca tion and homology of par tic u larel e ment types. Two el e ments from sam ple Md−14 may rep re sent the plat form se ries (Fig. 27A, B) if theytruly be long to the spe cies (they lack the tuberculation oc cur ring in un doubted spec i mens). Al ter na tively, theplat form se ries is rep re sented by el e ments with rel a tively straight pro cesses but oth er wise sim i lar to other el −


Fig. 26. Bi zarre prob a ble prioniodinid Guizhoudella triangularis Wang et Wang, 1978 from the P. jugosus Zone at Ostrówka(sam ple Ost 185, A, B, D–F and Ost−265, G), the K. crepida Zone at Jabłonna (C, sam ple J−46; L, bed 27; J, sam ple J−66), the lateK. triangularis Zone at Wietrznia (I, sam ple Wtr−27) in the Holy Cross Moun tains, and the P. jugosus Zone at Dzikowiec (H,sam ple Dz−8) in the Sudetes. P1 (A?), P2 (B?), S0 (C), S1–2 (D and F), S3–4 (E, G, and H), and M (I–L) el e ments; spec i mens ZPAL

cXVI/1806, 1808, 1775, 1807, 1809–1811, 1862, 1909, 1901, 1861, and 1892, re spec tively.

e ment types (Fig. 27G and I). The el e ments S0 and S1 may or may not have the me dial pro cess, but it seems to be al ways rather short in length, un like the prob a bly homeomorphic Car bon if er ous Dinodus.

Oc cur rence. — Rare from the be gin ning of the Famennian at Kowala through the K. crepida Zone atJabłonna, Kadzielnia and Kowala to the L. styriacus Zone at Miedzianka.

Family Gondolellidae Lindström, 1970Di ag no sis. — The P1 el e ments with more or less re duced dor sal pro cess, oth er wise the ap pa ra tus with a

gen er al ized polygnathid−like struc ture; the prioniodinid af fin i ties dis closed only by the “enantiognathus” ap −pear ance of S1 el e ments.

Re marks. — The old est un doubted gondolellid Gondolella are known from the late Car bon if er ous (vonBit ter and Merrill 1998). Its ap pa ra tus dif fers from that of Branmehla only in the mor phol ogy of el e mentlack ing dor sal pro cess but bear ing a plat form along the ven tral pro cess. Ju ve nile el e ments of Branmehlaclosely re sem ble the paedomorphic Perm ian and Tri as sic gondolellid Neospathodus.

Genus Branmehla Hass, 1959Type spe cies: Spathodus inornatus Branson et Mehl, 1934 from the Fammennian Saverton Shale, Mis souri.

Di ag no sis. — Non−plat form P1 el e ments with a short dor sal pro cess, dom i nant cusp to gether with neigh −bor ing denticles gives the el e ment an an gu lar out line; trans versely ex pand ing short basal cavity.

Branmehla bohlenana (Helms, 1959)(Figs 28, 29, and 124)

Type ho ri zon and lo cal ity: Bed 8 in the lime stone quarry at Bohlen near Saalfeld, lower Clymenia beds.

Ma te rial. — 1,664 spec i mens.

52 JERZY DZIK

Fig. 27. Bi zarre prob a ble prioniodinid Guizhoudella dinodontoides sp. n. from the L. styriacus Zone at Miedzianka (sam ple Md−14,A, B, H, J, T), Jabłonna (C–F, K, M, O–S, sam ple J−45a; L, bed 12), and Kowala (G, I, sam ple Ko−18; N, sam ple Ko−154) in theHoly Cross Moun tains. P1 (A?), P2 (B?), S0 (C and L), S1 (D, E, F, and K), S2 (H? and I?), S3–4 (G, and I)), and M (M, N, O–T) el e −ments; spec i mens ZPAL cXVI/1801, 1802, 1813–1815, 1333, 1799, 1809–1811, 1862, 1909, 1901, 1861, and 1892, respectively.

Di ag no sis. — Elon gated and rather ro bust P1 el e ment with rel a tively low cusp.Re marks. — The geo log i cally old est pop u la tions of Branmehla from Łagów (sam ple ŁSł73−2; Fig. 28)

closely re sem ble the ear li est Famennian Pluckidina lipperti (Fig. 21A–E) and that spe cies is likely to be itsan ces tor. The main dif fer ence is in the mor phol ogy of the M el e ment, which is higly arched in B.bohlenana (note that the geo log i cally later Pluckidina lipperti from the K. crepida Zone ex hib its the sameten dency; Fig. 21N).


Fig. 28. Early gondolellid Branmehla bohlenana (Helms, 1959) from the C. quadrantinodosa Zone at Łagów (A–F, sam pleŁSł73−2) and the K. crepida zones at Wietrznia (G, sam ple Wtr−21) and the L. styriacus Zone at Miedzianka (H–R, sam pleMd−14) and Ostrówka (S, T, sam ple Ost−5; W–AA, sam ple Ost−15,) in the Holy Cross Moun tains. P1 (A, B, I–K, T–V, and AA),P2 (E, H, L, and U), S0 (D, M, and W), S1 (N), S2 (E, O, and X), S3–4 (P and Z), and M (F–G, Q, R, and Y) el e ments; spec i mensZPAL cXVI/1970, 1969, 1971–1974, 1968, 1978, 1975, 1977, 1976, 1979–1985, 1988, 1986, 1990, 1987, 1991, 1992, 1994,

1993, and 1989, re spec tively.

The tran si tion from B. bohlenana to B. inornata is grad ual and it is vir tu ally im pos si ble to de limit thesespe cies even if for most of their range they oc cur allopatrically. In the rather representive sam ple Ost−12 from the L. styriacus Zone some el e ments can be found that are in dis tin guish able from those of the type spe cies ofthe ge nus. They grade into the modal mor phol ogy of B. bohlenana and are thus con sid ered ex trememorphologies within the same spe cies. In fact, the mor phol ogy of B. inornata is typ i cal mostly for ju ve nilespec i mens. In the course of on tog eny denticles are added with out in creas ing the el e ment height and as a re −sult the dominantion of the cusp seems to dis ap pear. Ontogenetic ex ten sion of the dor sal pro cess is con nected with its lat eral bend ing, which gives it the characteristic appearance.

The spe cies is rather vari able, es pe cially in re spect to the denticulation of P1 el e ments. Denticles are wideand short, densely dis trib uted in some spec i mens (Fig. 28S), or elon gate and sparsely dis trib uted in oth ers(Fig. 29D).

Panderodella subrecta of Holmes (1928, p. 31, pl. 10:15) from the Chat ta nooga Sshale may be con −specific and thus a se nior syn onym of the spe cies.

Oc cur rence. — Pos si bly an ces tral pop u la tion in the K. crepida Zone at Wietrznia, the typ i cal form fromthe C. quadrantinodosa Zone to the end of the Famennian at Łagów, Miedzianka, Kowala, Ostrówka, andDzikowiec.

Branmehla inornata (Branson et Mehl, 1934)(Figs 30 and 124)

Type ho ri zon and lo cal ity: Prob a bly the Saverton Shale near Mon roe City, Mis souri (Klapper in Ziegler 1975).

Ma te rial. — 6,049 spec i mens.Di ag no sis. — P1 el e ments with high, sharp denticles, which de crease lin early in size with dis tance from

the cusp, to give the el e ment a po lyg o nal profile.Re marks. — Ma ture P1 el e ments of this spe cies re sem ble ju ve niles of B. bohlenana and its or i gin is

clearly paedomorphic. They rarely oc cur sympatrically but then ju ve niles are dif fi cult to tell apart.Oc cur rence. — From the P. trachytera Zone to the end of Famennian at Kowala, Ostrówka, and

Dzikowiec.

54 JERZY DZIK

Fig. 29. Ju ve nile spec i mens of early gondolellid Branmehla bohlenana (Helms, 1959) from the P. trachytera Zone at Kowala(sam ple Ko−8a) in the Holy Cross Moun tains. P1 (A–F), P2 (G, H), S0 (I), S1 (J), S2 (K), S3–4 (L), and M (M and N) el e ments; spec −

i mens ZPAL cXVI/2339, 2338, 530, 2337, 2336, 529, 526, 2340–2344, 2335, and 521, re spec tively.

Branmehla suprema (Ziegler, 1962)(Figs 31B–I and 124)

Type ho ri zon and lo cal ity: Sam ple 2c from the Hönnetal road sec tion in the Rhenish Slate Moun tains, top most costatusZone.

Ma te rial. — 1,539 spec i mens.Di ag no sis. — Dor sal pro cess of P1 el e ment lat er ally bent, widely gap ing basal cone.Re marks. — P2 and S el e ments are rel a tively ro bust, as com pared with other spe cies of the ge nus.

Spathodus fissilis Branson et Mehl, 1934 from the Saverton Shale near Mon roe City, Mis souri may beconspecific.

P1 el e ment of this spe cies some what re sem bles that of B. inornata in pro file view and many mor pho log i −cally tran si tional spec i mens occur.

Oc cur rence. — Prob a ble an ces tral pop u la tion near the end of the C. marginifera at Kowala; un doubtedfrom the L. styriacus Yone to the end of Famennian at Jabłonna, Kowala and Dzikowiec.

Branmehla disparilis (Branson et Mehl, 1934)(Figs 31A and 124)

Type ho ri zon and lo cal ity: Prob a bly the Saverton Shale near Mon roe City, Mis souri (Ziegler 1975).


Fig. 30. Paedomorphic early gondolellid Branmehla inornata (Branson et Mehl, 1934) from the P. jugosus Zone at Jabłonna (A,F, G, and S, bed 27) and Ostrówka (B–E, I–R, and T, U, sam ple Ost−185; H, sam ple Ost−5) in the Holy Cross Moun tains.P1 (A–H), P2 (I), S0 (J, K), S1 (L), S2 (M–O), S3–4 (P, Q), and M (R–U) el e ments; spec i mens ZPAL cXVI/1995, 2000, 2001, 1999,

2002, 1996, 1997, 2003, 2004–2008, 2011, 2009, 2010, 2014, 1998, 2012, and 2013, re spec tively.

Ma te rial. — Four spec i mens.Di ag no sis. — Base of P1 el e ments lat er ally ex panded to form denticulated trans verse pro cesses.Re marks. — Even more prom i nent lat eral pro cess may de velop in P1 el e ments, as shown by Perri and

Spaletta (1991). Prob a bly the spe cies orig i nated from B. suprema, the lat eral pro cess be ing de riv a tive of theasym met ric ex pan sion of the base.

Oc cur rence. — The lat est Famennian at Dzikowiec and Ostrówka.

Family Spathognathodontidae Hass, 1959Di ag no sis. — Biramous S0 el e ment, other el e ments of S se ries with rel a tively short pro cesses.Re marks. — De spite the great time gap be tween the Car bon if er ous oc cur rences of cono donts with

biramous sym met ri cal el e ments in the ap pa ra tus (Hindeodus and Synclydognathus) and sim i lar Si lu rianOzarkodina, it was pro posed by my self that they rep re sent the same clade (Dzik 1991). The oc cur rence ofSynclydognathus in the early Famennian sig nif i cantly re duces this gap and con firms the hy poth e sis of theirclose re la tion ship. One of the new Famennian spe cies ap pears closely sim i lar in its ap pa ra tus or ga ni za tion toco eval Apatognathus, earlier classified among prioniodinids.

Genus Synclydognathus Rexroad et Varker, 1992

Type spe cies: Prioniodus geminus Hinde, 1900 from the Viséan of Scot land.

Di ag no sis. — M and S1–2 el e ments mor pho log i cally sim i lar, with high−arched ar range ment of pro cessesthat are subequal in length.

Re marks. — Ap pa ra tus re con struc tions of the Car bon if er ous spe cies were pre sented by Sweet (1988;as Hindeodus scitulus), Rexroad and Varker (1992), and Dzik (1997, un der mis spelled name Synclado −gnathus). In the high−arched ap pear ance of S el e ments the ge nus is closely sim i lar to Apatognathus, towhich it seems closely re lated. The main dif fer ence is in the mor pho log i cally non−sim pli fied el e ments ofthe plat form se ries.

56 JERZY DZIK

Fig. 31. Ad vanced late Famennian spe cies of Branmehla from the D. trigonica Zone at Ostrówka (sam ple Ost−3) in the HolyCross Moun tains. A. Branmehla disparilis (Branson et Mehl, 1934). P1 el e ment; spec i men ZPAL cXVI/2016. B–I. Branmehlasuprema (Ziegler, 1962). P1 (B, C), P2 (D), S0 (E), S1 (F), S2 (G), S3–4 (H), and M (I) el e ments; spec i mens ZPAL cXVI/2017–

2024, re spec tively.

Synclydognathus ancestralis sp. n.(Figs 32A–F and 121)

Holotype: Spec i men ZPAL cXVI/1824 (Fig. 32F).

Type ho ri zon and lo cal ity: Sam ple Wtr−32, early Famennian K. crepida Zone at Wietrznia, Holy Cross Moun tains.

Der i va tion of name: Re fer ring to an ces tral po si tion to other spe cies of the ge nus.

Ma te rial. — 109 spec i mens.Di ag no sis. — Flat el e ments with elon gated sharp denticles.Re marks. — The new spe cies dif fers from the type spe cies (Rexroad and Varker 1992) in the del i cate,

rel a tively un de rived ap pear ance of el e ments. Falcodus, as in ter preted by my self (Dzik 1997) may be an otherde scen dant lin eage re lated to this species.

Oc cur rence. — The K. crepida Zone at Wietrznia and Jabłonna.

Synclydognathus triramosus sp. n.(Figs 32G–K and 121)

Holotype: Spec i men ZPAL cXVI/1841 (Fig. 32K).

Type ho ri zon and lo cal ity: Bed 24, mid Famennian P. trachytera Zone at Jabłonna, Holy Cross Moun tains.

Der i va tion of name: Re fer ring to triramous ap pear ance of non−P el e ments.

Ma te rial. — 14 spec i mens.Di ag no sis. — All el e ments of the ap pa ra tus with a nar row plat form ex tend ing along pro cesses and ro bust

reg u lar denticulation; S0 el e ment with a short denticulated medial process.Re marks. — From the mor pho log i cally clos est Car bon if er ous type spe cies (Rexroad and Varker 1992),

dif fers in arched pro file of P1 and incipiently triramous S0 and M elements.Oc cur rence. — Only the type ho ri zon and lo cal ity.


Fig. 32. Famennian ozarkodinids Synclydognathus from the Holy Cross Moun tains. A–F. Synclydognathus ancestralis sp. n.from the K. crepida Zone at Wietrznia (sam ple Wtr−32). P1 (A), P2 (B), S0 (C), S1–2 (D), S3–4 (E), and M (F) el e ments; spec i mensZPAL cXVI/1819–1824 (holotype, F), re spec tively. G–K. Synclydognathus triramosus sp. n. from the P. trachytera Zone atJabłonna (bed 24). P1 (G), P2 (H), S0 (I, J), and M (K) el e ments; spec i mens ZPAL cXVI/1838, 1842, 1839, 1840 (holotype, K),re spec tively. I–M. Synclydognathus sp. from the L. styriacus Zone at Ostrówka (sam ple Ost−12). P2 (J), and M (I, K–M)

elements; spec i mens ZPAL cXVI/2384–2386.

Synclydognathus sp.(Fig. 32I–M)

Ma te rial. — Five spec i mens.Re marks. — Rare spec i mens from sam ple Ost−12 may rep re sent a new spe cies of Synclydognathus rep −

re sent ing an other link be tween the early Famennian and Carboniferosus mem bers of the lineage.Oc cur rence. — The L. styriacus Zone at Ostrówka.

Genus Apatognathus Branson et Mehl, 1934Type spe cies: Apatognathus varians Branson et Mehl, 1934 from the Famennian of cen tral Mis souri.

Di ag no sis. — Nar row ro bust pro cesses and sparsely dis trib uted denticles of all el e ments; closely sim i larmor phol ogy of el e ments within M, S, and P series.

Re marks. — Nicoll (1980) pre sented a suc ces sion of Apatognathus spe cies in the Can ing Ba sin that isprob a bly an evo lu tion ary se ries. The change is ex pressed mostly in the mor phol ogy of the M el e ment. In A.provarians from the K. crepida Zone al ter nat ing denticles of the same as pect as those in S el e ments oc curred. In A. klapperi from the C. marginifera Zone they formed a prom i nent fan with out al ter na tion of size. Onedenticle dom i nated in A. varians from the P. trachytera to D. trigonica zones. Nicoll (1980) pointed out alsoa grad ual change in the pat tern of denticle al ter na tion. This suc ces sion is con firmed by the ma te rial from theHoly Cross Moun tains al though the avail able ma te rial of pos si ble A. klapperi is too lim ited to de ter mine thisspe cies with con fi dence.

Apatognathus provarians Nicoll, 1980(Figs 33A–F, I–M, and 121)

Type ho ri zon and lo cal ity: Lower Napier For ma tion in the Can ning Ba sin of Aus tra lia, late K. crepida Zone.

Ma te rial. — 203 spec i mens.Di ag no sis. — Pro cesses of S and M el e ments arched, larger denticles of M el e ment of rel a tively uni form

size.

58 JERZY DZIK

Fig. 33. Famennian ozarkodinids Apatognathus from the Holy Cross Moun tains. A–F, I–M. Apatognathus provarians Nicoll,1980 from the late K. crepida Zone at Łagów (A–F, sam ple ŁSł73−2; M, sam ple Ł−27) and the K. triangularis Zone at Jabłonna(I–L, sam ple J−45a). P (A, B), S0 (I), S1–2 (C, J), S3–4 (D, E and K, L), and M (F) el e ments; spec i mens ZPAL cXVI/1825–1827,1830, 1829, 1828, 1834–1837, and 1831, re spec tively. G, H. Apatognathus varians Branson et Mehl, 1934 from the P. jugosus

Zone at Ostrówka (sam ple Ost−185). S1–2 (G) and M (H) el e ments; spec i mens ZPAL cXVI/1833 and 1832.

Re marks. — Ap pa ra tus re con struc tion was pre sented by Nicoll (1980) and is fully con firmed by the newEu ro pean ma te rial (Fig. 121).

Oc cur rence. — The K. crepida Zone at Jabłonna, Wietrznia, Kadzielnia and Kowala.

Apatognathus varians Branson et Mehl, 1934(Fig. 33G, H)

Type ho ri zon and lo cal ity: Famennian at Dixie, Mis souri (Ziegler 1977, p. 313).

Ma te rial. — 106 spec i mens.Di ag no sis. — Pro cesses of S and M el e ments nar rowly dis posed, M el e ment with one large denticle of al −

most the cusp size.Re marks. — Ap pa ra tus re con struc tion was pre sented by Nicoll (1980) and the few avail able spec i mens

from the Holy Cross Moun tains seem to rep re sent its part (Fig. 121). The di ag nos tic fea tures are shown byone of the type spec i mens of Branson and Mehl (1934, pl. 17:3).

Oc cur rence. — The K. marginifera zone at Jabłonna; rare in all the stud ied lo cal i ties from the P.trachytera to D. trigonica zones.

Family Francodinidae fam. n.Di ag no sis. — A ten dency to bi fur cate pro cesses of S el e ments start ing from S1 and ex pand ing to wards

S4, all el e ments more or less arched in pro file view, M−el e ments tend ing to de velop A−shaped ap pear ance,triramous sym met ri cal el e ment of the apparatus.

Re marks. — This is a well−de fined clade but dif fi cult to di ag nose mor pho log i cally. The branch prob a blystarted with a Mehlina−like (or Pandorinellina−like, de pend ing of which mor phol ogy of M el e ments is prim i −tive) an ces tral form hav ing arched pro file of denticulation in P1 el e ments, strongly ab oral ly bent ex ter nal pro −cess in S el e ments and a rather prom i nent ex ter nal pro cess in M el e ments. From this hy po thet i cal form (sim i −lar to Vogelgnathus proclinatus sp. n.) as a re sult of dim i nu tion of the adult size (paedomorphosis) the lin −eage of Vogelgnathus emerged char ac ter ized by a sur pris ingly prim i tive ap pear ance of the ap pa ra tus for thelate De vo nian. Urbanekodina undata gen. et sp. n. seems to be the end mem ber of this evo lu tion ary trend(Fig. 125), with its el e ments show ing a bi zarre mix ture of palmatolepidid and spathognathodontid as pects.

The lin eage of Francodina at its be gin ning pre served large size of el e ments in her ited from a P.? vogel −gnathoides−like an ces tor (Fig. 126). The prioniodinid−like ap pa ra tuses of Francodina spe cies prob a bly orig i −nated as a re sult of re leased de vel op men tal con trol of the el e ment mor phol ogy. The most sur pris ing as pect ofthis is the trans for ma tion of M el e ments, oth er wise the most mor pho log i cally sta ble el e ment in the post−Or do vi −cian cono dont ap pa ra tuses. In ad vanced F. franconica both its pro cesses achieved a sim i lar length, the ex ter nalpro cess be com ing rib bon−like in Sweetodina lagoviensis sp. n. The most un usual as pect of the ap pa ra tus ofPlanadina plana is the undenticulated in ter nal pro cess of M el e ments – the sit u a tion op po site to that typ i cal forvir tu ally all ozarkodinine cono donts (but sim i lar to the Or do vi cian prioniodontids, e.g., Prioniodus elegans).De spite its seem ingly Ligonodina−like ap pear ance it is ap par ently a francodinid, and its prom i nent ex ter nal pro −cess orig i nated by mod i fi ca tion of a rib bon−like struc ture sim i lar to that of S. lagoviensis.

Genus Vogelgnathus Norby et Rexroad, 1985Type spe cies: Spathognathodus campbelli Rexroad, 1957 from the Namurian (Chesterian) of Il li nois (see Norby and

Rexroad, 1985; van dem Boogard 1992).

Di ag no sis. — Min ute and mor pho log i cally gen er al ized el e ments of a spathognathodontid ap pear ance but with triramous S0 elements.

Vogelgnathus variabilis sp. n.(Figs 34 and 125)

Holotype: Spec i men ZPAL cXVI/1859 (Fig. 34R).Type ho ri zon and lo cal ity: Sam ple J−53a, early Famennian K. triangularis Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: Re fer ring to ex treme vari abil ity in shape of P1 el e ments.

Ma te rial. — 748 spec i mens.Di ag no sis. — M el e ments with rel a tively short and trans versely ar ranged in ner pro cess; cusp of P1 el e −

ment only slightly lon ger than other denticles.


Re marks. — This is the geo log i cally old est spe cies of the ge nus. Its un usual vari abil ity and small sizesug gests a paedomorphic or i gin and re lease of se lec tion pres sure on the ap pa ra tus mor phol ogy. Pos si bly theas so ci ated Mehlina kielcensis sp. n. is close to its an ces tor, as sug gested by sim i lar M and P1 el e ments butother el e ments of the ap pa ra tus are non−paedomorphic. Al ter na tive to this an ces try is the re la tion ship toPandorinellina? vogelgnathoides but this would re quire a cryp tic oc cur rence of the lat ter lin eage dur ing theK. triangularis Zone. As gen er ally the francodinids show punc tu ated dis tri bu tion, this is not un likely.

In late pop u la tions of the spe cies (e.g., J−58, Fig. 34S–BB) most M el e ments show a rounded tip of the ex −ter nal pro cess, which is typ i cal for its suc ces sor, V. proclinatus.

Oc cur rence. — The K. triangularis Zone at Płucki and Jabłonna.

Vogelgnathus proclinatus sp. n.(Figs 35 and 125)

Holotype: Spec i men ZPAL cXVI/1932 (Fig. 35N).Type ho ri zon and lo cal ity: Sam ple Wtr−21, early Famennian K. crepida Zone at Wietrznia, Holy Cross Moun tains.

60 JERZY DZIK

Fig. 34. Ear li est francodinid Vogelgnathus variabilis sp. n. from the early Famennian K. triangularis Zone at Jabłonna (A–R,sam ple J−53a; S–BB, sam ple J−58) in the Holy Cross Moun tains. P1 (A–C and S, T), P2 (D–F and U–W), S0 (G and X), S1 (H, I,and Z), S2 (J–L and Y), S3–4 (M–P and AA), and M (Q, R, and BB) el e ments; spec i mens ZPAL cXVI/1844, 1843, 1845, 1847,1848, 1846, 1849, 1850, 1852, 1851, 1853–1856, 1858, 1857, 1860, 1859 (holotype, R), 1946, 1947, 1950, 1949, 1948, 1951,

1953, 1952, 1954, and 1952, re spec tively.

Der i va tion of name: Re fer ring to the proclined cusp of P1 el e ment.

Ma te rial. — 2,727 spec i mens.Di ag no sis. — Rel a tively long and high−arched in ner pro cess of M el e ment, high and proclined cusp of P1

el e ment.


Fig. 35. Paedomorphic francodinid Vogelgnathus proclinatus sp. n. from the early Famennian K. crepida Zone at Jabłonna(A–G, sam ple J−45a; O–Z, sam ple J−44) and Wietrznia (H–N, sam ple Wtr−21) in the Holy Cross Moun tains. P1 (A, H, Q–S, andX), P2 (B, I, and Y), S0 (C, J, and P), S1 (D, K, and U), S2 (E, L, and T), S3–4 (F, M, O, and W), and M (G, N, V, and Z) el e ments;spec i mens ZPAL cXVI/1919–1932 (holotype, N), 1933, 1942, 1938, 1937, 1939, 1943, 1942, 1934, 1944, 1935, 1940, and 1945,

re spec tively.

Re marks. — Spec i mens of this spe cies from sam ple J−44 at tained a much larger size than typ i cal forVogelgnathus (Fig. 35O, V, X). They are ro bust and bear long and sharp tips of outer pro cesses in S3–4 andM el e ments but pre serve the unique paedomorphic as pects of the ge nus gen er ally con nected with small el −e ment size.

Oc cur rence. — The K. crepida to C. marginifera zones at Jabłonna, Kadzielnia, Kowala, and Wietrznia.

Vogelgnathus arcuatus sp. n.(Figs 36A–J and 125)

Holotype: Spec i men ZPAL cXVI/1959 (Fig. 36D).Type ho ri zon and lo cal ity: Sam ple Wtr−32, early Famennian K. crepida Zone at Wietrznia, Holy Cross Moun tains.Der i va tion of name: Re fer ring to the arched S3–4 el e ment.

Ma te rial. — 47 spec i mens.Di ag no sis. — S0 and S3–4 el e ments bear ing very long rib bon−like pro cesses with arched pro file.Re marks. — P1 el e ments (Fig. 36A) ten ta tively at trib uted to this spe cies re sem ble some large spec i mens

of V. proclinatus in bear ing nu mer ous denticles and an un dif fer en ti ated cusp.Oc cur rence. — The K. crepida Zone at Wietrznia and Miedzianka.

62 JERZY DZIK

Fig. 36. De rived spe cies of the francodinid Vogelgnathus from the early Famennian of the Holy Cross Moun tains. A–J. V. arcu a −tus sp. n. from the K. crepida Zone at Wietrznia (A, H–J, sam ple Wtr−21; B–G, sam ple Wtr−32). P1 (A, B), P2 (C), S0 (D, E),S2 (H), S3–4 (F and I), and M (G and J) el e ments; spec i mens ZPAL cXVI/1966, 1956, 1957, 1959 (holotype, D), 1958, 1960,1964, 1963, and 1965, re spec tively. K–FF. V. unicus (Klapper et al., 2004) in co−oc cur rence with K. rhomboidea from Kowala(K, M, sam ple Ko−164; L, N, UY, Z, and DD, sam ple Ko−163) and Łagów (O–Q, SS–T, VX, AA–CC, EE, FF, sam ple Ł−28).P1 (K–Q), P2 (R–T), S0 (U, V), S1 (W), S2 (X, Y), S3–4 (BB–DD), and M (AA and DD–FF) el e ments; spec i mens ZPALcXVI/2040, 2033, 2041, 2034, 2045, 2044, 2043 (holotype, Q), 2035, 2046, 2047, 2036, 2048, 1373, 2049, 2037, 2039, 2052,

2050, 2051, 2038, 2054, and 2053, re spec tively.

Vogelgnathus unicus (Klapper, Uyeno, Armstrong et Telford, 2004)(Figs 36K–FF and 125)

Type ho ri zon and lo cal ity: Sam ple COB−19 from depth 101.5–101.7 m, up per mem ber of the Long Rap ids For ma tion ofOn tario, rhomboidea Zone.

Ma te rial. — 1,801 spec i mens.Di ag no sis. — Short denticles of P1 el e ments, merg ing to gether into a blade with un du lated mar gin in

adult spec i mens; M el e ments with short pro cesses of subequal length.Re marks. — The type se ries of P1 el e ments of Mehlina? uni ca Klapper et al., 2004 is char ac ter is tic enough

in the pat tern of denticulation to make iden ti fi ca tion of this spe cies rather safe, even if the rest of its ap pa ra tuswas not iden ti fied by Klapper et al. (2004). The sim i lar ity of P1 el e ments to geo log i cally older Mehlinarobustidentata sp. n. may be ac ci den tal but a pos si bil ity can not be ex cluded that V. unicus and V. variabilis in −de pend ently paedomorphically de vel oped from closely re lated spe cies clas si fied here in Mehlina. If truly thiswas the case, as sug gested by the sympatric oc cur rence of these spe cies of Vogelgnathus, the range of the ge nus should be enlarged to preserve its monophyly.

Oc cur rence. — The K. crepida Zone at Jabłonna; the C. quadrantinodosa to P. trachytera zones atŁagów, Miedzianka, and Kowala.

Vogelgnathus branmehloides sp. n.(Figs 37A–R and 125)

Holotype: Spec i men ZPAL cXVI/2066 (Fig. 37K).Type ho ri zon and lo cal ity: Bed 27, late Famennian P. jugosus Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: From sim i lar ity of P1 el e ments to those of Branmehla inornata.

Ma te rial. — 2,560 spec i mens.Di ag no sis. — High arched pro file of P1 el e ments, with dom i nant cusp.


Fig. 37. Late spe cies of the francodinid Vogelgnathus from the mid Famennian of the Holy Cross Moun tains. A–R. V.branmehloides sp. n. from the L. styriacus Zone at Ostrówka (A–J, sam ple Ost−12) and the P. jugosus Zone at Jabłonna (K–R,bed 27). P1 (A, B and K, L), P2 (C and M), S0 (D and N, O), S1 (E), S2 (F, G, and P), S3–4 (H–I and Q), and M (J and R) el e ments;spec i mens ZPAL cXVI/2059, 2056, 2060–2063, 2057, 2058, 2064–2066 (holotype, K), 2067–2073, re spec tively. S–X. V.werneri (Ziegler, 1962) from the L. styriacus Zone at Ostrówka (sam ple Ost−12). P1 (S–V), P2 (W?), and M (X?) el e ments; spec i −

mens ZPAL cXVI/2077, 2075, 2076, 2074, 2078, 2079, re spec tively.

Re marks. — The ap pa ra tus re con struc tion of this spe cies has been pro posed by my self (Dzik 1991, fig.12B) un der the name Pinacognathus(?) sp. Its pro vi sional tax o nomic af fil i a tion re ferred to the sim i lar ity of its P1el e ments to Car bon if er ous el e ments at that time clas si fied in Pinacognathus, which later ap peared to rep re sent P2el e ments prob a bly of Siphonodella (Dzik 1997). The M and S el e ments are closely sim i lar to stratigraphicallyolder V. unicus, be ing dif fer ent only in bear ing less nu mer ous, usu ally not al ter nat ing denticles. But P1 el e mentsof these spe cies are un like each other, rather pre clud ing any di rect an ces tor− de scen dant re la tion ship.

Oc cur rence. — The P. trachytera to P. jugosus zones at Jabłonna, Kowala, Miedzianka, and Ostrówka.

Vogelgnathus werneri (Ziegler, 1962)(Figs 37S–X and 125)

Type ho ri zon and lo cal ity: Sam ple 1132 from the Hönnetal sec tion in the Rhenish Slate Moun tains, up per velifer Zone.

Ma te rial. — 959 spec i mens.Di ag no sis. — Con i cal cusp of P1 el e ment dom i nat ing over nearby denticles but lower than denticles near

the end of ven tral pro cess; dor sal pro cess with ru di men tary denticles.Re marks. — Min ute el e ments in dis tin guish able from Spathognathodus breviatus Wang et Wang, 1978

co−oc cur with other ones typ i cal of this spe cies and prob a bly also their type spec i mens are ju ve niles of thisspe cies. Al ready Van dem Boogard (1992) no ticed a sim i lar ity of Spathognathodus werneri Ziegler, 1962 toVogelgnathus campbelli. Ap pa ra tus com po si tion of the spe cies is poorly known and non−P1 el e ments arehere only provisionally attributed to it.

Oc cur rence. — From the C. marginifera Zone at Miedzianka to the L. styriacus Zone at Kowala,Ostrówka and Jabłonna. Eco log i cally as so ci ated with V. branmehloides. The spe cies oc curs also in theDębnik area (Baliński 1995).

Genus Urbanekodina gen. n.Type spe cies: Urbanekodina undata sp. n. from the mid Famennian of the Holy Cross Moun tains.Der i va tion of name: Af ter Adam Urbanek, to rec og nize his con tri bu tion to un der stand ing the palaeo bi ol ogy and evo lu tion

of co lo nial clonal or gan isms.

Di ag no sis. — Min ute el e ments with trun cated pro cesses.

Urbanekodina undata sp. n.(Figs 38 and 125)

Holotype: Spec i men ZPAL cXVI/2026 (Fig. 38B).Type ho ri zon and lo cal ity: Sam ple Md−8/9, mid Famennian P. trachytera Zone at Miedzianka, Holy Cross Moun tains.Der i va tion of name: Re fer ring to un du lat ing course of denticles in S el e ments.

Ma te rial. — 206 spec i mens.Di ag no sis. — As for the ge nus.Re marks. — S el e ments of the ap pa ra tus of this spe cies are su per fi cially sim i lar to ju ve nile ho mol o gous

el e ments of the ap pa ra tus of the palmatolepidid Conditolepis, with which they co−oc cur. They dif fer in in −

64 JERZY DZIK

Fig. 38. Ex tremely paedomorphic francodinid Urbanekodina undata sp. n. from the P. trachytera Zone at Miedzianka (sam pleMd−8/9) in the Holy Cross Moun tains. P1 (A, B), P2 (C), S0 (D), S1 (E), S2 (F), S3–4 (G), and M (H) el e ments; spec i mens ZPAL

cXVI/2025, 2026 (holotype, B), and 2027–2032, re spec tively.

verse an gu lar ap pear ance, the ex ter nal pro cess be ing “trun cated” in such a way that the first denticles are thelarg est in the row. The al most com plete re duc tion of the me dial pro cess in S0 el e ments is prob a bly apaedomorphic fea ture. The or i gin of this bi zarre cono dont from Vogelgnathus seems likely but not proven.The al ter na tive palmatolepidid re la tion ship would require even deeper transformations.

Oc cur rence. — Pos si bly in the late K. triangularis Zone at Kowala, rare in all lo cal i ties ex cept forMiedzianka, rang ing from the C. quadrantinodosa to L. styriacus Zone.

Genus Francodina gen. n.

Type spe cies: Ligonodina franconica Sannemann, 1955 from the Cheiloceras Stufe of Frankenwald.Der i va tion of name: Af ter Frankenwald, from where most of the el e ments of the ap pa ra tus were first de scribed by

Sannemenn (1955a).

Di ag no sis. — S1 and S2 el e ments in the ap pa ra tus triramous.Re marks. — Mor phol ogy of P se ries and M el e ments of the old est spe cies of this ge nus is sim i lar to that

of Vogelgnathus variabilis sp. n., which ap pears some what ear lier in the Holy Cross Moun tains Famennian.Al though the tran si tion re mains to be dem on strated, it seems likely. Poorly pre served spec i mens re sem bling


Fig. 39. Large−size francodinid Francodina santacrucensis sp. n. from the K. crepida Zone at Jabłonna (bed 8) in the Holy CrossMoun tains. P1 (A), P2 (B), S0 (C), S1 (D), S2 (E), S3–4 (F), and M (G) el e ments; spec i mens ZPAL cXVI/1756–1761 (holotype, E).

F. santacrucensis sp. n. oc cur at Wietrznia in sam ple Wtr−19 rep re sent ing the late K. triangularis Zone.Triramous el e ments are miss ing among them and it is pos si ble, al though far from proven, that they rep re sentthe ancestral population of the lineage.

Francodina santacrucensis sp. n.(Figs 39, 40A–E, and 126)

Holotype: Spec i men ZPAL cXVI/1760 (Fig. 39E).Type ho ri zon and lo cal ity: Bed 8, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: From Latin name of the Holy Cross Moun tains.

Ma te rial. — 1,121 spec i mens.Di ag no sis. — S el e ments with pro cesses emerg ing un der a wide angle.Re marks. — Un like the prob a bly re lated spe cies of Vogelgnathus, el e ments of F. santacrucensis are of

large size, even as com pared with the larg est known cono donts of the Famennian. There seems to be an in −crease in ma ture size in the evo lu tion of the lin eage, prob a bly a re ver sal from the ten dency to paedo morphism,which was prob a bly be hind the or i gin of the branch of francodinids. Pandorinellina? vogel gnathoides sp. n.(Fig. 125) is some what sim i lar to this old est Francodina in show ing a ten dency to bi fur ca tion of the ex ter nalpro cess of S1 el e ments and arched con tour of P1 el e ments. Its S el e ments, how ever, are still of a rather gen er al −ized polygnathid ap pear ance and the spe cies is known from sig nif i cantly youn ger strata.

Oc cur rence. — The K. crepida Zone at Jabłonna, Wietrznia, and Kadzielnia.

66 JERZY DZIK

Fig. 40. Spe cies of Francodina from the K. crepida Zone at Jabłonna in the Holy Cross Moun tains. A–E. Pop u la tion tran si tionalbe tween F. santacrucensis sp. n. and F. franconica (Sannemann, 1955) (sam ple J−46). P1 (A), P2 (B), S1 (C), S3–4 (D), and M (E)el e ments; spec i mens ZPAL cXVI/1770–1774. F–L. F. franconica (Sannemann, 1955) (bed 8). P1 (F), P2 (G), S0 (H), S1 (I), S2

(J), S3–4 (K), and M (L) el e ments; spec i mens ZPAL cXVI/1763–1769.

Francodina franconica (Sannemann, 1955)(Figs 40F–L and 126)

Type ho ri zon and lo cal ity: Black lime stone with Nehdenites verneuili at Breitengrund, Frankenwald (Sannemann 1955a).

Ma te rial. — 1,593 spec i mens.Di ag no sis. — M el e ment with equal length of high−arched pro cesses, ex ter nal pro cess of S el e ments

strongly recurved.Re marks. — Sannemann’s (1955a) Apatognathus inversus, Ligonodina franconica, Roundya franca,

Tripodellus flexuosus and Helms’ (1959) Roundya prava and Tripodellus tenuis all were based on el e ments ofthe spe cies, which are very char ac ter is tic and easy to iden tify even in as so ci a tion with other francodinids andprioniodinids. El e ments of the P se ries are sim i lar to each other in pro file view but dif fer in cur va ture: P1 el e −ments are twisted whereas P2 are lat er ally bent. S1 el e ment mim ics el e ments of the palmatolepidid Tripodellus.The spe cies co−oc curs sympatrically for most of its range with F. santacrucensis, which is ap par ently moreprim i tive and closer mor pho log i cally to the com mon an ces tor. A pos si bly relic pop u la tion tran si tional be tweenthem oc curs at Jabłonna (J−46, Fig. 40A–E) and may rep re sent an other spe cies of the ge nus.

Oc cur rence. — Pos si bly the lat est K. triangularis Zone at Wietrznia; the K. crepida Zone at Jabłonna,Wietrznia and Kadzielnia.

Genus Sweetodina gen. n.Type spe cies: Ligonodina monodentata Bis choff et Ziegler, 1956 from the late Famennian of the lime stone quarry NE

Weitershausen.Der i va tion of name: In rec og ni tion of the great con tri bu tion to the pa le on tol ogy of cono donts by Wal ter C. Sweet (The Ohio

State Uni ver sity, Co lum bus).

Di ag no sis. — S el e ments with a ten dency to pla nar dis po si tion of pro cesses, the ex ter nal one pointed.


Fig. 41. Ad vanced francodinid Sweetodina from the late K. crepida Zone at Łagów in the Holy Cross Moun tains. A–G. S. lago −viensis sp. n. (sam ple Ł−28). P1–2 (A), S0 (B), S1 (C), S2 (D), S3–4 (E), and M (F, G) el e ments; spec i mens ZPAL cXVI/1787–1792(holotype, F). H–K. Early pop u la tion of S. monodentata (Bis choff and Ziegler, 1956) (sam ple ŁSł73−2). P1–2 (H), S2 (I), S3–4 (J),

and M (K) el e ments; spec i mens ZPAL cXVI/1793–1796.

Re marks. — Dis tinc tions be tween el e ment types of Sweetodina are oblit er ated by the unusualy highvari abil ity. There seems to be no morphologic dif fer ence be tween el e ments of the P series.

Sweetodina lagoviensis sp. n.(Figs 41A–G and 126)

Holotype: Spec i men ZPAL cXVI/1791 (Fig. 41F).Type ho ri zon and lo cal ity: Sam ple Ł−28, mid Famennian C. quadrantinodosa Zone at Łagów, Holy Cross Moun tains.Der i va tion of name: From latinized name of the type lo cal ity.

Ma te rial. — 436 spec i mens.Di ag no sis. — M el e ment with elon gated pro cesses, the outer pro cess rib bon−like, in ner strongly denticu lated.Re marks. — This is prob a bly a lo cal, allopatrically de vel oped spe cies as it is pre ceded in the Łagów sec −

tion by the less de rived (in re spect to M, but not S3 el e ment mor phol ogy) S. monodentata and suc ceeded bythe same spe cies in the re gion. The bi zarre ap pear ance of the M el e ment makes it dif fer ent from all other De −vo nian cono donts and only the sim i lar ity to F. franconica al lows rec og ni tion of its af fin ity.

Oc cur rence. — The C. quadrantinodosa to C. marginifera (pos si bly P. trachytera) zones at Łagów andMiedzianka.

Sweetodina monodentata (Bischoff et Ziegler, 1956)(Figs 41H–K, 42, and 126)

Type ho ri zon and lo cal ity: Late Famennian lime stone at quarry NE Weitershausen.

Ma te rial. — 480 spec i mens.

68 JERZY DZIK

Fig. 42. Late pop u la tions of the francodinid Sweetodina monodentata (Bis choff et Ziegler, 1956) from the late L. styriacus Zone atŚciegnia (Wzdół Plebański; sam ple Wzd−13, A–C, F, J, K) in the Holy Cross Moun tains and late P. jugosus and D. trigonica zones atDzikowiec (sam ple Dz−8, D, E, Dz−7, G, I, J, and Dz−10, H) in the Sudetes. P1–2 (A–C, I), S0 (D), S1–2 (E), S3 (H and L), S4 (K), and M(J and M) el e ments; spec i mens ZPAL cXVI/1863–1965, 1873, 1874, 1867, 1870, 1875, 1869, 1868, 1866, 1871, 1872, re spec tively.

Di ag no sis. — M el e ment with subequal length of short pro cesses, outer pro cess undenticulated and sharplzpointed, in ner one with a few denticles; outer pro cess of S3–4 el e ments with en larged prox i mal denticle.

Re marks. — Along with the type spe cies, also the types of Ozarkodina homoarcuata and Neoprionioduspostinversus of Helms (1959) be long to the same ap pa ra tus. The early pop u la tion of the spe cies from Łagów(Fig. 41H–K) dif fers from that of the late Famennian only in more highly arched pro cesses of the M el e ment,in this re spect re sem bling prob a bly an ces tral Francodina and its suc ces sor, S. lagoviensis. En larged denticlein S3 el e ment in di cates some de gree of ad vance ment so S. lagoviensis prob a bly did not orig i nate in placefrom this pop u la tion but only shared with it com mon an ces try. That this el e ment type rep re sents only a sin gle lo ca tion in S. monodentata is sug gested by the oc cur rence of sim i lar el e ments lack ing denticulation on theouter pro cess and with denticles of the in ner pro cess al most uni form in size. They are thus tran si tional mor −pho log i cally be tween S and M el e ment types. In ner pro cess in M el e ments is short and oriented almostperpendicular to the cusp in late populations.

Oc cur rence. — Prob a bly the C. quadrantinodosa Zone at Kowala and Łagów, wide spread from the P.trachytera Zone to the end of Famennian.


Fig. 43. The ad vanced francodinid Planadina plana (Helms, 1959) from the early P. jugosus Zone at Jabłonna (A, D, and F, bed27), the C. quadrantinodosa Zone at Łagów (B, C, sam ple ŁSł73−5), and the L. styriacus Zone at Ostrówka (E, H, and J, sam pleOst−5; G and I, sam ple Ost−12) in the Holy Cross Moun tains. P1 (A?), S0 (J?), S1 (B?), S2 (C, I), S3–4 (D), and M (E–H) el e ments;

spec i mens ZPAL cXVI/1776, 1779, 1780, 1777, 1783, 1778, 1785, 1782, 1784, and 1781, re spec tively.

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