dtlpddth , D ppt th 0 lt , lt ll P lt th lpd ll dttdtl , lthpldp, lld

Genus Conditolepis van den Boogaard et Kuhry, 1979Type spe cies: Palmatolepis marginifera Helms, 1959 from the Cheiloceras Stufe of the Rhenish Slate Moun tains.

Di ag no sis. — Ap pa ra tus with biramous S0 el e ments, lat er ally sin u ous P2 el e ments with low cusp andusu ally uni form dis tri bu tion of denticles, M el e ment hav ing proclined cusp, lin early ar ranged pro cesses withra di at ing denticles of the dorsal process.

Re marks. — Panderolepis Helms, 1963 has a pri or ity over Conditolepis van den Boogard and Kuhry,1979 but it was based on the type spe cies Polygnathus? serratus Hinde, 1879, which was found to be anomen dubium by Hud dle (1968).

Conditolepis lobicornis (Schülke, 1995)(Figs 101A–M and 136)

Type ho ri zon and lo cal ity: Bed 29 of Seßacker trench near Dillenburg in the Rhenisch Slate Moun tains (Schülke 1995).

Ma te rial. — 403 spec i mens.Di ag no sis. — Smooth P1 el e ment with a nar row ex ten sion of the pos te rior lobe and gently bent P2 el e −

ment with in cip i ent plat form and denticles of al most uni form height and size.Re marks. — Ap pa ra tus was re stored by Schülke (1999) who iden ti fied it as the first mem ber of the

branch of palmatolepidids with re duced ad di tional pro cesses in S0 el e ments, in di cat ing also its prox im ity toK. linguiloba sp. n. rep re sent ing a con tin u a tion of its lin eage. As shown by Schülke (1999) Klapperilepissubperlobata (Branson et Mehl, 1934 sensu Schülke 1999) is an ces tral to C. lobicornis. Its re la tion ship to,and pos si ble iden tity with, K. triangularis remains to be clarified.

The shape of the plat form of P1 el e ments is very vari able in each of the stud ied sam ples, with end−mem −bers of tran si tion se ries ap proach ing Palmatolepis initialis sp. n. (Fig. 101 C–E), C. tenuipunctata (Fig.101O), and C. linguiloba sp. n. (Fig. 101E). This ap par ent sim i lar ity is the re sult of a phylo gen etic prox im ityto all these spe cies, in the first case the shared fea tures of the ap pa ra tus mor phol ogy are plesiomorphic, intwo oth ers these could be con sid ered apomorphic for the ge nus. There is a clear morphologic gap be tweenthe ranges of the plat form shape vari abil ity of the type pop u la tions of C. lobicornis, C. linguiloba sp. n., andC. tenuipunctata.

Klapper et al. (2004) con sid ered C. lobicornis conspecific with Palmatolepis subperlobata helmsiOvnatanova, 1976, here re named C. linguiloba sp. n. How ever, the holotype of C. lobicornis with its nar rowpos te rior lobe is unsimilar to spec i mens of C. linguiloba, and no spec i men in the type se ries (Schülke 1995,pl. 40) fits its mor phol ogy. In stead, they tend to be sim i lar in the shape of the plat form to un de rived spe cies of Palmatolepis.

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

Conditolepis linguiloba sp. n.(Figs 101N, O and 136)

Holotype: Spec i men ZPAL cXVI/2794 (Fig. 101N).Type ho ri zon and lo cal ity: Sam ple J−48, early Famennian K. crepida Zone at Jabłonna, Holy Cross Moun tains.Der i va tion of name: Re fer ring to the tongue−like shape of the plat form lobe.

Ma te rial. — 125 spec i mens.Di ag no sis. — Smooth P1 el e ment with a wide tongue−like ex ten sion of the pos te rior lobe.Re marks. —Al though no other el e ment of the ap pa ra tus of this spe cies was iden ti fied with con fi dence,

co−oc cur rence of its rel a tively nu mer ous P1 el e ments with spe cies of Conditolepis sug gests that the P1 andother el e ments of their ap pa ra tuses were in dis tin guish able, thus more ad vanced than those of C. lobicornis.C. lobicornis prob a bly rep re sents the same lin eage but dif fers in the more pointed tip of the plat form lobe inthe P1 el e ments, its P2 el e ments vir tu ally lack a platform, and occur in older strata.

This spe cies has been re ferred to as Palmatolepis subperlobata by i.a., Wolska (1967), Dreesen andDusar (1974), Helms and Ziegler in Clark (1981, fig. 52), Wang (1989), and Metzger (1994), but it def i nitelyrep re sents a dif fer ent lin eage, as shown both by the mor phol ogy of P1 el e ments and ap pa ra tus com po si tion(Schülke 1999). Klapper et al. (2004) used the name C. lobicornis for this spe cies as a re place ment forPalmatolepis subperlobata helmsi Ovnatanova, 1976, a hom onym of Palmatolepis helmsi Ziegler, 1962.

Oc cur rence. — The K. crepida and C. quadrantinodosa zones at Jabłonna, Kowala, Miedzianka, andŁagów.

138 JERZY DZIK

Conditolepis tenuipunctata (Sannemann, 1955)(Figs 101P–CC and 136)

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

Ma te rial. — 3,535 spec i mens.Di ag no sis. — P1 el e ments elon gate and sin u ous, with wide smooth plat form bear ing rounded tri an gu lar

ex ten sion in the middle.Re marks. — The spe cies may be re lated to stratigraphically older C. lobicornis, from which it dif fers in

the re duced lobe of the plat form. At Wietrznia in sam ple Wtr−27 an early pop u la tion of this spe cies oc curs

FAMENNIAN CONO DONTS AND AMMONOIDS 139

Fig. 101. Early spe cies of the palmatolepidid Conditolepis. A–M. Conditolepis lobicornis (Schülke, 1995) from the late K. trian −gularis Zone at Jabłonna (A, B, F, sam ple J−65) and Wietrznia (C–E, G–L, sam ple Wtr−27). P1 (A–E), P2 (F–J), S1 (K), S3–4 (L),and M (M) el e ments; spec i mens ZPAL cXVI/2875, 2878, 2803, 2804, 2879, 1417, 2801, 2805, 2802, 1418–1420, re spec tively.N, O. C. linguiloba sp. n. from the K. crepida Zone at Jabłonna (sam ple J−48). P1 el e ments; spec i mens ZPAL cXVI/2794(holotype) and 2793. P–BB. Conditolepis tenuipunctata (Sannemann, 1955) from the K. crepida Zone at Jabłonna (sam pleJ−45a). P1 (P–R), P2 (S, T), S0 (U, V), S1 (W), S2 (X, Y), S3–4 (Z), and M (AA, BB) el e ments; spec i mens ZPAL cXVI/1303, 1301,

1302, 1304–1306, 1311, 1308, 1307, 1309, 1310, and 2829, re spec tively.

with P2 el e ments vir tu ally lack ing the plat form, sim i lar to those of C. falcata, that is with out a rec og niz ablecusp. Such an el e ment was al ready at trib uted to the spe cies by den Boogaard and Kuhry (1979, fig. 8; thespec i men on Fig. 7 be longs to Klapperilepis quadrantinodosolobata, as pointed out by Schülke 1999).Youn ger sam ples with P1 el e ments typ i cal for the spe cies con tain P2 el e ments with very wide, oval plat form,closely sim i lar to that in more ad vanced Conditolepis spe cies. I sug gest to dis tin guish early and late forms ofthe spe cies on this ba sis. A ro bust ap pear ance of P1 el e ments char ac ter izes pop u la tions with the plat form ofP2 el e ments al ready well de vel oped but not reaching the tip of the inner process (sample Md−27).

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

Conditolepis prima (Ziegler et Huddle, 1969)(Figs 102 and 136)

Type ho ri zon and lo cal ity: Red dish lime stone at the base of the up per Cheiloceras Stufe at Amönau near Mar burg, RhenishSlate Moun tains (Ziegler 1975).

Ma te rial. — 7,165 spec i mens.Di ag no sis. — Nar row P1 el e ments with short an te rior lobe hav ing el e vated rounded mar gin and no pos te −

rior lobe.Re marks. — P2 el e ments of the ap pa ra tus were iden ti fied by van den Boogaard and Kuhry (1979) and

Metzger (1994) and its com plete res to ra tion was of fered by Schülke (1999).The spe cies is tran si tional be tween C. tenuipunctata and C. glabra but usu ally is un der stood in a ty po logi −

cal sense. Schülke (1999) synonymized it with Palmatodella unca of Sannemann (1955a). How ever, C.tenuipunctata and P. unca share the same type lo cal ity and there is noth ing in the text of their orig i nal de −scrip tion (Sannemann 1955a) that would sug gest that they oc cur in dif fer ent ho ri zons. More over, Sanem −mann (1955a) il lus trated typ i cal C. glabra from the same lo cal ity. Per haps Schülke (1999) was in pos ses sion

140 JERZY DZIK

Fig. 102. Conditolepis prima (Ziegler et Hud dle, 1969), an early mem ber of the C. glabra lin eage, from the K. crepida Zone atJabłonna (I–L, sam ple J−50) and Kadzielnia (E, F, and H, sam ples Ka−3; G, I–K, sam ple Ka−4) in the Holy Cross Moun tains.P1 (A, B, E, F), P2 (C, G, H), S0 (I), S2 (J), and M (K, L) el e ments; spec i mens ZPAL cXVI/2831, 2830, 2832, 2833, 2867, 2868,

2871, 2869, 2872–2874, and 2870, re spec tively.

of ad di tional in for ma tion that the third spe cies is rep re sented in the same as sem blage. In my ma te rial ap pa ra −tuses of C. tenuipunctata and C. glabra, as well as pop u la tions pos si bly tran si tional be tween them, do not dif −fer in their ap pa ra tus com po si tion. This re fers also to the M el e ments and the spec i men il lus trated by Schülke(1999, pl. 3: 18) as be long ing to C. tenuipunctata may be atyp i cal. Nar row−plat form el e ments clas si fied bySchülke (1999) in P. unca co−oc cur with typ i cal el e ments of C. tenuipunctata in its late sam ples and the same


Fig. 103. Ad vanced spe cies of the palmatolepidid Conditolepis glabra lin eage from the Holy Cross Moun tains. A–P. Condito −lepis glabra Ulrich et Bassler, 1926 from the C. quadrantinodosa Zone at Łagów (A–H, sam ple Mak−3−4; I, J, sam ple Ł−26; L–N, sam ple Ł−28) and Miedzianka (O, P, sam ple Md−2). P1 (A–C, I–M), P2 (D, N, O), S0 (E), S2 (F), S3–4 (G, P?), and M (H) el e ments;spec i mens ZPAL cXVI/2837, 2836, 2835, 2838–2842, 2864, 2863, 1394, 1393, 1395, 1397, 2865, 1393, and 2866, re spec tively.Q, R. Conditolepis distorta (Branson et Mehl, 1934) from the C. marginifera Zone at Łagów (sam ple Ł−9). P1 el e ments; spec i −

mens ZPAL cXVI/2850 and 2851.

is in the Montagne Noire ma te rial of Schülke (1999, p. 91). This sug gests that these are just morphologieswithin monospecific pop u la tions in stead of dis tinct spe cies and that in the course of evolution narrow− platform elements became more and more numerous.

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

Conditolepis glabra Ulrich et Bassler, 1926(Figs 103A–P and 136)

Type ho ri zon and lo cal ity: “Hardin sand stone” near Mount Pleas ant, Ten nes see (Hud dle 1968).

Ma te rial. — 2,230 spec i mens.Di ag no sis. — P1 el e ments with a ridge along the an te rior mar gin of the plat form, ven trally de part ing from

the blade and trun cated with an angulation per pen dic u lar to it.Re marks. — In sev eral large sam ples stud ied there is a com plete gra da tion be tween morphologies tra di −

tion ally be lieved to char ac ter ize sep a rate spe cies, in clud ing “Palmatolepis” prima (Fig. 103L), “P.” pecti −nata (Fig. 103K–M) or “P.” acuta (Fig. 103J). I con sider them vari ants within pop u la tion vari abil ity, al −though the type pop u la tions of some of them may truly be dis tinct from C. glabra.

El e ments of the ap pa ra tus of this spe cies are closely sim i lar to those of C. quadrantinodosa and it is dif fi −cult to sep a rate them in sam ples. The dif fer ence is that in C. glabra P2 el e ments are more arched and M el e −ments more elon gated than in the other lineage.

The spe cies is ap par ently an ces tral to C. distorta and prob a bly to C. falcata. Evo lu tion from C. prima toC. glabra and from C. glabra to C. distorta is con nected with in creased vari abil ity and tran si tional pop u la −tions are poly mor phic in re spect to mor phol ogy of P1 el e ments. De lim i ta tion of these prob a ble chronospecies is thus rather conventional.

Oc cur rence. — The C. quadrantinodosa Zone at Łagów, Kowala, Jabłonna, and Miedzianka; also in theDębnik area (Baliński 1995).

Conditolepis distorta (Branson et Mehl, 1934)(Figs 103Q, R and 136)

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

Ma te rial. — 890 spec i mens.Di ag no sis. — P1 el e ments elon gate and si nu soi dal, with nar row plat form bear ing a prom i nent an te rior

ridge, close to the blade along its whole length.Re marks. — Tran si tion from C. glabra to C. distorta took place early in the C. marginifera Zone. As al −

ready shown in the Can ning Ba sin suc ces sion in Aus tra lia by Glenister and Klapper (1966, text−fig. 3), thechange is grad ual. Pop u la tions in ter me di ate in age show more and more com mon el e ments hav ing the nar −row fur row be tween the ridge on the an te rior lobe and the blade, in clud ing those usu ally clas si fied as C. pec −tinata.

Oc cur rence. — Typ i cal pop u la tions are com mon in the C. marginifera Zone at Łagów, Kowala, andMiedzianka. Rare spec i mens from higher ho ri zons at Ostrówka and Miedzianka may be reworked.

Conditolepis falcata (Helms, 1959)(Figs 104 and 136)

Type ho ri zon and lo cal ity: Up per part of bed 5 at Bohlen near Saalfeld, Thuringia (Helms, 1959).

Ma te rial. — 41,366 spec i mens.Di ag no sis. — P1 el e ments very elon gate and nar row, with tri an gu lar, ab oral ly point ing an te rior lobe;

P2 el e ments sin u ous, with ru di men tary plat form, S0 el e ments with prominent processes.Re marks. — P2 el e ments of the ap pa ra tus were iden ti fied by van den Boogaard and Kuhry (1979), the

com plete ap pa ra tus by my self (Dzik 1991). Metzger (1994) con firmed iden ti fi ca tion of the sym me try tran si −tion se ries but ques tioned the iden ti fi ca tion of the M el e ment, transfering it in cor rectly to P. schindewolfi(this would be the only el e ment of that spe cies dif fer ent from the ap pa ra tus of re lated P. perlobata). C.falcata is the most abun dant palmatolepidid spe cies in the Holy Cross Moun tains and its ap pa ra tus re con −struc tion leaves lit tle doubt. Sam ple Ost−12 of fers the larg est ma te rial of this spe cies be ing de void of anyother Conditolepis species.

142 JERZY DZIK

The or i gin of this spe cies re mains in doubt. The spe cies emerged from an un known source in the late K.crepida Zone at Kadzielnia to gether with equally cryp tic C. klapperi. Al though rel a tively lit tle change isnec es sary in the mor phol ogy of the P1 el e ment to de rive it from that of C. glabra, the whole ap pa ra tus isquite dif fer ent. It can not be ex cluded that the lack of a plat form in P2 el e ments is an an ces tral fea ture in her −ited from C. lobicornis and early C. tenuipunctata and that the lin eage of C. falcata is in de pend ent of thatof C. glabra.

Oc cur rence. — Com mon from the C. quadrantinodosa Zone, most abun dant in the P. trachytera and L.styriacus zones of all stud ied lo cal i ties. Rare spec i mens found to gether with P. jugosus at Ostrówka andMiedzianka may be reworked.


Fig. 104. Palmatolepidid Conditolepis falcata (Helms, 1959) from the C. quadrantinodosa Zone at Łagów (C, F, I, J, M, and P,sam ple Ł−28) and the early Lagovignathus styriacus Zone at Ostrówka (A, B, D, E, G, H, K, L, N, O, and Q, sam ple Ost−12) in theHoly Cross Moun tains. P1 (A–C), P2 (D–F), S0 (G–I), S1–2 (J, K), S3–4 (L–N), and M (O–Q) el e ments; spec i mens ZPALcXVI/2852, 2853, 1392, 2854, 2855, 1396, 2857, 2856, 1398, 1399, 2858, 2859, 1400, 2860, 2862, 1401, and 2861, re spec tively.

Conditolepis klapperi (Sandberg et Ziegler, 1973)(Figs 105A–G and 136)

Type ho ri zon and lo cal ity: 42 m above the base of the West Range Lime stone at the Bactrian Moun tains, Ne vada (Ziegler1975).

Ma te rial. — 2,846 spec i mens.Di ag no sis. — An te rior mar gin of the P1 el e ment with a mas sive ramp−like ridge, pos te rior mar gin of the

plat form non−lobate, strongly sinuous.Re marks. — This is the first spe cies of the C. marginifera lin eage as in di cated by its ro bust ap pear ance

with mas sive an te rior ridge of the plat form. Sim i larly to the P1 el e ment, other el e ments of the ap pa ra tus are of large size, mas sive ap pear ance and reg u lar dis tri bu tion of sharp denticles, oth er wise they re sem ble those ofother typ i cal Conditolepis species.

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

Conditolepis quadrantinodosa (Branson et Mehl, 1934)(Figs 105H–N and 136)

Type ho ri zon and lo cal ity: Saverton Shale seven miles north of Mon roe City, Mis souri (Ziegler 1977, p. 367).

Ma te rial. — 139 spec i mens.Di ag no sis. — Very vari able P1 el e ment, oval in out line with gently con vex an te rior lobe cov ered with tu −

ber cles or form ing a ridge.Re marks. — The range of pop u la tion vari abil ity of this spe cies en com passes, with com pletely smooth

gra da tion, morphologies typ i cal of C. inflexa (see Dreesen and Dusar 1974; Dreesen 1976) but also C.marginifera and C. inflexoidea. They sep a rated into dis tinct bi o log i cal spe cies some what later in theirevolution.

Oc cur rence. — Zone of its own at Łagów, Kowala, and Miedzianka.

Conditolepis marginifera (Helms, 1959)(Figs 105O, P, S–V and 136)

Type ho ri zon and lo cal ity: Bed 13 of red dish lime stone with Cheiloceras enkebergense in trench II at Seßacker nearOberscheld, Rhenisch Slate Moun tains (Ziegler 1977, p. 325).

Ma te rial. — 1,430 spec i mens.Di ag no sis. — Short P1 el e ment with subcircular dor sal end, bear ing a sharp ridge or row of tu ber cles

along the an te rior mar gin in prox im ity to the blade.Re marks. — Be cause of the tre men dous mor pho log i cal vari abil ity of the P1 el e ments of this spe cies it is

dif fi cult to de ter mine it re li ably based on sin gle spec i mens. In fact, the co−oc cur rence of C. inflexoidea, sep a −rated by a gap in morphologic vari abil ity, gives a help ful hint that one is deal ing with a sep a rate spe cies andnot with an ex treme mor phol ogy of the stratigraphically pre ced ing C. quadrantinodosa.

Oc cur rence. — Zone of its own at Łagów, Kowala, Ostrówka, and Miedzianka.

Conditolepis inflexoidea (Ziegler, 1962)(Figs 105Q, R and 136)

Type ho ri zon and lo cal ity: Bed 13 of red dish lime stone with Cheiloceras enkebergense in trench II at Seßacker nearOberscheld, Rhenish Slate Moun tains (Ziegler 1975).

Ma te rial. — 145 spec i mens.Di ag no sis. — Elon gate P1 el e ment with gently con vex smooth an te rior lobe.Re marks. — The prob lem of the evo lu tion ary or i gin of this spe cies re mains to be solved. As shown by

Dreesen and Dusar (1974) and Dreesen (1976) there is a tre men dous poly mor phism in the pop u la tions tran si −tional be tween those with dom i nant C. quadrantinodosa and C. marginifera morphologies, which oblit er ates the true na ture of evo lu tion ary change. Whether the ge netic bar rier de vel oped sympatrically in the wholearea of oc cur rence of the spe cies or, al ter na tively, an allopatric speciation event not ex pressed in mor phol ogy pre ceded spa tial re uni fi ca tion of the lin eage, re mains un known. Per haps de tailed biometrical stud ies in suc −ces sions from dif fer ent places in the world may help in finding an answer to this question.

Oc cur rence. — The C. marginifera Zone at Łagów, Kowala, and Miedzianka.

144 JERZY DZIK


Fig. 105. Palmatolepidids of the Conditolepis marginifera lin eage from the Holy Cross Moun tains. A–G. C. klapperi (Sandberg etZiegler, 1973) from the K. crepida Zone at Kadzielnia (sam ple Ka−10). P1 (A–C), P2 (D), S0 (E), S1–2 (F), and S3–4 (G) el e ments;spec i mens ZPAL cXVI/2845, 2844, 2846, 2847, 2849, and 2848, re spec tively. H–N. C. quadrantinodosa (Branson et Mehl, 1934)from its nom i nal zone at Łagów (sam ple Ł−28). P1 (H–J), P2 (K), S0 (L), S1–2 (M), and S3–4 (N) el e ments; spec i mens ZPALcXVI/1382–1386, 1389, and 1390, re spec tively. O, P, S–V. C. marginifera (Helms, 1959) from its nom i nal zone at Łagów (sam pleŁSł73−3). P1 (O, P), P2 (S), S0 (T), S1–2 (U), and M (V) el e ments; spec i mens ZPAL cXVI/2821, 2825, 2826, 2823, 2827, and 2828,

re spec tively. R. C. inflexoidea (Ziegler, 1962) from the same sam ple. P1 el e ments; spec i mens ZPAL cXVI/2825 and 2824.

Family Cavusgnathidae Austin et Rhodes, 1981Di ag no sis. — P1 el e ments with ax ial sym me try, bear ing elab o rate icrion or high−po si tioned platform.Re marks. — The three Famennian cono dont lin eages with more or less de vel oped ax ial sym me try in

their plat form el e ments dis cussed be low, prob a bly share their com mon an ces try in a Pandorinellina−likeform of early Famennian age. The Pseudopolygnathus lin eage con tin ued with out any dras tic mor pho log i calchanges from the Famennian to the Tournaisian when it di ver si fied sig nif i cantly (Dzik 1997). Alterno −gnathus is ap par ently only a homeomorph of Pinacognathus (sensu Dzik 1994) and early Siphonodella(Ziegler and Sandberg 1984; Sweet 1988). Omolonognathus, known from the late Famennian of the Cra cowarea (Baliński 1995) may be an ex tremely shal low−wa ter off shoot of the Alternognathus lin eage. As shownby Beinert et al. (1971) there is a con ti nu ity be tween the mid Famennian Scaphignathus and TournaisianClydagnathus lin eages. It has to be ad mit ted that there is some un cer tainty whether the ax ial sym me try orig i −nated only once at the base of the whole clade. Whereas Pandorinellina seems to be truly the best root for theAlternognathus branch, the old est mem ber of the Pseudopolygnathus lin eage (P. jugosus) shows ex pandedbasal cone sim i lar rather to Dasbergina stabilis, the spe cies stratigraphically pre ced ing the lin eage ofPseudopolygnathus but not Alternognathus. The family defined as above may thus be polyphyletic.

146 JERZY DZIK

Fig. 106. Alternognathus pseudostrigosus (Dreesen et Dusar, 1974), prob a bly the ear li est cavusgnathid, from the K. crepidaZone at Kadzielnia (sam ple Ka−10) in the Holy Cross Moun tains. P1 (A–C), P2 (D), S0 (E), S1 (F), S2 (G), S3–4 (H), and M (I) el e −

ments; spec i mens ZPAL cXVI/2236–2244 (holotype B), re spec tively.

Genus Alternognathus Ziegler et Sandberg, 1984

Type spe cies: Alternognathus regularis Ziegler et Sandberg, 1984 from the Famennian Three Forks For ma tion of Montana.

Di ag no sis. — P1 el e ment with plat form de vel op ing trans verse ridges al most at the same level as the tipsof nearby denticles of the blade, dis tinct shal low lar val basal cav ity of fusiform outline.

Re marks. — The old est oc cur rence of the Alternognathus lin eage in the Holy Cross Moun tains is sam pleMd−2 from Miedzianka, where it co−oc curs with C. (C.) inflexoidea, thus older than the en try of P. trachytera.P1 el e ments in this sam ple usu ally show very weak de vel op ment of the plat form, which re mains in cip i ent evenin large spec i mens. A plat form−icrion de vel oped stron ger on one side of the blade and ax ial sym me try may bein volved there. In bed 24 at Jabłonna the lin eage co−oc curs with P. trachytera, but the plat form re mains rel a −tively nar row. Ma ture el e ments with very wide plat form oc cur abun dantly in sam ple Wzd−10 at Ściegnia. Ju ve −niles are vir tu ally miss ing there. It re mains thus un de cided whether the dif fer ence in width of the plat form be −tween sam ples re sults from evo lu tion or only expresses differences in population dynamics.

The P2 el e ments of Alternognathus dif fer from those of as so ci ated Pseudopolygnathus, Dasbergina andPandorinellina in a sig nif i cantly more prom i nent dor sal pro cess. The dif fer ences be tween these gen era in the


Fig. 107. Early cavusgnathids from the Holy Cross Moun tains. A, B. Alternognathus pseudostrigosus (Dreesen et Dusar, 1974)from the C. quadrantinodosa Zone at Miedzianka (sam ple Md−2). P1 el e ments; spec i mens ZPAL cXVI/2346, 2347. C–H. Alter −no gnathus regularis Ziegler et Sandberg, 1984 from the P. trachytera Zone at Jabłonna (bed 24). P1 (A–F), P2 (G), and M (H) elements; spec i mens ZPAL cXVI/2351, 2350, 2349, 2348, 2352, and 2353, re spec tively. I–L. Scaphignathus velifer Helms,

1959 from the same sam ple. P1 el e ments; spec i mens ZPAL cXVI/2355–2357 and 2354.

rest of the ap pa ra tus are mi nor and it is hardly pos si ble to sep a rate them in sam ples where they oc cur insimilar numbers.

Alternognathus pseudostrigosus (Dreesen et Dusar, 1974)(Figs 106, 107A, B, and 137)

Type ho ri zon and lo cal ity: Sam ple 4 from the Hamoir−Fairon sec tion near Liege, Bel gium.

Ma te rial. — 114 spec i mens.Di ag no sis. — Ir reg u lar, strongly asym met ric in cip i ent plat form lo cated high on the blade slightly be low

the tips of its denticles in ma ture P1 el e ments; el e ments of the ap pa ra tus ro bust of Pandorinellina type. Re marks. — This is the ear li est of the more or less cer tain mem bers of the Alternognathus lin eage. In its

strati graphic oc cur rence it sig nif i cantly pre dates other spe cies of Alternognathus and spe cies of Pseudo −polygnathus with ro bust icrion. The struc ture re ferred to as an in cip i ent plat form may ap proach icrion insome el e ments and is so ir reg u larly de vel oped that it is hard to say whether ax ial sym me try is al ready de vel −oped in P1 el e ments of this spe cies, or not. There seems to be an evo lu tion ary change in the Holy CrossMoun tains ma te rial; the stratigraphically youn ger spec i mens show a better de vel oped plat form orig i nat ingear lier in the on tog eny and not so high on the blade. The el e ments are also less robust and somewhat“arthritic” in appearance.

Oc cur rence. — The late K. crepida Zone at Kadzielnia and the C. quadrantinodosa Zone at Miedzianka(Md−2).

Alternognathus regularis Ziegler et Sandberg, 1984(Figs 107C–H, 108, and 137)

Type ho ri zon and lo cal ity: Top of the Tri dent Mem ber of Three Forks For ma tion at Sheep Moun tain in Centennian Rangeof Montana (Ziegler and Sandberg 1984).

Ma te rial. — 1,390 spec i mens.Di ag no sis. — Well de vel oped but rather nar row plat form of P1 el e ment with large nodes along the

margin.Re marks. — The spe cies was re ferred to as Scaphignathus subserratus un til Ziegler and Sandberg

(1984) ques tioned the spe cies iden tity of the lectotype. The in cip i ent ax ial sym me try of the el e ments re fersonly to the shape of the plat form and is more clearly vis i ble on ju ve niles than on large spec i mens with a wider plat form. The blade re mains curved in mirror−image pairs.

Oc cur rence. — The C. quadrantinodosa to early P. jugosus zones at Łagów, Kowala, Ostrówka, andMiedzianka.

Alternognathus? sp.(Fig. 110H–L)

Ma te rial. — Seven spec i mens.Re marks. — Rare P1 el e ments from Dzikowiec are of ju ve nile A. regularis size and mor phol ogy but

their free blade is shorter, re sem bling thus much more ontogenetically ad vanced and larger spec i mens of thatspe cies. They may thus rep re sent adults of a sep a rate species.

Oc cur rence. — The late P. jugosus Zone at Dzikowiec in the Sudetes.

Alternognathus beulensis Ziegler et Sandberg, 1984(Figs 9A, 109, and 137)

Type ho ri zon and lo cal ity: Top of the Hemberg Lime stone at the top of Beul Moun tain near Balve in the Rhenish SlateMoun tains (Ziegler and Sandberg 1984; Beinert et al. 1971).

Ma te rial. — 202 spec i mens.Di ag no sis. — Wide plat form or na mented with ridges and nodes.Re marks. — This is the most ad vanced spe cies of the Alternognathus lin eage, prob a ble a suc ces sor of A.

regularis and this is con sis tent with the strati graphic suc ces sion of these spe cies in the Holy Cross Mountains.Al though the non−plat form el e ments of the Alternognathus ap pa ra tus are of a rather gen er al ized mor phol −

ogy, P2 el e ments with their straight long pro cesses may be of some use in de ter min ing the ge neric affinity.Oc cur rence. — The P. trachytera to L. styriacus zones at Jabłonna, Ściegnia, Miedzianka, and Ostrówka.

148 JERZY DZIK

Genus Scaphignathus Helms, 1959Type spe cies: Scaphignathus velifera Helms, 1959 from the mid Famennian of the Saalfeld area.


Fig. 108. Cavusgnathid Alternognathus regularis Ziegler et Sandberg, 1984 from the early P. trachytera Zone at Kowala (A–G,sam ple Ko−8a) and the L. styriacus Zone at Miedzianka (H–R, sam ple Md−14) in the Holy Cross Moun tains. P1 (A, H–P), P2 (Q),and M (R) el e ments; spec i mens ZPAL cXVI/527, 528, 531–533, 535, 534, 2358, 2360, 2365, 2359, 2367, 2368, 2366, 2364,

2361, 2363, and 2362, re spec tively.

Di ag no sis. — Strongly asym met ric icrion in P1 el e ments, with tu ber cles of lat eral and mid dle rows at ap −prox i mately the same level, high free blade.

Re marks. — Scaphignathus dif fers from Alternognathus mainly in vir tu ally equal height of tu ber cles ofthe plat form and those of the cor re spond ing part of the blade. In fact, at early stages of the on tog eny this isnot a true plat form but rather an icrion. This dif fer ence is ap par ent even in the geo log i cally old est sam pleswith spe cies of those gen era co−oc cur ring (e.g., J−24), al though some spec i mens re main dif fi cult to tell apart.There was prob a bly no dif fer ence in the ap pa ra tus struc ture be tween those erliest mem bers of the lin eage. Ingeo log i cally youn ger sam ples two kinds of P2 el e ments seem to oc cur, one with a rather gen er al ized mor −phol ogy be long ing to Alternognathus and the other with a very reg u lar and low denticulation, prob a bly rep −re sent ing Scaphignathus. Early oc cur rences of Scaphignathus in the Holy Cross Moun tains dif fer from thosefrom youn ger strata in a much nar rower icrion. In the icrion be ing de vel oped dor sally mostly on one side, theP1 el e ments ex hibit an ax ial sym me try, al though some bend ing of the blade allows recognition of particularmembers of the mirror image pair.

Scaphignathus velifer Helms, 1959(Figs 110G and 137)

Type ho ri zon and lo cal ity: 4.5 m be low the top of the finely nod u lar lime stone Unit 5 at Bohlen, as sem blage dom i nated byPalmatolepis schindewolfi and Conditolepis falcata (Helms 1959).

150 JERZY DZIK

Fig. 109. Cavusgnathid Alternognathus beulensis Ziegler et Sandberg, 1984 from from the P. trachytera Zone at Ściegnia(Wzdół Plebański; sam ple Wzd−10; homeomorph of the spe cies?) and the L. styriacus Zone at Miedzianka (A, B, sam ple Md−14) and Ostrówka (C–I, sam ple Ost−15) in the Holy Cross Moun tains. P1 (A–G) and P2 (H, I) el e ments; spec i mens ZPAL

cXVI/2370–2376, 2378, and 2377, re spec tively.

Ma te rial. — 50 spec i mens.Di ag no sis. — Icrion in P1 el e ments grad u ally ta per ing dor sally, with nar row tip.Re marks. — Alternognathus costatiformis of Matyja (1993), with its vir tu ally equal height of lat eral and

me dial denticles in the icrion and a dis tinct sep a ra tion of icrion from the free blade, is so sim i lar toScaphignathus that it prob a bly be longs in this genus.

Oc cur rence. — The C. quadrantinodosa to P. trachytera zones at Łagów and Jabłonna.

Scaphignathus leptus Ziegler et Sandberg, 1984(Figs 110A–F and 137)

Type ho ri zon and lo cal ity: Up per trachytera Zone, Hemberg Lime stone at Ballberg near Balve (Ziegler and Sandberg1984).

Ma te rial. — 77 spec i mens.Di ag no sis. — Par al lel mar gins of the icrion and rel a tively blunt dor sal end in ma ture P1 el e ments.Re marks. — Be cause of sig nif i cant changes in the on tog eny of P1 el e ments and pop u la tion vari abil ity

the avail able ma te rial of rel a tively few spec i mens from many sam ples is too small to be sure of spe cies dis −tinc tions in this ge nus. The pro posed di ag no ses re main thus pro vi sional un til a more pre cise pic ture of theirevolution is achieved.

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


Fig. 110. Cavusgnathids. A–F. Scaphignathus leptus (Ziegler et Sandberg, 1984) from the L. styriacus Zone at Miedzianka (sam −ple Md−14, A) and Ostrówka (B, D–F, sam ple Ost−12; C, sam ple Ost−1a) in the Holy Cross Moun tains. P1 (A–E) and P2 (F) el e −ments; spec i mens ZPAL cXVI/2369, 2379, 2504, and 2380–2382, re spec tively. G. Scaphignathus velifer Helms, 1959 from theC. marginifera Zone at Łagów (G, sam ple Ł−5) in the Holy Cross Moun tains. P1 el e ment; spec i men ZPAL cXVI/2503.H–L. Alternognathus? sp. from the P. jugosus Zone at Dzikowiec (H, sam ple Dz−10; I–L, sam ple Dz−54) in the Sudetes. P1 el e −ments, spec i mens ZPAL cXVI/2387, 2990–2992, and 2989. M, N. Pinacognathus? praesulcatus (Sandberg, 1972) from the D.

trigonica Zone at Kowala (sam ple Ko−166) in the Holy Cross Moun tains. P1 el e ments, spec i mens ZPAL cXVI/2389 and 2390.

Genus Pseudopolygnathus Branson et Mehl, 1934Type spe cies: Pseudopolygnathus prima Branson et Mehl, 1934 from the Tournaisian of Mis souri.

Di ag no sis. — Icrion in P1 el e ments com posed of rows of tu ber cles and fre quently a lon gi tu di nal ridge,basal cav ity wide and shallow.

Re marks. — The De vo nian spe cies at trib uted here were clas si fied in Bispathodus Müller, 1962, al −though its type spe cies Spathodus spinulicostatus E. R. Branson, 1934 is of Early Car bon if er ous age. Itsholotype, sim i lar to that of S. costatus, is prob a bly the end mem ber of the vari abil ity range of the co−occuringtype spe cies of Pseudopolygnathus, P. pri mus, as sug gested by sam ples from else where (Dzik 1997). What −ever is the real na ture of this spec i men, these gen era are clearly closely re lated and there is no way to dis tin −guish them in a satisfactory way.

Ac cord ing to Ziegler et al. (1974), the icrion de vel oped in de pend ently in the branch of P. jugosus and P.ziegleri from the an ces tral sit u a tion of P. stabilis. I have not been able to con firm or ex clude this. How ever,P. ostrovkensis, with its ro bust and wide icrion, ap pears ear lier at Kowala than P. jugosus, much less ad −vanced in this re spect. Per haps Pseudopolygnathus, as un der stood here, is truly polyphyletic and the P.jugosus lin eage should be clas si fied in Dasbergina.

Pseudopolygnathus jugosus (Branson et Mehl, 1934)(Figs 111, 112, and 137)

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

Ma te rial. — 3,111 spec i mens.Di ag no sis. — Icrion in the dor sal part of ma ture P1 el e ment com posed of two rows of denticles and ad di −

tional me dial row of min ute denticles.Re marks. — Icrion de vel oped in this spe cies by wid en ing denticle tips in ju ve niles and shows ax ial sym −

me try ow ing to asym met ric po si tion of the sharp ridge along the ven tral por tion of the blade. Mir ror−im agesym me try is still clearly ex pressed in cur va ture of the blade.

Oc cur rence. — The P. jugosus and D. trigonica zones at Kowala, Jabłonna, Ostrówka, and Dzikowiec.

Pseudopolygnathus ostrovkensis sp. n.(Figs 113 and 137)

Holotype: Spec i men ZPAL cXVI/2298 (Fig. 113F).Type ho ri zon and lo cal ity: Sam ple Ost−185, late Famennian P. jugosus Zone at Ostrówka, Holy Cross Moun tains.Der i va tion of name: From the name of the type lo cal ity.

Ma te rial. — 2,927 spec i mens.Di ag no sis. — Usu ally wide icrion of P1 el e ments; in the dor sal part com posed of three rows of ro bust

denticles, the me dial row be ing of the same height as the lat eral ones.Re marks. — P1 el e ments of this spe cies from the bore hole Kowala have al ready been il lus trated as

Bispathodus ultimus by Nehring−Lefeld (1990, pl. 2: 13, 14). Prob a bly also spec i mens iden ti fied with the

152 JERZY DZIK

Fig. 111. Early prob a ble cavusgnathid Pseudopolygnathus jugosus (Branson et Mehl, 1934) from its nom i nal Zone at Jabłonna(A–C, bed 27) and Ostrówka (D, E, sam ple Ost−185) in the Holy Cross Moun tains. P1 el e ments; spec i mens ZPAL cXVI/2212,

2211, 2210, 2295, and 2296, re spec tively.


Fig. 112. Ad vanced prob a ble cavusgnathid Pseudopolygnathus jugosus (Branson et Mehl, 1934) from its nom i nal Zone atJabłonna (sam ple J−51) in the Holy Cross Moun tains. P1 (A–E), P2 (F, G), S0 (H–J), S1 (K, L), S2 (M, N), S3–4 (O, P), and M (Q, R) el e ments; spec i mens ZPAL cXVI/2225, 2226, 2286–2288, 2227, 2289, 2290, 2228, 2229, 2230, 2291, 2231, 2292, 2232, 2293,

2233, and 2294, re spec tively.

Tournaisian Pseudopolygnathus vogesi by Dreesen et al. (1976, pl. 4: 1–5) from the Famennian of Bel giumare conspecific. There is a pos si bil ity that this is the be gin ning of the P. pri mus lin eage, as sug gested by find −ings of pop u la tions mor pho log i cally close and tran si tional in time (e.g., Bouckaert and Groessens 1976;Sanz−López et al. 1999). How ever, an in de pend ent or i gin of the Car bon if er ous Pseudopolygnathus from aless elab o rate lat est Famennian Pseudopolygnathus seems more likely be cause of the sub stan tial gap in con −

154 JERZY DZIK

Fig. 113. Prob a ble cavusgnathid Pseudopolygnathus ostrovkensis sp. n. from the P. jugosus Zone at Ostrówka (sam pleOst−185) in the Holy Cross Moun tains. P1 (A–I), P2 (J, N), S0 (K), S1 (L), S2 (M), S3–4 (O), and M (P) el e ments; spec i mensZPAL cXVI/2213, 2214, 2303, 2297, 2301, 2298 (holotype, F), 2299, 2302, 2300, 2215, 2305–2307, 2304, 2308, and 2309,

re spec tively.

ti nu ity in the late Famennian and much wider pop u la tion vari abil ity of P. pri mus (cov er ing also morpho −logies typ i cal for the late Famennian) than is ob served in P. ostrovkensis sp. n.

Oc cur rence. — The lat est L. styriacus to D. trigonica zones at Jabłonna, Ostrówka, Kowala, andDzikowiec.

Pseudopolygnathus aculeatus (Branson et Mehl, 1934)(Figs 114A–D and 137)

Type ho ri zon and lo cal ity: Prob a bly the Saverton Shale near Mon roe City, Mis souri (Ziegler 1975, p. 17).

Ma te rial. — 1,014 spec i mens.Di ag no sis. — Denticles de vel oped only on one side of the sharp ridge formed by the blade; left and right

el e ments only slightly dif fer in their ax ial sym me try.Re marks. — This is prob a bly a suc ces sor of P. jugosus, but it ap par ently orig i nated as a re sult of

allopatric speciation and their ranges par tially over lap in Pol ish sec tions. They prob a bly were adapted to dif −fer ent en vi ron ment, but the na ture of these dif fer ences re main to be clarified.

Oc cur rence. — Late P. jugosus to the end of the Famennian in all stud ied lo cal i ties.


Fig. 114. Prob a ble cavusgnathids Pseudopolygnathus in the Holy Cross Moun tains. A–D. P. aculeatus (Branson et Mehl, 1934)from the P. jugosus (A, B, sam ple Ko−54) and D. trigonica (C, D, sam ple Ko−110) zones at Kowala. P1 el e ments; spec i mensZPAL cXVI/2332, 2333, 2321, and 2320, re spec tively. E–N. P. ziegleri (Rhodes, Aus tin et Druce, 1969) from the D. trigonicaZone at Ostrówka (sam ple Ost−3). P1 (E–I), P2 (J), S0 (K), S2 (L), S3–4 (M), and M (N) el e ments; spec i mens ZPAL cXVI/2322,

2324, 2325, 2323, and 2326–2331, re spec tively.

Pseudopolygnathus ziegleri (Rhodes, Austin et Druce, 1969)(Figs 114E–N and 137)

Type ho ri zon and lo cal ity: Up per costatus Zone in the Hönnetal road cut near Balve, Rhenish Slate Moun tains (Ziegler1975).

Ma te rial. — 112 spec i mens.Di ag no sis. — Icrion in the dor sal part of P1 el e ments com posed of three rows of denticles; the me dial

higher than the lat eral ones that tend to de velop trans verse ridges.Re marks. — Left and right el e ments show dif fer ent mor phol ogy of the icrion, as al ready shown by

Ziegler and Sandberg (1984) clas si fy ing the lat ter un der the name of Bispathodus ultimus. In Kowala P.ziegleri was re placed by P. ultimus be tween de po si tion of strata rep re sented by sam ples Ko−114 and Ko−113,thus af ter emer gence of D. trigonica, whereas at Dzikowiec P. ultimus ap pears be fore D. trigonica. Prob a blythis change was con trolled eco log i cally and not di rectly re lated to evo lu tion, as sug gested by the re−ap pear −ance of P. ziegleri be fore the end of the Famennian at Kowala.

Oc cur rence. — The late P. jugosus Zone at Kowala, Ostrówka, and Dzikowiec.

Pseudopolygnathus ultimus (Bischoff, 1957)(Figs 115 and 137)

Type ho ri zon and lo cal ity: The type lo cal ity of the Wocklumeria beds at Wocklum, Rhenish Slate Moun tains (Bis choff1957; Ziegler 1975).

Ma te rial. — 1,333 spec i mens.Di ag no sis. — The lat eral rows of denticles in the dor sal part of the icrion of P1 el e ments trans formed into

wide trans verse ridges sep a rated in the mid dle by a lon gi tu di nal es carp ment, some times form ing also a lowcrest.

Re marks. — Left and right el e ments dif fer in width of icrion and prom i nence of its rib bing. The spe ciesseems to be a di rect suc ces sor of P. ziegleri but the tran si tion was prob a bly long−last ing and pop u la tion vari −abil ity wide. It is thus hardly of much cor re la tive value in stratigraphy.

Oc cur rence. — The lat est P. jugosus and D. trigonica zones at Kowala and Dzikowiec.

Family Elictognathidae Austin et Rhodes, 1981Di ag no sis. — Sec ond arily lost ax ial sym me try in P1 el e ments.

156 JERZY DZIK

Fig. 115. Prob a ble cavusgnathid Pseudopolygnathus ultimus (Bis choff, 1957) from the P. jugosus Zone at Dzikowiec (A–H,sam ple Dz−17) in the Sudetes and the D. trigonica (I, J, sam ple Ko−110) Zone at Kowala in the Holy Cross Moun tains. P1 (A–E, I,

J), P2 (F), S2 (G), and S3–4 (H) el e ments; spec i mens ZPAL cXVI/2310–2319, re spec tively.

Re marks. — Clas si fi ca tion of Siphonodella in this fam ily was pro posed by Sweet (1988) based on theas sump tion that its ap pa ra tus P2 lo ca tion was oc cu pied by el e ments ear lier clas si fied as Elictognathus. Thiswas fol lowed by my self (Dzik 1994), al though only one of the Siphonodella−like Tournaisian cono donts may pos si bly fit such an ap pa ra tus con cept. P2 el e ments of Elictognathus mor phol ogy co−oc cur with ramiform el −e ments clas si fied in Dinodus, the ge neric name hav ing pri or ity. The De vo nian el e ments with a mor phol ogyre sem bling Dinodus can hardly be long to an ap pa ra tus re lated to the Siphonodella lin eage and prob a bly arerather of prioniodinid af fin i ties (Guizhoudella). The de rived mor phol ogy of ap pa ra tus el e ments of Siphono −della prob a bly did not orig i nate be fore the Tournaisian. Pos si bly, the ear li est elictognathids were quite dif fer −ent and their roots are to be looked for in Immognathus. All this in tro duces much uncertainty to classificationof those conodonts and requires clarification.

Genus Pinacognathus Branson et Mehl, 1934Type spe cies: Pinacodus profundus Branson et Mehl, 1934 from the Bushberg Sand stone of Mis souri.

Di ag no sis. — P1 el e ments with lan ceo late, flat or in verted basal cav ity sep a rated by an es carp ment fromthe rest of the plat form base; ramiform el e ments with rel a tively short processes.

Pinacognathus? praesulcatus (Sandberg, 1972)(Figs 110M, N and 129)

Type ho ri zon and lo cal ity: Top of the Sappington Mem ber of Three Forks For ma tion at Lick Creek Road, Lit tle Belt Moun −tains, Montana.

Ma te rial. — 17 spec i mens.Re marks. — Ma te rial at my dis posal is rather poor and only sam ple Ko−116 con tains enough spec i mens

of var i ous ontogenetic age to see their vari abil ity. It ap pears that ju ve niles have a plat form usu ally sep a ratedfrom the blade by a deep de pres sion and the el e ment has the great est width at the dor sal end of the plat form.In ma ture spec i mens the mid dle part of the plat form is the wid est and it is al most flat with the denticles of theblade some what el e vated. El e ments show mir ror sym me try that makes un likely any re la tion ship to Alterno −gnathus. Also the flat or some what in verted basal cav ity makes it dif fer ent from Alternognathus, where thebase re mains slightly con cave and the mar gin of the basal cone re mains sharp. Vari abil ity is ex pressedmostly in more or less ro bust trans verse rib bing of the plat form and in the dor sal mar gin of the plat form, inju ve niles be ing thin and wide or rather narrow and robustly tuberculated. The apparatus remains unknown.

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

Family Idiognathodontidae Harris et Hollingsworth, 1933Di ag no sis. — Gen er al ized late Palaeozoic cono donts with wide basal cav ity in P1 el e ments and ten dency

to de velop a plat form at the basal cone or icrion in the dor sal part of the el e ment; in post−De vo nian forms theex ter nal pro cess of S3–4 el e ments is arched and recurved.

Genus Protognathodus Ziegler, 1969Type spe cies: Gnathodus kockeli Bis choff, 1957 from the Gattendorfia Stufe in the Rhenish Slate Moun tains.

Di ag no sis. — Widely gap ing and rather deep basal cav ity reach ing the dor sal tip of P1 el e ment, gentlyarched pro file of the blade.

Protognathodus kockeli (Bischoff, 1957)(Figs 116M, N and 138)

Type ho ri zon and lo cal ity: Lower part of the Gattendorfia Lime stone at Wocklum in the Rhenish Slate Moun tains.

Ma te rial. — 42 spec i mens.Re marks. — Spec i mens from the top most Famennian strata at Kowala are dif fi cult to clean from clayish

cover and thus are not suit able for SEM pho tog ra phy. They show tuberculation of the basal cone typ i cal ofthe early Tournaisian pop u la tions of Protognathodus (e.g., Dzik 1997).

Oc cur rence. — Ter mi nal Famennian at Kowala (sam ples Ko−51 and Ko−24; Dzik 1997) and Dzikowiec(Dz−75).


Genus Dasbergina Schäfer, 1976

Type spe cies: Dasbergina ziegleri Schäfer, 1976 from the bed 93 of a trench at Dasberg, lower costatus Zone.

Di ag no sis. — Very shal low con i cal basal cav ity, var i ously de vel oped plat form around the basal cone, ap −pa ra tus of gen er al ized morphology.

Re mark. — Early spe cies of the ge nus, with weakly de vel oped plat form, are tran si tional to the prob a blyan ces tral spe cies of Pandorinellina, P. vulgaris. The dif fer ence be tween tran si tional pop u la tions is only inthe width of the basal cone, in fact, very variable.

Dasbergina stabilis (Branson et Mehl, 1934)(Figs 116 and 138)

Type ho ri zon and lo cal ity: Prob a bly the Saverton Shale near Mon roe City, Mis souri (Ziegler 1975, p. 47).

Ma te rial. — 4,491 spec i mens.Di ag no sis. — P1 el e ment lack ing plat form, al most flat cav ity of the basal cone with eye−drop out line,

con tin u ing to the dor sal tip of the blade.

158 JERZY DZIK

Fig. 116. Early idiognathodontids. A–L. Dasbergina stabilis (Branson et Mehl, 1934) from the P. jugosus Zone at Jabłonna(A–D, bed 27) and Ostrówka (E–L, sam ple Ost−185) in the Holy Cross Moun tains. P1 (A–C, E, F), P2 (D, G), S0 (H), S1 (I), S2 (J),S3–4 (K), and M (L) el e ments; spec i mens ZPAL cXVI/2206–2209 and 2261–2268, re spec tively. M, N. Protognathodus kockeli

(Bis choff, 1957) from Dzikowiec (sam ple Dz−75). P1 el e ments, spec i mens ZPAL cXVI/2965 and 2964.

Re marks. — The ap pa ra tus of the spe cies was re stored by Over (1992). The basal cav ity var ies from be −ing vir tu ally flat to dis tinctly con i cal. Spec i mens with very flat base dif fer from those of Pandorinellinavulgaris in their much lower blade and a ten dency to de velop an in cip i ent plat form of low el e va tion con tin u −ing to the tip of the el e ment. From as so ci ated ju ve nile spec i mens of Dasbergina micropunctata such el e −ments dif fer in hav ing their base (and “plat form”) grad u ally nar row ing to the tip while in D. micropuntata the base has a pear−like out line and ter mi nates significantly in front of the element tip.

The old est spec i mens of D. stabilis in the densely sam pled part of the Kowala sec tion have been found insam ple Ko−187. Un for tu nately spec i mens clas si fied in P. vulgaris from the sam ple im me di ately be low arefrag men tary, ham per ing biometrical pre sen ta tion of the change.

Oc cur rence. — The P. trachytera to D. trigonica zones at Jabłonna, Ostrówka, Kowala, and Miedzianka,the Holy Cross Moun tains, and Dzikowiec, the Sudetes.

Dasbergina micropunctata (Bischoff et Ziegler, 1956)(Figs 117 and 138)

Type ho ri zon and lo cal ity: Dark lime stone of toV from the “Rote Scheid” quarry near Mar burg (Bis choff and Ziegler 1956).


Fig. 117. Idiognathodontid Dasbergina micropunctata (Bis choff et Ziegler, 1956) from the P. jugosus Zone at Jabłonna (A–C, sample J−52) and Ściegnia (D–L ,sam ple Wzd−13) in the Holy Cross Moun tains. P1 (A–F), P2 (G), S0 (I, J), S1 (K), S2 (H), S3–4 (L),and M (M) el e ments; spec i mens ZPAL cXVI/2235, 2197, 2196, 2234, 2251, 2250, 2252, 2254, 2256, 2255, 2253, and 2257, 2258,

re spec tively.

Ma te rial. — 1,733 spec i mens.Di ag no sis. — The basal cone of P1 el e ment with a smooth tri an gu lar plat form, basal cav ity nar rows in

front of the dor sal tip of the blade.Re marks. — Some P1 el e ments have plat form with a com pletely smooth sur face, oth ers show the re tic u −

late pat tern of ameloblast im prints. The mean ing of this dif fer ence is un clear. Triramous S1 el e ments may be −long to this species.

Oc cur rence. — The L. styriacus to D. trigonica zones at Jabłonna, Ostrówka, Kowala, Miedzianka, theHoly Cross Moun tains, and Dzikowiec, the Sudetes.

160 JERZY DZIK

Fig. 118. Idiognathodontid Dasbergina granulosa (Ziegler, 1962) from the L. styriacus Zone at Ostrówka (A–C, I–K, sam ple 1a; D–H, sam ple Ost−12) in the Holy Cross Moun tains. P1 (A–H), P2 (J), and S3–4 (I, clus ter of two el e ments, K) el e ments; spec i mens

ZPAL cXVI/2271, 2270, 2269, 2275, 2278, 2276, 2277, 2274, 2345, 2272, and 2273, re spec tively.

Dasbergina granulosa (Ziegler, 1962)(Figs 118 and 138)

Type ho ri zon and lo cal ity: Bed 0 in the Sessacker II trench, P. trachytera Zone.

Ma te rial. — 1,732 spec i mens.Di ag no sis. — The plat form of ir reg u larly an gu lar shape or na mented with nu mer ous tu ber cles, basal cav −

ity forms lobes on both sides of the P1 element.Re marks. — This is one of the most ro bust and larg est Famennian cono donts. Ma ture P1 el e ments bear a

very wide plat form but the con i cal basal cav ity grows to ma tu rity. Some spec i mens from Kowala andDzikowiec, with very vari able or na men ta tion and the plat form ex tend ing far out side the basal cone, but re −stricted to the cen ter of the el e ment, may be tran si tional to Dasbergina sp. aff. D. kayseri.

Oc cur rence. — The P. trachytera to P. jugosus zones at Jabłonna, Ostrówka, Kowala, Miedzianka, theHoly Cross Moun tains, and Dzikowiec, the Sudetes.

Dasbergina kowalensis sp. n.(Figs 119A–C and 138)

Holotype: Spec i men ZPAL cXVI/2256 (Fig. 119B).Type ho ri zon and lo cal ity: Sam ple Ko−1, late Famennian early P. jugosus Zone at Kowala, Holy Cross Moun tains.Der i va tion of name: From the latinized name of the type lo cal ity.

Ma te rial. — 27 spec i mens.Di ag no sis. — In cip i ent plat form of ir reg u larly an gu lar shape or na mented with a few large tu ber cles,

basal cav ity forms wide lobes on both sides of the P1 element.Re marks. — This seems to be the old est mem ber of the D. trigonica lin eage, per haps a suc ces sor of D.

stabilis, al though the tran si tion has not been dem on strated. The basal cone or na mented with tu ber cles makesthe spe cies sim i lar to Protognathodus. This is prob a bly only a con ver gence; the dif fer ence is that the basalcav ity is ex panded only in prox im ity to the cusp and strongly narrows dorsally.

Oc cur rence. — The early P. jugosus Zone at Kowala.

Dasbergina marburgensis (Bischoff et Ziegler, 1956)(Figs 119D–I and 138)

Type ho ri zon and lo cal ity: Dark lime stone of toV unit from the quarry NE Weitershausen in the Rhenish Slate Moun tains.

Ma te rial. — 190 spec i mens.Di ag no sis. — Tri an gu lar plat form of P1 el e ment or na mented with ro bust mar ginal denticles, tend ing to

form ir reg u lar trans verse rows near the el e ment cen ter; trans versely elon gated lobes of basal cav ity withparallel margins.

Re marks. — Per haps the most char ac ter is tic fea ture of the spe cies is the basal cav ity of the P1 el e ment.Its nar rowly elon gated an te rior lobe has a rounded tip, whereas the pos te rior lobe var ies from nar rowlyrounded to an gu lar, with the ven tral an gle more prominent.

Oc cur rence. — The late P. jugosus Zone at Ostrówka, Kowala, and Dzikowiec.

Dasbergina trigonica (Ziegler, 1962)(Figs 119J–N and 138)

Type ho ri zon and lo cal ity: Sam ple 3a from the Hönnetalstraße sec tion in the Rhenish Slate Moun tains.

Ma te rial. — 204 spec i mens.Di ag no sis. — The basal cav ity of P1 el e ment bi fur cates an te ri orly.Re marks. — In suc ces sive sam ples in the up per part of the Dzikowiec sec tion the an te rior lobe of the

basal cav ity of the P1 el e ment, orig i nally round (Dz−18), grad u ally becames more an gu lar (Dz−10, Dz−17,Dz−20b) and then its tip nar rows and the lobe bi fur cates (Dz−19, Dz−47, Dz−4). This marks the tran si tion from D. marburgensis to D. trigonica. This char ac ter shows a sig nif i cant pop u la tion vari abil ity and ju ve niles mayshow an gu lar wide lobes even in sam ples from high above in the section (e.g., Dz−8).

The spe cies oc curs in the bore hole Kowala, as shown by Nehring−Lefeld (1990).Oc cur rence. — Zone of its own at Kowala and Dzikowiec.


162 JERZY DZIK

1

2

Fig. 119. Idiognathodontids of the Dasbergina trigonica and D. kayseri lin eages. A–C. D. kowalensis sp. n. from the early P.jugo sus Zone at Kowala (sam ple Ko−1). P1 el e ments; spec i mens ZPAL cXVI/2245–2247 (holotype, B). D–I. Dasberginamarburgensis (Bis choff and Ziegler, 1956) from the late P. jugosus Zone at Ostrówka (D, E, sam ple Ost−185) and Dzikowiec(F–I, sam ple Dz−10; late form). P1 (D–G, I) and P2 (H) el e ments; spec i mens ZPAL cXVI/2280, 2279, 2977–2980, and 2979, ®

Dasbergina sp. aff. D. kayseri (Bischoff et Ziegler, 1956)(Fig. 119O, P)

Re marks. — At Kowala be low the oc cur rence of Dasbergina kayseri an un named spe cies oc curs with awide asym met ric basal cav ity re sem bling that of adult Dasbergina granulosa and with a higly vari abletuberculation of the plat form that may be al most smooth in ju ve niles (e.g., Perri and Spalletta 1991, pl. 9: 5).It even seems that in suc ces sive sam ples the basal cone be came smaller and more asym met ric, al though toofew spec i mens are known to be sure that this is not a case of very wide pop u la tion vari abil ity. A pos si bil itythus emerges that the lat est Famennian D. keyseri lin eage is an outshoot of Dasbergina, homeomorphic withthe early Famennian ancyrognathids.

Dasbergina kayseri (Bischoff et Ziegler, 1956)(Figs 119Q–Y and 132)

Type ho ri zon and lo cal ity: Dark lime stone of toV from the quarry NE Weitershausen near Mar burg (Bis choff and Ziegler1956).

Ma te rial. — 80 spec i mens.Di ag no sis. — Rel a tively large, asym met ric basal cone (pit); rhomboidal out line of plat form in P1 el e ment

reach ing al most the ven tral end of the blade in ma ture specimens.Re marks. — One spec i men from Kowala (Fig. 119Q) shows a large basal cav ity, tran si tional in size and

shape be tween that in the pre ced ing spe cies and typ i cal D. kayseri. Rare ramiform el e ments as so ci ated withP1 el e ments typ i cal of the spe cies are too gen er al ized to prove its ge neric af fil i a tion, but their mor phol ogydoes not con tra dict af fin i ties with ad vanced Dasbergina.

Oc cur rence. — The P. jugosus Zone at Ostrówka and Kowala.

Dasbergina brevipennata (Ziegler, 1962)(Figs 120A–U and 138)

Type ho ri zon and lo cal ity: Sam ple 1327 from the road sec tion at Hönnetal, late L. styriacus Zone.

Ma te rial. — 782 spec i mens.Di ag no sis. — Plat form of P1 el e ment with reg u larly an gu lar out line and flat sur face or na mented with nu −

mer ous tubercles.Re marks. — Ju ve nile P1 el e ments of the spe cies show a more lan ceo late out line of the plat form than pre −

ced ing spe cies and in this re spect they are rather sim i lar to ju ve niles of D. ziegleri.Oc cur rence. — The L. styriacus Zone at Jabłonna, Kowala, and Ściegnia.

Dasbergina ziegleri Schäfer, 1976(Figs 120V–Y and 138)

Type ho ri zon and lo cal ity: Bed 93 in a trench at Dasberg, lower costatus Zone

Ma te rial. — 310 spec i mens.Di ag no sis. — Wide ro bust plat form ex tend ing al most to the whole length of P1 el e ment, or na mented with

ro bust tu ber cles and ridges.Re marks. — Pseudopolygnathus controversus Sandberg et Ziegler, 1979 is prob a bly conspecific with

this spe cies. No ta bly, the early pop u la tion of D. ziegleri from the L. styriacus Zone (Fig. 120V, W), with arather an gu lar out line of the plat form and its fine tuberculation, re sem bles D. brevipinnata. The late pop u la −tion from the P. jugosus Zone (Fig. 120X, Y) is sim i lar in those re spects rather to Pseudopolygnathus


re spec tively. J–N. Das bergina trigonica (Ziegler, 1962) from its nom i nal zone at Ostrówka (J, K, sam ple Ost−3) in the HolyCross Moun tains and Dzikowiec (L, sam ple Dz−7; M, N, sam ple Dz−74) in the Sudetes. P1 el e ments; spec i mens ZPALcXVI/2281–2283, 2966, and 2967. O, P. Dasbergina sp. aff. kayseri (Bis choff et Ziegler, 1956) from the P. jugosus Zone atKowala (sam ple Ko−192), pos si ble tran si tional forms from un de rived Dasbergina with wide basal cone to D. kayseri. P1 el e −ments; spec i mens ZPAL cXVI/2996 and 2997. Q. Dasbergina kayseri (Bis choff et Ziegler, 1956) early form with wide basalcav ity from the P. jugosus Zone at Kowala (sam ple Ko−197). P1 el e ment; spec i men ZPAL cXVI/2995. R–Y. Typ i calDasbergina kayseri (Bis choff et Ziegler, 1956) from P. jugosus Zone at Ostrówka (H, K, sam ple Ost−2; I, J, L, M, sam pleOst−185) and Kowala (N, O, sam ple Ko−198) in the Holy Cross Moun tains. P1 (R–W), P2 (X), and S3–4 (Y) el e ments; spec i mens

ZPAL cXVI/2998, 2603, 2598, 2599, 2999, and 2600–2602 re spec tively.

164 JERZY DZIK

Fig. 120. Idiognathodontids of the Dasbergina ziegleri branch from the Holy Cross Moun tains. A–U. Dasbergina brevipennata(Zie gler, 1962) from the early P. jugosus Zone at Jabłonna (A–I, bed 27) and the late L. styriacus Zone at Kowala (J–U, sam pleKo−191). P1 (A–C, J–O), P2 (D, P), S0 (E, Q), S1 (F, R), S2 (G, S), S3–4 (H, T), and M (I, U) el e ments; spec i mens ZPALcXVI/2216–2224, 3000–3011, re spec tively. V–Y. Dasbergina ziegleri Schäfer, 1976 from the L. styriacus Zone at Ostrówka(V, W, sam ple Ost−7) and the early P. jugosus Zone at Kowala (X, Y, sam ple Ko−1). P1 el e ments; spec i mens ZPAL cXVI/2284,

2285, 2248, 2249.

ostrovkensis. It seems thus that this is an outshoot of the Dasbergina lin eage homeomorphic with Pseudo −polygnathus (in fact, both are closely related).

Oc cur rence. — The P. jugosus Zone at Kowala and Ostrówka.

FAUNAL DYNAMICS OF THE FAMENNIAN CONODONTS IN POLAND

While re view ing dis tri bu tion of par tic u lar groups of fos sil or gan isms one has to face the con flict be tweenecol ogy and phy log eny. Ob vi ously, the supraspecific tax o nomic units are by def i ni tion based on evo lu tion −ary (that is, ge netic) re la tion ships among spe cies, which do not need to be re lated to their en vi ron men tal pref −er ences. How ever, many Famennian cono dont taxa seem to pre serve some eco log i cal coherencey and the re −view in the in tro duc tory chap ters of this work pre sent ing cono dont ecostratigraphy in the area (Fig. 5, 6) mayserve as an use ful ap prox i ma tion. To ex tract the un bi ased ev i dence on the evo lu tion of lo cal en vi ron ments itwould be nec es sary to ad dress dis tri bu tion of each spe cies sep a rately. This kind of ev i dence is of fered byquan ti ta tive data on their fre quency dis tri bu tion in the part of the work show ing the strati graphic dis tri bu tionof spe cies (Figs 121–138). Some gen eral com ments on the global en vi ron men tal change as seen from a Pol −ish per spec tive are pre sented. This will be fol lowed by a re view of the Famennian evo lu tion ary di ver si fi ca −tion of conodonts, with an attempt to be free as much as possible from its preservational and ecological bias.

SUCCESSION OF THE FAMENNIAN CONODONT FAUNAS

Cono dont spe cies with ap pa ra tuses con tain ing plat form el e ments rep re sent only a frac tion of the com −plete late De vo nian cono dont di ver sity. This is to a great de gree oblit er ated by un bal anc ing of fos sil sam ples,that is their en rich ment in ro bust plat form− or icrion−bear ing el e ments as a re sult of hy dro dy namic sort ingand/or frag men ta tion of frag ile ramiform el e ments of the ap pa ra tus. To re store the orig i nal nu mer i cal con tri −bu tion of spe cies to the bi o log i cal pro duc tiv ity of the cono dont com mu nity this bias should be re moved. Letus at tempt to do this by re plac ing counts of plat form el e ments to a se lected sam ple by num bers de rived fromcounts of non−plat form el e ments of the same spe cies. In the stan dard cono dont ap pa ra tus with 15 el e ments,the P1 el e ments rep re sent ap prox i mately 13% of all el e ments. To com pen sate for the pos si bly greater fra gil −ity of ramiform el e ments in such ap pa ra tuses in re spect to those with out plat forms, lets as sume that the num −ber of P1 el e ments ac tu ally found rep re sents 25 % of their orig i nal num ber prior to taphonomic dis tor tion. For rare spe cies known only from plat form spec i mens and for Icriodus their number in the count below has beenonly halved.

If re in ter preted in this way, the ter mi nal Frasnian Kellwasserkalk sam ple Pł−391 rep re sented by 17,804spec i mens (see Dzik 2002) gives a not es pe cially im pres sive pic ture of bi o log i cal di ver sity of the com mu nity(more pre cisely: its bi o log i cal pro duc tion). Only three spe cies sig nif i cantly con trib uted to this: the dom i nantPolygnathus webbi (in clud ing its va ri ety in for mally listed as P. “tuberculatus”, do not re lated to true P.tuberculatus Hinde, 1879) with al most 35% of all el e ments, Icriodus iowaensis with about 30% (per haps more,as the way in which this es ti mate was done is es pe cially un re li able in this case), and the palmatolepididLagovilepis bogartensis with 20% con tri bu tion. Five spe cies con trib uted from 1.5 to 3.9% each, so some ofthem had a chance to be rep re sented in sam ples of stan dard size (spe cies of Ctenopolygnathus, Ancyrodella,Klapperilepis, Ligonodina and Dyminodina). All the re main ing seven spe cies of this high−di ver sity com mu nityeven af ter such com pen sa tion are rep re sented by less then 1%. They are thus un likely to be iden ti fied in a stan −dard cono dont sam ple taken for strati graphic pur poses (spe cies of Pelekysgnathus, Pluckidina, Mehlina,Ancyrognathus, and the palmatolepidids Manticolepis winchelli, M. rhenana and “Conditolepis” lingui formis).

All this shows also how un re li able ranges of ver ti cal dis tri bu tion of taxa may be, even in sit u a tions wherethe fos sil re cord is complete.

Frasnian–Famennian bound ary event. — The dra matic change at the Frasnian–Famennian bound aryseen from this per spec tive fades very much (cf. House 2002; Bambach et al. 2004; Racki 2005). It is still asig nif i cant de crease in di ver sity, but not un prec e dented in the Frasnian re cord. The three ear li est Famenniansam ples (Pł−20, Pł−16, and Pł−15; Dzik 2002), counted to gether in the same way to com pen sate for un bal anc −ing, give a sur pris ingly sim i lar com mu nity struc ture. Again, only three spe cies dom i nate: Icriodus (dif fer ent


spe cies than in the Frasnian, I. alternatus) with its 39% con tri bu tion, Polygnathus (P. praecursor, pos si bly asuc ces sor of the Frasnian P. webbi) con tri bu tion about 30% and the only palmatolepidid Klapperilepis ul timacon tin u ing from the Frasnian, of al most 20% share in the com mu nity struc ture. Also other lin eages prob a blycon tinue from the Frasnian, with two of rea son able con tri bu tion (Ctenopolygnathus, about 10% and Mehlina4%). Pelekysgnathus, Ligonodina, Dyminodina and Pluckidina all have be low a 1% con tri bu tion. It ap pearsthus that ex cept for the ter mi na tion of Lagovilepis bogartensis and re plac ing one spe cies of Icriodus by an −other, noth ing truly un usual hap pened at this al leg edly “one of the great est ex tinc tion events in the World his −tory”, in con trast to the in ter na tion ally−ac cepted def i ni tion of the F–F mass ex tinc tion bound ary (see ref er −ences above). In the Holy Cross Moun tains, where the stromatoporoid−coral reef com mu ni ties of theFrasnian are es pe cially well rep re sented, their growth ter mi nated sig nif i cantly be low this change in cono dont com mu ni ties. There is hardly any rea son to be lieve in a sud den cat a strophic event of ex tra ter res trial cause atthe Frasnian–Famennian bound ary. No doubt, how ever, that a pro found re build ing of ma rine eco sys tems and long−dis tance mi gra tions of pe lagic com mu ni ties took place that time, sug ges tive of a deep change in theglobal cli mate (e.g., Dzik 2002; Joachimski and Buggisch 2002; Joachimski et al. 2001, 2004).

Within−Famennian en vi ron men tal changes. — The ear li est Famennian strata in the Płucki sec tion dif −fer from those be low in the in creased con tri bu tion of black shale in ter ca la tions. It is thus in struc tive to com −pare the first Famennian cono dont com mu nity with that rep re sent ing a sim i lar en vi ron ment of black shale inthe mid Famennian. Sam ple Ko−8a of 1058 spec i mens from the lime stone con cre tions, col lected im me di ately above the black “pa per” shale ex posed in the Kowala quarry rep re sent ing the Platyclymenia annulata event,of fers such an op por tu nity (Fig. 4, 6). Counted in the same way it shows an overdomination of theprioniodinid (or the ear li est non−plat form gondolellid) Branmehla (57%), two other spe cies, the palmato −lepidid Tripodellus schleizius and the cavusgnathid Alternognathus regularis con trib uted 12% and 10%, re −spec tively. All other spe cies (of gen era Icriodus, Mitrellataxis, Idioprioniodus, Vogelgnathus, Pandorinel −lina, Mehlina, Polynodosus, Hemilistrona, and Neopolygnathus) are of in sig nif i cant im por tance in the com −mu nity. Less then half of them had platform elements in their apparatuses.

Par a dox i cally, a much more pro found re build ing of the cono dont com mu nity oc curred in time slices notso well noted in con nec tion with pur ported cat a strophic events. The first such fun da men tal trans for ma tion ofthe cono dont com mu ni ties in the Holy Cross Moun tains dur ing the Famennian took place near the end of theK. triangularis Zone. Many new lin eages im mi grated at that time, but most of the pre ced ing di ver sity sur −vived the change. Ob vi ously, this was con nected with a transgressive event im me di ately af ter the sea leveldrop (John son et al. 1985) that in the Holy Cross re gion re sulted in the sub mer sion of the ear lier eroded mas −sive lime stone at Kadzielnia, the sed i men tary dis con ti nu ity sur face at Wietrznia, and the redeposition oflimestone pebbles at Miedzianka.

Less ap par ent and gen er ally stepwise changes in com po si tion of the cono dont com mu nity by im mi gra tion of some and dis ap pear ance of other lin eages took place within the C. quadrantinodosa and C. marginiferazones. Also these events were re lated to pe ri odic low er ing of sea level, as sug gested by the sed i men tary dis −con ti nu ity within this time span in the Łagów sec tion. The change was not con nected with com pe ti tion be −tween mem bers of dif fer ent lin eages but re sulted rather from lat eral shifts of eco sys tems as a result of globalenvironmental changes.

Sev eral events of the sea level rise and in crease in bi o log i cal pro duc tiv ity marked by bi tu mi nous shale de −po si tion (see Walliser 1996; House 2002) are well dis played in the Kowala sec tion. These were iden ti fiedwith the Platyclymenia annulata, Epinette and Hangenberg events by my self (Dzik 1997). They did notcause any dra matic change in the cono dont com mu nity. The changes in their rel a tive con tri bu tion were notmore sig nif i cant than in the whole up per part of the sec tion, where cyclicity of prob a bly Milanković na tureseems to be re corded (Fig. 6). There was a se ries of two sud den drops in di ver sity of cono donts near the endof the Famennian. The first one cor re sponds to the be gin ning of the D. trigonica Zone; from this mo ment theicrion−bear ing Pseudo polygnathus dom i nated the as sem blages. The whole pe lagic fau nal di ver sity was re −duced to a cat a strophic level at the time of sed i men ta tion of the Hangenberg black shale (Dzik 1997; see alsoBrand et al. 2004).

In some as pects this sit u a tion is sim i lar to that at the Frasnian–Famennian tran si tion. The fau nal re place −ment was even more sig nif i cant near the end of the Famennian and the new dom i nant taxa (Protognathoduskockeli and Acutimitoceras) seem to mark an in cur sion of cold wa ters to the area (Dzik 1997). No doubt,how ever, that else where sev eral Famennian lin eages sur vived and par tic i pated in re stor ing the warm−wa ter

166 JERZY DZIK


Fig. 121. Strati graphic dis tri bu tion of the pos si ble protopanderodontid Jablonnodus, icriodontids, and spathognathodontids inthe Pol ish Famennian. Per cent con tri bu tion to sam ples in a com pos ite suc ces sion as sem bled with rep re sen ta tive sec tions in theHoly Cross Moun tains (sam ple num bers given in the left col umn; for spe cies not rep re sented in these sec tions only ver ti cal linede notes their dis tri bu tion) and di a gram matic rep re sen ta tion of iden ti fied ap pa ra tus el e ments is shown. Prob a ble chronospeciesof the same lin eage are ar ranged in the same line; bases of lines cor re spond to im mi gra tion events. Note highly ir reg u lar fre −quency dis tri bu tion of el e ments in par tic u lar lin eages, which makes ob served ex tent of ranges very sen si tive to dif fer ences in

sam ple size and usu ally not truly mean ing ful.

high−pro duc tiv ity com mu ni ties of the Gattendorfia Lime stone (Dzik 1997). Their ex act course of evolutionremains to be traced.

168 JERZY DZIK

Fig. 122. Strati graphic dis tri bu tion of the prioniodinids Ligonodina and Lagovidina in the Pol ish Famennian (the same kind ofpre sen ta tion as on Fig. 121).

EVOLUTION OF THE FAMENNIAN CONODONTS

Most of the change in the fau nal dy nam ics of cono donts in the Famennian of Po land ap pears thus to becaused by mi gra tions, not un like other ex am ples of the fos sil re cord in trop i cal en vi ron ments. Only in a fewcases a phyletic evo lu tion ary change is re corded clearly enough to en able quan ti ta tive stratophenetic de scrip −tion. It is be yond the scope of the pres ent work to use stratophenetics to show the evo lu tion of the fauna. Itsmain goal is to pres ent ap pa ra tus re con struc tions. Their knowl edge is still in so early a stage that more ba sicques tions on their re la tion ships haves to be clar i fied first. How ever, it has to be pointed out that the avail ableev i dence sup ports a gen er ally grad ual pat tern of evo lu tion, un less it was trun cated by sud den changes in eco −log i cal con di tions or non−de po si tion events (Dzik 2005; com pare with Donoghue 2001). A brief over view ofprob a ble phylogeny of particular Famennian conodont taxa is given below.

Mitrellataxids. — The first coni form Mitrellataxis emerges in the Holy Cross Moun tains to gether withPalmatolepis initialis. More elab o rate M. ornata fol lowed soon and this may be an evo lu tion ary suc ces sion.It oc curred in prox im ity of the Karczówka mudmound anal o gous to the Frasnian belodellids. The or der in ap −pear ance of smooth “sim ple cones” is from M. circularis, through Jablonnodus erec tus to J. oistodiformis(Fig. 121). The end mem ber of the Jablonnodus lin eage is prob a bly the un named late Famennian spe cieswith geniculate el e ments il lus trated by Sandberg and Dreesen (1984). If taken lit er ally this would sug gestthat the mor pho log i cal dis tinc tions within the ap pa ra tus typ i cal of the last spe cies de vel oped grad u ally. How −ever, no real tran si tion has been dem on strated and this suc ces sion may have beeen con trolled eco log i cally.The main lin eage of M. circularis, al though rel a tively well rep re sented through out the suc ces sion, is too in −dif fer ent mor pho log i cally to be in ter preted in evo lu tion ary terms. Any way, a sec ond ary homeomorphy withthe Or do vi cian distacodontids re mains a valid alternative to inheritance of ancient characters in a lineage of“living fossils”.

Icriodontids. — Such “liv ing fos sils” are prob a bly the most ar chaic Famennian icriodontids Lateri −criodus, ep i sodic im mi grants from an un known source area.

The pop u la tion of the ear li est Famennian Icriodus alternatus (sam ple Pł−16) dif fers mor pho log i cally fromthe lat est Frasnian I. iowaensis pop u la tion, and may have roots some what deeper in the Frasnian (Dzik 2002).The rep re sen ta tion of Icriodus through out the early and mid Famennian is not com pletely con tin u ous (theseshal low−wa ter cono donts were very sen si tive to lo cal en vi ron men tal changes) but it is al most cer tain that thechange from I. alternatus to I. cornutus was grad ual. Also the suc ces sion of Pelekysgnathus, which re placedIcriodus in times of its low fre quency and was ap par ently an open−sea an i mal, is of that kind. The low num berof spec i mens and their dis junc tive oc cur rence leaves some un cer tainty whether Dollymae? guizhouensis is inthe di rect an ces tor−de scen dant re la tion ship to older Pelekysgnathus or immigrated from another region.

De vo nian spathognathodontids. — Biramous sym met ri cal el e ments in the ap pa ra tus are an an cient fea −ture of the ozarkodinid cono donts. At the be gin ning of the Si lu rian in the prioniodinid lin eage, and at the be −gin ning of the De vo nian in polygnathids, a me dial pro cess de vel oped in the S0 el e ment to be pre served al −most with out any sig nif i cant change to the end of the Tri as sic. Re ver sal to the biramous con di tion was rec og −nized only in the Frasnian lin eage of the palmatolepidids (Dzik 2002). There fore it was puz zling thatthrough out the Car bon if er ous and Perm ian the lin eage of Synclydognathus and Hindeodus oc curred with anap pa ra tus groundplan closely sim i lar to that of the Si lu rian spathognathodontid Ozarkodina. It was pro posedby my self that they rep re sent the same clade (Dzik 1991) and this has found sup port in the sig nif i cant re duc −tion of the stratigraphi gap as a re sult of find ing Synclydognathus in the early Famennian of the Holy CrossMoun tains. This find ing dis closed also the af fin ity of Apatognathus, ear lier clas si fied among prioniodinids.There is a ten dency to wards re duc ing morphologic dif fer ences be tween el e ments in the ap pa ra tus in this lin −eage. Oc cur rences of the spathognathodontids re main few and stratigraphically isolated and it is thus difficult to resolve their phylogeny.

Prioniodinids and an ces tral gondolellids. — Among the prioniodinids only two lin eages seem to dis −play phyletic evo lu tion in situ, all oth ers show a dis junc tive, eco log i cally punc tu ated dis tri bu tion. The mostim pres sive case of evo lu tion ary trans for ma tion is of fered by Lagovidina. Its or i gin from ear lier Ligonodinare mains to be dem on strated, but is likely. Like other prioniodinids, these cono donts show a great vari abil itythat ham pers evo lu tion ary stud ies with low−num ber sam ples, but the lat est pop u la tions of the lin eage showrather stable organization of the apparatus.


The Famennian part of the lin eage of Pluckidina seems to lead to wards a gracile ap pear ance of el e mentswith a sharp, del i cate denticulation. Un for tu nately, only in a few sam ples are they rep re sented in rea son ablenum ber. The late Car bon if er ous to Tri as sic branch of the gondolellids (Or chard 2005) for a long time was

170 JERZY DZIK

Fig. 123. Strati graphic dis tri bu tion of the prioniodinids Pluckidina, Idioprioniodus and Uncadina in the Pol ish Famennian (same kind of pre sen ta tion as on Fig. 121).

con sid ered to be of cryp tic or i gin. The most char ac ter is tic as pect of their ap pa ra tus is the un usual shape of the S1 el e ment (“enantiognathus”), a re duced dor sal pro cess in the P1 el e ments, and ex tremely gracile ap pear −ance of other el e ments. These are fea tures of the lat est De vo nian mem bers of Branmehla. Its or i gin fromPluckidina is likely and it is only a mat ter of con ve nience where to place the boundary between theprioniodinids and gondolellids.

The most bi zarre lin eage of prob a ble prioniodinids is rep re sented by Uncadina of un known or i gin andgeo graphic source area. Some poorly known spe cies of Idioprioniodus show re mote sim i lar ity but the com −plete lack of any morphologic dif fer en ti a tion of el e ments in the ap pa ra tus was prob a bly an evolutionarynovelty.

Francodinids. — The most un ex pected re sult of this ap pa ra tus study is rec og ni tion of a whole branch ofcono donts with ap pa ra tus com po si tion rang ing from min ute forms of al most Si lu rian ap pear ance (Vogel −gnathus, ear lier known only from the Car bon if er ous) to ro bust ap pa ra tuses with el e ments mim ick ing the Car −


Fig. 124. Strati graphic dis tri bu tion of the ad vanced prioniodinid Guizhoudella and ear li est gondolellid Branmehla in the Pol ishFamennian (the same kind of pre sen ta tion as on Fig. 121).

bon if er ous prioniodinid Kladognathus (Planadina). They share a ten dency to paedomorphic shapes and areprob a bly monophyletic, al though the ear li est stages of their evo lu tion re main un known. All of sev eral lin −eages of the fam ily, ex cept for that of Vogelgnathus variabilis ® V. proclinatus, rep re sent brief in cur sions ofex otic, en vi ron men tally sen si tive forms. Mor pho log i cally the most in trigu ing is the lin eage of Urbaneko −dina, long−rang ing only at Miedzianka, with ex tremely small and sim pli fied el e ments. A fun da men tal re or ga −ni za tion of the ground plan of the M el e ment, oth er wise the most sta ble in the evo lu tion of cono donts, is of −fered by the Francodina ® Sweetodina ® Planadina succession, unfortunately punctuated by migrationsand allopatric speciation events.

Mehlinid or i gin of or nate plat forms. — An other un ex pected as pect of the evo lu tion of the Famennian cono donts is the prob a ble cen tral role of Mehlina in the evo lu tion of plat form−bear ing polygnathids. A key

172 JERZY DZIK

Fig. 125. Strati graphic dis tri bu tion of paedomorphic francodinids in the Pol ish Famennian (the same kind of pre sen ta tion as onFig. 121).

to un der stand ing their re la tion ship was of fered by the de vel op ment of a brush−like peniculus in the earlyhistogeny of the P2 el e ments. This char ac ter dis closes re la tion ships of sev eral lin eages of Polynodosus,ear lier placed in the an cient plat form−bear ing Polygnathus branch. It re mains to be re solved how far this


Fig. 126. Strati graphic dis tri bu tion of large francodinids in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

reached and whether the even more elab o rate Hemilistrona also be longs there. The end mem ber of thisbranch is of bi zarre mor phol ogy with al most com pletely re duced blades of both types of the P elements(Fig. 128).

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Fig. 127. Strati graphic dis tri bu tion of the non−plat form polygnathids and the least de rived plat form−bear ing Polynodosus spe cies in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

There is a hope that in crease in knowl edge of the ap pa ra tus struc ture will in tro duce a long−awaited or derin the phy log eny of or nate plat form−bear ing polygnathids. Stud ies on the evo lu tion of the Polygnathus−group cono donts are dif fi cult be cause they are highly sen si tive en vi ron men tally and have punc tu ated dis tri bu tionham per ing rec og ni tion of phyletic evo lu tion ary changes. More over, their pop u la tion vari abil ity is high. Un −for tu nately, ap pa ra tuses of the polygnathids are mor pho log i cally gen er al ized and usu ally only the P2 el e −ments are of sig nif i cance in di ag nos ing spe cies. How ever, a dis tinct ap pa ra tus com po si tion al lows sep a ra tion of the P. extralobatus–P. znepolensis group and proves that Polylophodonta (ac tu ally a rel a tive of Ancyro −gnathus) has lit tle to do with the Polynodosus confluens ® P. triphyllatus lineage.

Prob a ble ctenopolygnathid roots of siphonodellids. — There is much un cer tainty re gard ing re la tion −ships of the late Famennian and Tournaisian cono donts with a fusiform flat basal cav ity in the plat form el e −ments. They are be lieved to be of cru cial im por tance in de lim it ing, on an evo lu tion ary ba sis, the De vo −nian–Car bon if er ous bound ary. Un for tu nately, the ap pa ra tus re con struc tions of var i ous siphonodellids areno more than pro vi sional (Dzik 1997) and even re la tion ships within the clade (clas si fied in the Elicto −gnathidae, based on the as sumed con tro ver sial ap pa ra tus in ter pre ta tion) are not clear. It may be, how ever,of im por tance that the pre lim i nary ap pa ra tus re con struc tions of the ear li est siphonodellids (Dzik 1997)


Fig. 128. Strati graphic dis tri bu tion of the ad vanced Polynodosus spe cies and Hemilistrona in the Pol ish Famennian (the samekind of pre sen ta tion as on Fig. 121).

resemble the ap pa ra tus of the mid and late Famennian de riv a tive of the ro bust polygnathid Cteno poly −gnathus, here named Immognathus. In this lin eage the pe cu liar form of Siphonodella−like base de vel oped

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Fig. 129. Strati graphic dis tri bu tion of the ro bust polygnathids Ctenopolygnathus and Immognathus and pos si ble elictognathidspe cies in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121; draw ing of Polygnathus pennatulus of pos si ble re −

la tion ship to the Immognathus lin eage added).

in de pend ently of the Famennian Alternognathus. The lat ter ge nus, with its in cip i ent ax ial sym me try in theP1 pair, is an un likely relative of the siphonodellids, being rather ancestral to the Carboniferous cavus −gnathids.


Fig. 130. Strati graphic dis tri bu tion of the polygnathids Polygnathus and Neopolygnathus in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

The prob lem of Neopolygnathus. — The com mon early Car bon if er ous ge nus Neopolygnathus has De −vo nian roots. It is dif fi cult, how ever, to in di cate its ex act an ces try be cause its ear li est pop u la tions show agreat pop u la tion vari abil ity and the mor phol ogy of its el e ments is gen er ally de gen er a tive, marked by deepsim pli fi ca tion of the un known orig i nal mor phol ogy. Their P1 el e ments are of small size and sim ple mor phol −ogy but, par a dox i cally, a ten dency to wards sim pli fi ca tion and de vel op ing a smooth plat form (but with im −prints of ameloblasts) char ac ter izes one of the most ro bust polygnathids of the Famennian, P. semicostatus.No ta bly, un usu ally high pop u la tion vari abil ity is a fea ture of this spe cies. It seems thus that the char ac ter is ticcon cave base of P1 el e ments of Neopolygnathus is also a degenerative feature, an effect of loweredmineralization of the platform base.

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Fig. 131. Strati graphic dis tri bu tion of the polygnathid Lagovignathus and its pos si ble rel a tives among Polygnathus spe cies in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

Ad vanced Car bon if er ous spe cies of Neopolygnathus are al most homeomorphic to an other, un re latedpolygnathid lin eage, here clas si fied in Lagovignathus. Sev eral lin eages of these cono donts with a thin plat −form in the P1 el e ments oc cur in the mid Famennian, mostly of eco log i cally punc tu ated dis tri bu tion. Their or −i gin re mains un known but pos si bly the ear li est Famennian Polygnathus procerus may con nect them with theFrasnian spe cies char ac ter ized by an al most sym met ri cal, thin plat form. Only two lin eages show a prob a blyphyletic evo lu tion in situ. One of them L. fallax ® L.? dissimilis is in com pletely dem on strated. The other, L.lagoviensis ® L. granulosus ® L. styriacus, offers a succession of great correlative value.

Palmatolepidids. — Only one lo cal lin eage of Famennian palmatolepidids was rep re sented at the be gin −ning of the Famennian. The en vi ron men tal change that re sulted in the dis ap pear ance of the typ i cally Frasnian lin eages of Lagovilepis and Manticolepis had no de tect able in flu ence on the phyletic evo lu tion of theKlapperilepis lin eage ex cept for a some what de layed in crease in its pop u la tion vari abil ity (Dzik 2002). Thisseems to be a re sult of a re leased se lec tion pres sure ear lier gen er ated by com pe ti tion of other palmatolepididspe cies. New lin eages emerged se quen tially by im mi gra tion from their places of orig i na tion. The ap par entde crease in pop u la tion vari abil ity of K. ul tima was prob a bly caused by the ap pear ance in the area of spe cies


Fig. 132. Strati graphic dis tri bu tion of the ancyrognathids and their idiognathodontid homeomorphs in the Pol ish Famennian (thesame kind of pre sen ta tion as on Fig. 121).

that still had sim i lar eco log i cal pref er ences. Re mark ably, the plat form el e ments of the spe cies new in the area only a lit tle in creased the to tal range of shapes ear lier rep re sented in the sin gle spe cies, K. ul tima. How ever,un der the com pe ti tion from other sympatric spe cies they all had to re duce their vari abil ity. Prob a bly the lo calpop u la tion was forced to ac com mo date to the new con di tions of par tially over lap ping eco log i cal niches (thephe nom e non of char ac ter dis place ment). The sub se quent phy log eny of the Famennian palmatolepidids wasre viewed by my self else where (Dzik 2005). The early Famennian is marked by allopatric di ver si fi ca tion andthe sub se quent geo graphic merg ing of sev eral lin eages of Klapperilepis. They pre served an un de rivedapparatus structure but with various morphologies of the P1 elements.

In the mid Famennian they were re placed by a few lin eages of the grad u ally di ver si fy ing branch ofConditolepis, with sec ond arily biramous S0 el e ments of the ap pa ra tus. The branches of Palmatolepis andTripodellus con tinue through out the Famennian, usu ally with not more than two lin eages sympatric. Phyleticevo lu tion has been ob served in sev eral lin eages of the Famennian palmatolepidids. Among those of Klap −peri lepis the most ex ten sive is rep re sented by the lin eage K. schuelkei ® K. spathula ® K. quadrantino do −solobata. The suc ces sion K. wolskae ® K. circularis is also rather com plete but the ob served morphologicchange is not pro found. A sim i lar ex tent of trans for ma tions char ac ter izes the lin eage K. robusta ® K.crepida, the lin eage cov er ing early and late, rather dif fer ent K. ter mini, and the stratigraphically rather punc −tu ated lin eage K. protorhomboidea ® K. rhomboidea.

The main lin eage of Tripodellus of fers prob a bly the most ex ten sive morphologic change and com plete re −cord (Fig. 134). Ap par ently grad ual and con tin u ous is also the change in the long lin eage from Palmatolepisinitialis up to P. schindewolfi. Less con tin u ous but likely to be grad ual is the se ries P. ampla ® P. trachytera® P. rugosa. In Palmatolepis and Conditolepis the pro cesses of the M el e ments ac quired a po si tion closelysim i lar to that in the Frasnian Manticolepis. It ap pears thus that the pe lagic palmatolepidids, un like the gen er −ally shal low−wa ter polygnathids, of fer a much more com plete record of their evolution.

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Fig. 133. Strati graphic dis tri bu tion of ad vanced spe cies of the palmatolepidid Klapperilepis in the Pol ish Famennian (the samekind of pre sen ta tion as on Fig. 121).

Or i gin of ax ial sym me try of el e ments. — Left and right el e ments of many cono donts are not mir ror im −ages. This ap par ently im proved their fit in oc clu sion. In rare cases, how ever, this ten dency went as far as tode velop left and right el e ments of vir tu ally the same shape, seem ingly not form ing pairs. This is a case of ax −


Fig. 134. Strati graphic dis tri bu tion of ad vanced spe cies of the palmatolepidid Tripodellus and some Klapperilepis spe cies in thePol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

ial sym me try. Vir tu ally all the cono donts with the P1 el e ments of this kind oc cur in the late De vo nian andearly Car bon if er ous. One may thus won der whether the ax ial sym me try de vel oped once in the evo lu tion ofcono donts or sev eral times in de pend ently. The ear li est Famennian case of un doubted ax ial sym me try is rep −

182 JERZY DZIK

Fig. 135. Strati graphic dis tri bu tion of the palmatolepidid Palmatolepis and its an ces tral Klapperilepis spe cies in the Pol ishFamennian (the same kind of pre sen ta tion as on Fig. 121).

re sented by the mid Famennian Scaphignathus, a branch of the Alternognathus lin eage with an in cip i entasym me try of el e ments. There is no doubt that Scaphignathus gave rise to the Car bon if er ous branch ofCavusgnathus and Mestognathus, the model cases of axial symmetry.

There is an other group of cono donts with ax ial sym me try of their P1 el e ments. Their basal cav i ties are ba −si cally dif fer ent from that of Alternognathus, re sem bling rather Dasbergina in this re spect. How ever, the ex −act an ces try of this lin eage has not been traced and the sit u a tion re sem bles that among the Car bon if er ouscavusgnathids. The prob lem has to be clar i fied, which may not be so easy, how ever, be cause the Pseudo −polygnathus spe cies are en vi ron men tally sen si tive and their evo lu tion is poorly re corded. Per haps the mostcom plete suc ces sion is that of the P. ziegleri ® P. ultimus lineage.


Fig. 136. Strati graphic dis tri bu tion of the palmatolepidid Conditolepis in the Pol ish Famennian (the same kind of pre sen ta tion ason Fig. 121).

The De vo nian roots of the idiognathodontids. — A cru cial role in the evo lu tion of the late Pa leo zoiccono donts was played by a mid Famennian rel a tive of the most gen er al ized polygnathid lin eage of that age,Pandorinellina vulgaris. This was Dasbergina stabilis, the only de rived fea ture of which was the ex pandedbase with a shal low but wide cav ity. In sev eral lin eages that emerged from D. stabilis, the basal cone de vel −oped tuberculation or a kind of tuberculated plat form. Two rel a tively fast evolv ing lin eages are of im por tance in this group. The D. brevipinnata ® D. ziegleri lin eage de vel oped an elab o rate plat form with rows of tu ber −cles. In the D. kowalensis ® D. marburgensis ® D. trigonica lin eage, the orig i nally coarsely tuberculatedplat form grad u ally changed into a characteristic ramified structure.

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Fig. 137. Strati graphic dis tri bu tion of the cavusgnathids in the Pol ish Famennian (the same kind of pre sen ta tion as on Fig. 121).

Ap par ently from Dasbergina orig i nated also Protognathodus, which ap peared near the end of the De vo −nian from an un known source. Its lin eage prob a bly di ver si fied later into the main group of open−sea Car bon −if er ous cono donts, the idiognathodontids. An im por tant change in the ramiform el e ments of its ap pa ra tustook place in the Viséan (Dzik 1997).

Con clu sion. — The num ber of 142 cono dont spe cies says more about the no men cla tor ial his tory of theirre search than about their fau nal his tory. The count of iden ti fi able lin eages is some what more in for ma tive inthis re spect. Usu ally they rep re sent brief seg ments of evo lu tion la belled with a sin gle spe cies name, in someex am ples be ing a case of the more ex ten sive re cord of their evo lu tion, cor re spond ing to a few chronospecies.There are 101 such lin eages in the ma te rial avail able to me, 31 of them show ing more or less ap par ent evo lu −tion ary change. In the ma jor ity of cases the re cord is not comletely con tin u ous and it is not al ways clearwhether a lin eage com pletely dis ap peared from the area for some time or is not rep re sented in some sam plesonly because of its locally or periodically small contribution.

The di a grams de pict ing their time dis tri bu tion are not cal i brated in real time as the length of par tic u larunits ex presses only com plete ness of sam pled geo log i cal sec tions. Any way, if so crudely es ti mated rangesare com pared on the ba sis of their length, it ap pears that the suc ces sion show ing more or less cer tain evo lu −tion ary trans for ma tions rep re sent al most half (46%) of the sum ma rized ob served ranges of cono dont lin eages in the Holy Cross Moun tains and the Sudetes. Their re cord is not com plete and prob a bly not more than halfof it is suit able for stratophenetic stud ies. This is much less than in the tem per ate cli ma tic zone re cord rep re −


Fig. 138. Strati graphic dis tri bu tion of ad vanced spe cies of the idiognathodontids in the Pol ish Famennian (the same kind of pre −sen ta tion as on Fig. 121).

sented by the Or do vi cian suc ces sion in the same area (Dzik 1994), but this seems to be a dis tinc tion typ i cal of equatorial successions of various ages (Dzik 1983, 1997).

It is a con tro ver sial is sue how to es ti mate the com plex ity of liv ing forms but prob a bly in all as pects thatcan be con sid ered, the Famennian was the ep och of the great est an a tom i cal and spe cies di ver sity of cono −donts, com pa ra ble with only a few ep i sodes in their ear lier his tory (e.g., Llanvirn to early Caradoc; lateLlandovery to early Wen lock), and only one af ter the Famennian (late Tournaisian). The num ber ofsympatric spe cies (well de fined mor pho log i cal spe cies, not ar bi trarily se lected morphotypes!) was com −monly higher than 25. The Famennian ex pe ri enced also an in va sion of pe lagic en vi ron ments with ef fi cientlyswim ming large pred a tors. For the first time ac an tho dians, sharks, and ganoid fishes started to co−oc cur withcono donts (ear lier they in hab ited mostly near−shore en vi ron ments not ac ces si ble to cono donts; e.g., Blieck et al. 2000). The Famennian ma rine pe lagic fish show some spe cies di ver sity and sig nif i cant par tic i pa tion in bi −o log i cal pro duc tiv ity of the pe lagic com mu nity. Concavicarid and angustidontid crus ta ceans, with grasp ingap pend ages mim ick ing cono dont el e ments but an or der of mag ni tude larger, were also rep re sented byphosphatized man di bles and car a paces (see be low). Both fish and large pe lagic crus ta ceans prob a bly fed also on conodonts ( Rolfe and Dzik 2007). Perhaps this was a factor contributing to their late Palaeozoic decline.

AMMONOID CONCHS AND JAWS

The as pects of the ammonoid conch mor phol ogy eas i est to ac cess are the gen eral ge om e try and the con −tact of septa with the conch wall (su ture line), well dis cern ible on in ter nal moulds. Growth in cre ments on theconch sur face are less com monly pre served be cause the aragonitic conch wall is rarely trans formed into cal −cite with enough pre ci sion. Even less com monly pre served, but of much bi o log i cal and tax o nomic value, areini tial whorls of the conch. In the Late Palaeozoic goniatites and Me so zoic ammonites the first whorl of theconch has a smooth sur face de void any marks of growth in cre ments Kulicki et al. 2002). This is gen er ally ac −cepted as the ev i dence of its for ma tion within the egg cov ers and the ammonitella de vel op men tal stage pre −ced ing the pri mary shell con stric tion is con sid ered em bry onic. How ever, in the orthoceratid nautiloids (Dzik1981, 1885), bactritids (Doguzhaeva 2002), and ear li est ammmonoids (e.g., Klofak et al. 1999) the conch de −vel op ment stage cor re spond ing to ammonitella bears dis tinct growth in cre ments. Ap par ently ju ve nile earlycepha lo pods were free liv ing. The tran si tion from free−liv ing larva to de vel op ment of the larva in side the eggun til its meta mor pho sis was prob a bly grad ual and pro ceeded in de pend ently in many lin eages, like othermolluscs. In some goniatites with a smooth ammonitella conch, adult mor pho log i cal fea tures de velop sud −denly, show ing that the change was con nected with a meta mor pho sis of anat omy, not just a re tar da tion ofgrowth re sult ing from hatch ing. There fore I pre fer to ap ply the term “lar val” to conchs at the pre−meta mor −pho sis stage of the de vel op ment, equipped with two first septa. In the ma te rial from the Holy Cross Moun −tains a few phosphatized larval conchs have been found, the larval stage is also discernible in pyritic internalmoulds found in conodont samples.

Lit tle data are avail able on the anat omy of the De vo nian ammonoids. Rarely dis cern ible mus cle scar at −tach ments are known (Rich ter 2001, 2002; Rich ter and Fischer 2002). Only in a few cases the conchs can bematched with jaws.

ANAPTYCHI AND APTYCHI

The ammonoid jaws func tioned si mul ta neously also as opercula (Dzik 1981). To be pre served, the mostly or ganic jaws of the De vo nian ammonoids re quired anoxic con di tions of sed i men ta tion, rep re sented by theblack shale fa cies. The only ho ri zon in the Holy Cross Moun tains where ammonoid jaws oc cur in sig nif i cantnum ber is the cal car e ous shale at Kowala of ear li est Famennian (pos si bly also latest Frasnian) age.

Un for tu nately, no de ter mi na ble cephalopod conch has been found in as so ci a tion with them, so it can onlybe spec u lated, to which ammonoid taxa they be longed. A few sin gle−piece lower jaws (anaptychi) do not dif −fer in out line from those iden ti fied in the liv ing cham ber of the gephyroceratid Crickites by Matern (1931) orclos ing ap er ture in Sphaeromanticoceras (Clausen 1969). Al though the spec i mens were col lected from thescree, it is likely that they come from strata of lat est Frasnian age. At Kowala, the cephalopod lower jaws split

186 JERZY DZIK

me di ally into two valves (aptychi) are much more com mon. Their oc cur rence in the De vo nian is sur pris ing,as true aptychi are con fined to the late Me so zoic, ex cept for those of the Si lu rian arionoceratid nautiloids.The pos si bil ity that the fos sils from Kowala rep re sent bactritoid or orthoceratid jaws can be rather ex cludedbe cause of the size of the larg est spec i mens, many times ex ceed ing that of the late De vo nian bactritids (Fig.139B). Their as so ci a tion with the tornoceratids, the only ammonoids known from strata of this age in Eu rope, is more plau si ble. The lower jaw of the early Car bon if er ous goniatitid Girtyoceras (Doguzhaeva et al. 1997),as well as its late Car bon if er ous rel a tive Eoasianites (Closs 1967), sig nif i cantly dif fer in its shape, be ing sim −i lar rather to the jaws of the nautilids. Whether there were sev eral re ver sals in the evo lu tion of ammonoidbeaks or rather the goniatitids rep re sent a side branch of the main ammonoid lin eage, re mains to be clar i fied.The pen tag o nal shape of the prob a ble anaptychus Libodiscus from the early Tournaisian Exshaw Shale of Al −berta (Dzik 1997, p. 126) sup port the idea of con ti nu ity be tween the Frasnian and post−Car bon if er ousammonoid jaws with opercular func tion. Anaptychi of a some what sim i lar pen tag o nal shape have been de −scribed from the early Clymenia Stufe of the Rhenish Slate Moun tains by Korn (2004b) and the ClevelandShale of Ohio (Frye and Feldmann 1991).


Fig. 139. Prob a ble ammonoid jaws from the Late De vo nian at Kowala in the Holy Cross Moun tains. A. Hy po thet i cal me dial sec −tion of an ad vanced tornoceratid show ing po si tion of the jaws. B. Large aptychus with sep a rated valves from the same stra tum,spec i men ZPAL AmVII/366. B–F. Ju ve nile anaptychi pos si bly be long ing to Tornoceras, from the ear li est Famennian with theor ganic chitinous tis sue car bon ized or pyritized (C), spec i mens ZPAL AmVII/370, 371, 374, and 373. G. Car bon ized

anaptychus, pos si bly from the lat est Frasnian and be long ing to a gephyroceratid, spec i men ZPAL AmVII/367.

The names in tro duced for ammonoid jaws are valid from the no men cla tor ial point of view, and might po −ten tially com pete for pri or ity with names based on conchs. This ap proach has been ex e cuted al ready byMatern (1931), who iden ti fied the jaw (anaptychus) Spathiocaris koeneni Clarke, 1884 in the liv ing cham berof Crickites holzapfeli Wedekind, 1913. The type spec i mens of both taxa are from the same ho ri zon and re −gion, so their bi o log i cal iden tity raises lit tle doubt. This is not so ap par ent in the case of the type spe cies ofSpathiocaris, S. emersonii Clarke, 1882 from the Frasnian Por tage For ma tion at Na ples, On tario (Clarke1882). Po ten tially, Spathiocaris may ap pear to be a se nior syn onym of, say, Manticoceras Hyatt, 1884, but itwould not be prac ti cal to en force this no men cla tor ial ac tion be cause of rather in dif fer ent mor phol ogy of theDe vo nian ammonoid man di bles. For tu nately, no such prob lem is connected with the Famennian ammonoidsdiscussed below.

GROWTH AND FUNCTION OF THE CONCH

It is gen er ally be lieved that the cephalopod sep tum ge om e try is of func tional im por tance, es pe cially when it shows con vex ar eas act ing as vault ing in the gen er ally con cave sep tum (when seen from the conch ap er −ture). This is why in the ammonoid and nautiloid tax on omy much value is given to the course of the su tureline – a pla nar rep re sen ta tion of the con tact be tween sep tum and the conch wall. How ever, the cor re spon −dence be tween the su ture line and ge om e try of the sep tum is not straight for ward and sim i lar su tures may ex −press dif fer ent pat terns of con vex i ties and con cav i ties of the sep tum. In the sim plest sit u a tion, the vis ceral sac of an ammonoid body be haves as a bal loon un der pres sure of in ter nal flu ids and its end de vel ops a hemi −spher i cal shape in a cy lin dri cal body cham ber of the conch (Seilacher 1975). If the cross sec tion of the cham −ber, de ter mined by the se cre tive ac tion of the man tle mar gin at its ap er ture, is lat er ally flat tened, wide lat eralsi nuses may de velop in the su ture line. More com plex out lines may pro duce a quite so phis ti cated pat tern oflobes and sad dles with out any change in the con cav ity of the sep tum (Seilacher 1975; Dzik 1984; Korn 1992, 1999; Dzik and Gaździcki 2002). The sit u a tion be comes more com plex if the body is lo cally at tached to theshell wall with mus cles, which ham per its fast with drawal. Lobes not re lated to the conch cross−sec tion maythen de velop, fre quently pointed, and mar ginal parts of the sep tum may fi nally de velop vault ing (Seilacher1975). Yet an other mech a nism of vault ing and com plex septal ge om e try may be re lated to non−uni form stiff −ness of the free sur face of the vis ceral sac. This could have been caused by the translocation of some internalorgans towards the venter as a result of increased involuteness of the conch.

The conch involuteness and the whorl ex pan sion rate are in de pend ent as pects of the conch ge om e try. In −vo lute and evolute conchs may have the same whorl ex pan sion rate, and vice versa. The whorl ex pan sion rate var ied very much among the early ammonoids be ing clearly con nected with their swim ming abil i ties (Kluget al. 2004). The Agoniatitina, Gephuroceratina, Prolecanitida and Paraceltitina gen er ally show a high whorlex pan sion rate and sig nif i cant con trol of on tog eny, while Anarcestina, Pharciceratina, Tornoceratina, andGoniatitina show a low ex pan sion rate (Korn and Klug 2001). This is not an evo lu tion ary sta ble char ac ter and within the Tornoceratina there was a change from the tornoceratids with rel a tively high whorl ex pan sion rateto cheiloceratids with low ex pan sion rate, the char ac ter sub se quently in her ited by the Goniatitina, butchanged again dur ing the evo lu tion ary history of the Prolecanitina (Korn 1994).

Wrin kle layer. — The wrin kle layer of the ammonoid shell wall (House 1971) was se creted by the dor salsur face of the an i mal head while it ex tended from the liv ing cham ber. This is quite clearly shown, among oth −ers, by the spec i mens of Tornoceras typum from Jabłonna with pre served in tact ap er ture (Fig. 141A3). The ex −tent of the wrin kle layer is there clearly de lim ited. It formed a lobe out side the ap er ture. The area was in con tactwith the body only when with drawn from the conch. The se cret ing or gan was thus rather the an i mal’s head than a spe cial ized part of the man tle. This sug gests that the wrin kle layer is ho mol o gous to the “black layer” of thenautilids and ceratites, oc cur ring in ex actly the same re gion near the conch ap er ture (Klug et al. 2004).

Korn (2000) com pared ir reg u lar me an der ing pro tru sions on the shell sur face of some Famennian am −monoids with the wrin kle layer, us ing this as the ev i dence of their conchs be ing cov ered with man tle dur inglife. These struc tures re sem ble min er al ized periostracum pro tru sions, the fea ture of conchs of many mol −lusks, in clud ing the Mesozoic ammonites.

In ter nal thick en ings and con stric tions of the conch. — In ter nal thick en ings are spe cific for manyFamennian cheiloceratids and a high di ag nos tic value was at trib uted to them by Sobolew (1914). It is thus

188 JERZY DZIK

im por tant to un der stand their or i gin be fore their use in tax on omy is sup ported or aban doned. As shown bysome ma ture spec i mens of Cheiloceras subpartitum from Łagów, which died im me di ately af ter de vel op ingthe last in ter nal thick en ing of the shell, the thick en ing was pro duced at the mar gin of the ap er ture or im me di −ately be hind it. As a re sult, in un de rived cheiloceratids the thick en ings tend to fol low the growth in cre ments.In Prionoceras, there is even a con nec tion be tween ir reg u lar i ties at the conch sur face and the thick en ing be −low. In more de rived cheiloceratids such a cor re spon dence has dis ap peared and in their conchs the ap er turalsi nus may co−oc cur with con stric tions form ing a prom i nent ven tral sad dle (e.g., Cheiloceras inversum). Ap −par ently, the place of for ma tion of the thick en ings moved be hind ap er ture, al though most prob a bly its mostdis tal part was still ini ti ated at, or near, the ap er ture. This is sug gested by the unity of ex ter nal and in ter nalshell wall fea tures in Balvia. The mech a nism con trol ling de vel op ment of such con stric tions re mains un −known. It could have been sim i lar to that re spon si ble for the divaricate shell or na men ta tion in other mol lusks, that is the zone of for ma tion of the con stric tion mi grated along the ap er ture mar gin with its growth (this doesnot seem es pe cially likely) or a lin ear nar row zone of increased secretion of the lamellar aragonite layer of the conch extended across the mantle, initially in connection with its margin.

Terminalized conch growth. — It is gen er ally as sumed that a crowd ing of septa and growth in cre mentsnear the ap er ture is suf fi cient to prove ma tu rity of the ammonoid conch (e.g., Makowski 1991). Ac tu ally,these fea tures of the conch are just re flec tions of de layed growth, which may be ac ci den tal, sea sonal or con −nected with spawn ing, and may thus be mis lead ing. Re gret ta bly, an al ter na tive to this po ten tially sim plis ticrea son ing is of fered in the Famennian only by the conch ge om e try of the prolobitid goniatites and wocklu −meriid clymenias. In both groups the ma ture body cham ber shows a pro found change in its cross−sec tion andde vel ops periapertural mod i fi ca tions. Un like the post−Tri as sic ammonites, this does not seem to be an ex −pres sion of sex ual di mor phism. The body cham ber mod i fi ca tion of Prolobites, with a con stric tion near theap er ture strength ened with an in ter nal thick en ing, seems to be a pro tec tion agains pred a tors. The ma tureconch shape of the wocklumeriids probably improved their hydrodynamic properties.

DESCRIPTION AND CLASSIFICATION OF THE AMMONOIDS

In this work a re view of the ammonoid fau nal dy nam ics in the Famennian of the Holy Cross Moun tainsand Sudetes is pre sented rather than a for mal de scrip tion of the taxa. A re li able tax o nomic re search, as ex em −pli fied by works of Korn (e.g. 1994) and Becker (e.g., 1993b), would re quire ac cess to the mostly Ger mantype and topotype ma te rial. In the case of the taxa based on ma te ri als from Po land, very few type spec i menssur vived the World War II and these were al ready reillustrated by ear lier au thors. The types of Sobolew(1914) and Dybczyński (1913) have not been traced in known mu seum col lec tions and are prob a bly all lost.Their type strata are no lon ger ac ces si ble (with a few ex cep tions which were at tempted by my self to ex ploit).In the for mer Dzieduszycki’s Mu seum col lec tion in Lvov there are many spec i mens col lected by Dybczyński at Sieklucki’ brickpit (lo cated in the west ern part of Kielce, no lon ger ac ces si ble). The pyritized ammonoidspec i mens oc curred there, to gether with nu mer ous con cre tions, in the Qua ter nary sand cov er ing the Famen −nian shale ex ca vated in the pit (Sobolew 1912a). They were washed out from var i ous strati graphic units ofthe Famennian (pos si bly even Frasnian, if Beloceras acutodorsatum Dybczyński, 1913 truly rep re sents thege nus) in the Pleis to cene and are ob vi ously heterogenous from the strati graphic point of view. It is po ten −tially pos si ble to se lect neotypes from this ma te rial but these would be of lit tle use, as they in fact lack anystrati graphic in for ma tion. For tu nately, a closely sim i lar ma te rial has been re cov ered from black shale withinthe C. marginifera Zone at Kowala, which suggests that most (but not all) of the Sieklucki’s brickpit materialwas derived from this horizon.

Most of the ev i dence pre sented be low is de rived from mac ro scopic in ter nal moulds of the ammonoidconchs with frag ments of the orig i nal shell pre served. Cam era lucida draw ings of the su ture and growth lines,to gether with pho to graphs of spec i mens whit ened with am mo nia chlo ride il lus trate tax o nomic de scrip tions be −low. When ever there is enough ev i dence, the ge om e try of the sep tum is reconstructed in a diagrammatic way.

Tax o nomic no men cla ture of the Famennian ammonoids. — The type spe cies of the three rep re sen ta tivegen era of Pa leo zoic ammonoids, Cheiloceras, Prionoceras, and Imitoceras all have obliquely con vex pro file of


the conch ap er ture, as well as the same or closely sim i lar shape of the conch and the ex ter nal part of the su ture.They dif fer in a de gree of fold ing of the dor sal part of the sep tum and in the dis tri bu tion or lack of the in ter nalconch thick en ings, fea tures do not con sid ered to be di ag nos tic for high rank ammonoid taxa and dis tin guish ingrather spe cies within par tic u lar gen era. Yet, these are types of their own superfamilies! This tra di tional ap −proach is an ex pres sion of a gen eral trend in the evo lu tion of tax on omy to wards ex treme split ting of taxa andris ing their ranks. As a re sult of pro fes sional spe cial iza tion and low num ber of tax on o mists in par tic u lar ar eas of re search (in the case of De vo nian ammonoids rarely ex ceed ing a few stu dents in volved in re search at the sametime) there is lit tle in ter est in stan dard iz ing taxa. The dif fer ence be tween tra di tional tax on omy of the Famen −nian tornoceratine ammonoids (sev eral superfamilies) and that of the co eval palmato lepidid cono donts (singlegenus), both including a similar numbers of sympatric species, exemplifies this very well.

These dif fer ences in clas si fi ca tion meth od ol o gies would be prob a bly of lit tle im por tance if not the valuethat is given to count ing taxa in hope to dis close pat terns in the evo lu tion of the liv ing world. More over, thetoo deep dis crep ancy in at ti tudes makes the Lin nean tax on omy meth od olog i cally weak and prone to de struc −tion by fol low ers of more rad i cal phi los o phies of tax on omy. I be lieve that to make tax on omy truly use ful inits main func tion as a mea sure of di ver sity (even if it is un avoid ably sub jec tive) some kind of stan dard iza tionis nec es sary. This can be achieved by keep ing in mind that too many taxa within a taxon of higher rank makeclas si fi ca tion im prac ti cal, dif fi cult to mem o rize and to use. Ap prox i mately sim i lar range of morphologicvari abil ity should also be al lowed within taxa of the same rank. Ob vi ously, this can not be done too rig or −ously. I be lieve that a con trolled sub jec tiv ity is still a much better so lu tion than to go into the illusion thattaxonomy can be done in a strict and completely objective way (Dzik 2005).

Nomenclatorially in valid “ge neric” names of Sobolew (1914), cited in the text, are not ital i cized.

Phylum MOLLUSCA Linné, 1758Class CEPHALOPODA Cuvier, 1795Subclass AMMONOIDEA Zittel, 1884

Order GONIATITIDA Hyatt, 1884Suborder TORNOCERATINA Wedekind, 1918

Re marks. — The tornoceratine goniatites in her ited a low conch ex pan sion rate af ter their anarcestine an −ces tors (Korn and Klug 2001). The conchs are gen er ally in vo lute with the area of the sep tum on both sides ofthe pre ced ing whorl vaulted in the dorso−ven tral plane.

Much value is given to the pro file of ap er ture in the tornoceratine goniatites and an elab o rate ter mi nol ogyfor var i ous shapes of growth lines and ribs is avail able. To make the text com pre hen si ble also to non−spe cial −ists I avoid such terms and re fer only to prom i nence of ventrolateral pro tru sions of the ap er ture (au ri cles) anddepth of the ven tral infundibular si nus. Sche matic draw ings of growth lines of fer enough in for ma tion on their ex act course. Also while de scrib ing su tures pos si bly neu tral descriptives are used in hope that this will be un −der stand able to followers of various notation systems.

Family Tornoceratidae von Arthaber, 1911Di ag no sis. — Conch ap er ture with au ri cles and rel a tively high whorl ex pan sion rate, sep tum of sim ple

ge om e try de vel op ing only small lat eral vault ings (ex pressed as one set of flank lobes in the su ture) with aten dency to sim pli fi ca tion in secondarily evolute forms.

Re marks. — The main dif fi culty with the tax on omy of gen er al ized tornoceratids is the sim plic ity of their ex ter nal conch mor phol ogy and ge om e try of the sep tum. To in ter pret the lat ter it seems enough to in voke the“bal loon con cept” of the cephalopod sep tum as an in ter play of the whorl cross−sec tion and uni formly dis trib −uted hy drau lic pres sure within the an i mal soft body (Seilacher 1975). Only in more ad vanced tornoceratidsvaulted ar eas in the sep tum de vel oped, sug ges tive of a more com plex pat tern of mus cu lar at tach ments of thebody to the in ter nal conch wall. The septal and conch mor phol ogy of early tornoceratids seems to be sen si −tive of evo lu tion ary re ver sals and homeomorphy an thus of low tax o nomic value. In fact, the conch shapeand the su ture line con trolled by it seem vari able in both the early Frasnian (Bogoslowsky 1971) and lat estFrasnian (Dzik 2002) species represented by a large number of specimens.

190 JERZY DZIK

In such a sit u a tion the mor phol ogy of the first lar val sep tum, deeply con cave in at least some tornoceratids (Ruzhentcev 1962; Bogoslovsky 1969) may be of much tax o nomic value, be cause of its unique ness. The first sep tum is very dif fer ent from the suc ceed ing septa and ap par ently orig i nated un der dif fer ent con di tions of se −cre tion. One may spec u late that this was a re sult of se cre tion of the phragmocone cham ber liq uid be ing de −layed in re spect to se cre tion of the septal tis sue. The vol ume of the first reg u lar phragmocone cham ber wasmuch smaller than that of the protoconch, so sub se quent septa orig i nated in a more reg u lar rhyth mic pat tern.Any way, this pe cu liar mor phol ogy of the lar val (ammonitella) conch may be a use ful di ag nos tic char ac ter(synapomorphy) for the least derived trornoceratids.

The prob lem thus emerges when this con cave first sep tum orig i nated and how long it per sisted in the evo −lu tion of the tornoceratids. This char ac ter was first iden ti fied in a typ i cal Tornoceras spe cies from the earlyFrasnian Domanik For ma tion of Timan (Ruzhentcev 1962; Bogoslovsky 1969). House (1965), while in ter −pret ing pyritized protoconchs of the type spe cies of the ge nus from the Givetian Alden Marcasite ho ri zonwithin the Ledyard Shale of New York, sug gested the pres ence of an un usu ally large caecum. It seems pos si −ble that this was ac tu ally the first sep tum which de vel oped such a swol len struc ture. If true, this char ac termight de fine the or i gin of the tornoceratid clade more precisely than the conch morphology.

This bi zarre first sep tum mor phol ogy is shown by si lici fied tornoceratid spec i mens from the top mostFrasnian strata at Miedzianka in the Holy Cross Moun tains (Fig. 140) as so ci ated with a nor mally de vel oped(not swol len) siphuncle. Among postlarval tornoceratids of gen er al ized conch mor phol ogy only Lingua −tornoceras has been iden ti fied in these strata (Dzik 2002). A sim i lar, but not so strongly con cave first sep tumchar ac ter izes a tornoceratid from the sig nif i cantly youn ger Up per Łagów Beds at Dule (sam ple Ł−38). As so −ci ated protornoceratids show nor mal de vel op ment of the first sep tum. This sug gests a grad ual dis ap pear anceof this char ac ter and ex plains why the clymeniids do not show any signs of its pres ence, despite theirprobable tornoceratid ancestry.

Genus Tornoceras Hyatt, 1884

Type spe cies: Goniatites uniangularis Conrad, 1842 from the lat est Givetian Leicester Py rite of New York.

Di ag no sis. — In vo lute discoidal conch with closed um bi li cus; the main subventral area of the sep tumcon cave, with out lat eral vault ings; the first sep tum deeply con cave and swollen.


Fig. 140. Si lici fied ju ve nile ammonoid conchs from the lat est Frasnian at Miedzianka, sam ples Md−18 (A) and Md−21 (B–E) inthe Holy Cross Moun tains. A. Al most me di ally split protoconch of a tornoceratid (prob a bly Linguatornoceras, the only ge nuscon vinc ingly iden ti fied from strata of that age in the area; Dzik 2002) show ing swol len first cham ber, parts of spec i men ZPALAmVII/1808 in me dial and septal views. B. Sim i larly pre served spec i men ZPAL AmVII/1810. C. First sep tum of spec i menZPAL AmVII/1811. D. Spec i men pre served as a par tial in ter nal mould of the first cham ber, ZPAL AmVII/1812. E. Protoconch

of a gephyroceratid in lat eral view and its ven tral sur face with re mains of septa on the sec ond coil, ZPAL AmVII/1813.

192 JERZY DZIK

Fig. 141. The most gen er al ized tornoceratid Tornoceras typum (Sandberger et Sandberger, 1851) from the early Famennian K.crepida Zone at Jabłonna (A, B, D, F), Kadzielnia (C, G), and Wietrznia (E) in the Holy Cross Moun tains. Growth lines andviews of the conch with pre served shell and com plete ap er ture, mag ni fied to show the ex tend of wrin kle layer (A, bed 7), spec i −men ZPAL AmVII/299; su ture (B) of AmVII/259 (ex ter nal part; bed 7) and 301 (in ter nal part; bed 6); septal ge om e try (C) basedon spec i mens AmVII/266 (bed 8) and 301 (bed 6); ma ture male? conch with ex fo li ated shell (D, bed 5) ZPAL AmVII/262; in ter −nal mould of subadult conch (E, beds with Concavicaris) ZPAL AmVII/1692; ma ture fe male? conch (F, bed 9) ZPAL

AmVII/364; pos si ble macroconch phragmocone(G) with coarsely recrystallized shell ZPAL AmVII/458.

Re marks. — The Frasnian pop u la tions of the main lin eage of Tornoceras dif fer from those of theGivetian in a less acute out line of the ven tral sad dles (Bogoslovsky 1971) and from those from the earlyFamennian in su ture lobes re stricted to the subventral part of the whorl.

The sep tum ge om e try of Tornoceras is ex tremely sim ple, sim pler than one may in fer from the rel a tivelycom plex course of the su ture line. There are only con cav i ties in the sep tum that fits closely the baloon modelof Seilacher (1975). The dif fer ence be tween the cen tral con cav ity lo cated ven trally of the pre ced ing whorland con cav i ties on its side ap par ently re sulted from dif fer ent ease of with drawal of the body while de vel op −ing a new cham ber. This may have been re lated to the an i mal’s anat omy. One may guess that the in ter nal or −gans of higher stiff ness (liver?) oc cu pied the ven tral part of the whorl, re sist ing the cameral liquid pressuremore than the dorsolateral parts.

Tornoceras typum (Sandberger et Sandberger, 1851)(Figs 139B–E?, 141, 142A, and 159)

Type ho ri zon and lo cal ity: Prob a bly Frasnian at Seßacker in the Dillenburg re gion in the Rhenish Slate Moun tains (Becker1993b).

Ma te rial. — 23 spec i mens.Di ag no sis. — Smooth, in vo lute, discoidal conch, deep gently con cave flank lobe of the su ture line.Re marks. — This spe cies is usu ally re ferred to as T. sim plex (von Buch, 1832) the orig i nal draw ing of

the type of which shows a spec i men with open um bi li cus and is a nomen dubium (Becker 1993a, p. 182). T.incrassatus (Gürich, 1896), based on a spec i men from the early Famennian of Karczówka is prob a bly avail −able as a valid name. Al though the de pos i tory of the orig i nal spec i men re mains un known, pieces of the rockwith ammonoid conchs are pre served in the col lec tion of Dymitr Sobolew in Kharkov, Ukraine.

Ac cord ing to Becker (1993a) this mostly early Famennian spe cies is close to the Frasnian Lingua −tornoceras linguum (Sandberger et Sandberger, 1851), the only tornoceratid pre ced ing it stratigraphically inthe Holy Cross Moun tains (Dzik 2002). Makowski (1991; fol lowed by Becker 1993b) clas si fied the earlyFrasnian tornoceratids from the Holy Cross Moun tains in T. frechi (Wedekind, 1918) pro posed to be the typespe cies of the new ge nus Phoenixites by Becker (1993b). Un for tu nately, sim i larly as in the case of T. typum,the type ho ri zon of P. frechi is un cer tain. Orig i nally the spe cies was based on an early or mid Frasnian ma te −rial (Wedekind 1918) and Becker (1993a) pro posed the neotype from Seßacker near Oberscheld, prob a blyalso of Frasnian age. This would re quire that these two closely sim i lar tornoceratid lin eages co−oc curred inboth the Frasnian and Famennian, cross ing the bound ary be tween these ages. The di ag nos tic fea ture ofPhoenixites is the pres ence of ventrolateral fur rows at the ju ve nile growth stages. Noth ing like that has beenrec og nized in the spec i mens avail able to me, their smallest diameter being 5 mm.

There is a sig nif i cant vari a tion in the conch ge om e try and su ture line within par tic u lar sam ples. In somespec i mens the flank lobe is strongly con cave (as typ i cal for the spe cies), in oth ers shal low, al most ap proach −ing the sta tus typ i cal of the mid Famennian T. crebriseptum. Ju ve nile spec i mens may have rather flat or, in −stead, globose conchs. Se rial ven tral thick en ings of the ap er ture, con sid ered by Becker 1993b to de fine his


DIAMETER (mm)DIAMETER (mm)DIAMETER (mm) DIAMETER (mm)DIAMETER (mm)DIAMETER (mm)

Fig. 142. Vari abil ity of conch di men sions of Tornoceras A. T. typum (Sandberger et Sandberger, 1851) from the earlyFamennian K. crepida Zone at Jabłonna (beds 5, 6, and 7). B. T. crebriseptum Ray mond, 1909 from the mid Famennian C.

marginifera Zone at Łagów (sam ples Ł−5, 28, 29, 30, 31, and 33, prob a bly all from the same bed).

Phoenixites sulcatus and P. varicatus (Wedekind, 1918), co−oc cur with those with out thickenings (inJabłonna beds 7 and 24).

The tornoceratids show an ap par ent bimodality in size fre quency dis tri bu tion in ter preted as a case of sex ualdi mor phism (Makowski 1962, 1963, 1991). Al ready in the early Frasnian Domanik For ma tion of the Timanspec i mens of di am e ter about 35 mm with densely dis trib uted ter mi nal septa co−oc cur with conchs of at lest 90mm di am e ter (Bogoslowsky 1971). In my ma te rial from Jabłonna, a still not ma ture spec i men of 115 mm conch di am e ter co−oc curs with ma ture spec i mens of 27 mm size. Makowski (1991) pro posed to dis tin guish a sep a ratechronosubspecies T. frechi parvum based on the al leg edly small size of macroconchs but the il lus trated spec i −mens do not show con vinc ing ev i dence of growth termination and may be actually juveniles.

In the lam i nated marls of the ear li est Famennian at Kowala, nu mer ous cephalopod jaws oc cur. Their elon −gated shape sug gests that the conch ap er ture to which they were fit was rather com pressed, with a roundedventer. This would fit the mor phol ogy of Tornoceras, the only lin eage ex pected to oc cur in strata of this age.The most un ex pected as pect of these jaws is that they show a me dial su ture; these were ac tu ally aptychi (Fig.139B–E). Min ute spec i mens, prob a bly ju ve nile, show some times an iron min er al iza tion sug ges tive of sub −stan tial orig i nal thick ness of valves in their mar ginal parts. Large, prob a bly ma ture spec i men is split alongthe su ture and the ven tral tips of valves are rounded also admedially.

Dis tri bu tion. — Prob a bly the late K. triangularis Zone at Karczówka, the early K. crepida Zone atWietrznia, Kadzielnia, Jabłonna (beds 5–7), and Janczyce (Makowski 1991).

Tornoceras subacutum Makowski, 1991(Figs 143A–H and 159)

Type ho ri zon and lo cal ity: Bed 3 of the early Famennian at Janczyce, Holy Cross Moun tains.

Ma te rial. — 15 spec i mens.Di ag no sis. — Macroconchs with nar rowly trap e zoidal cross sec tion of the venter; microconchs with

rounded venter (Makowski 1991).Re marks. — The evo lu tion ary po si tion of the spe cies within the con tin u ous phyletic lin eage run ning

from T. typum to T. sublentiforme was pre sented by Makowski (1991). Su tures in three spec i mens (di am e ters 26 to about 30 mm) show a ter mi nal con den sa tion. The pyritized min ute spec i men of Gomi−monomeroceras(Tornoceras) kielcense Sobolew, 1914, from Sieklucki’s brickpit may be long to this spe cies al though its su −ture shows a rather un usual sym met ri cal shape of the flank lobe.

Dis tri bu tion. — The K. crepida Zone at Janczyce (bed 3 of Makowski 1991) and Kadzielnia.

Tornoceras sublentiforme (Sobolew, 1914)(Figs 143I–N and 159)

Type ho ri zon and lo cal ity: Re worked to Qua ter nary sed i ments at Sieklucki’s brickpit in Kielce, Holy Cross Moun tains.

Ma te rial. — Eight spec i mens.Di ag no sis. — Acute or very nar rowly tab u late venter of adult spec i mens; gen er al ized tornoceratid suture.Re marks. — This is the end mem ber of the lin eage of Tornoceras subacutum as shown by Makowski

(1991). Rel a tively large spec i men MD 11090 from Kadzielnia in the col lec tion of the for mer Dzieduszyckis’Mu seum in Lvov, about 70 mm in di am e ter, and ma ture spec i men ZPAL AmVII/991 in my col lec tion, of di −am e ter 57 mm, have an al most acute venter with a very nar row tab u la tion. The com plete lin eage seems to berep re sented at Kadzielnia, but the ma te rial co mes from loose blocks and the ex act strati graphic po si tion ofpar tic u lar specimens is hard to determine.

Dis tri bu tion. — The K. crepida Zone at Janczyce (beds 4–5 of Makowski 1991), Jabłonna (beds 10, 11,14), and Kadzielnia; re worked at Sieklucki’s brickpit in Kielce.

Tornoceras crebriseptum Raymond, 1909(Figs 142B, 144A–D and 159)

Type ho ri zon and lo cal ity: Mid Famennian (prob a bly Platyclymenia annulata Zone) Three Forks Shale north of ThreeForks, Montana (D. Korn, per sonal com mu ni ca tion 2005).

Ma te rial. — 33 spec i mens.Di ag no sis. — Rel a tively shal low flank lobe of the su ture.

194 JERZY DZIK


Fig. 143. Tornoceratids with acute conch venter from the early Famennian K. crepida Zone of the Holy Cross Moun tains.A–H. Torno ceras subacutum Makowski, 1991 from Kadzielnia; growth lines based on unnambered prob a ble macroconch fromMakowski’s col lec tion (A) and microconch ZPAL AmVII/287 (B), su ture line based on spec i men ZPAL AmVII/286 (C); prob a −ble microconchs ZPAL AmVII/990, 999, and 987 (D–F) and macroconchs ZPAL AmVII/991 (G) and MD 11090 (H).I–N. Torno ceras sublentiforme (Sobolew, 1914) from Janczyce (I, bed 5 of Makowski 1991) and Jabłonna (J–N); growth linesbased on unnambered prob a ble macroconch from Makowski’s col lec tion (I) and microconch ZPAL AmVII/473 (J, bed 11; alsoviews of the spec i men), su ture line based on spec i men ZPAL AmVII/477 (K, bed 11); ju ve nile ZPAL AmVII/482 (L, bed 14),

and prob a ble microconch ZPAL AmVII/ 472 (M).

Re marks. — Tornoceras applanatus (Gürich, 1896) based on a spec i men from the Up per Łagów beds(Sacculus−Bank) prob a bly re fers to the same spe cies. All avail able spec i mens are min ute in size, up to 22 mm in di am e ter, three of them with a con den sa tion of su tures sug ges tive of ma tu rity. Per haps also Gomi− re−monomeroceras (Tornoceras) simplicius subacutum Sobolew, 1914 from the clymeniid lime stone atŁagów− Dule be longs to this spe cies, al though only the shape of ap er ture makes it dif fer ent from as so ci atedProtornoceras simplicius. In the Czarnocki’s col lec tion housed at the State Geo log i cal In sti tute there is aprob a bly adult spec i men IG 284 II.290 about 52 mm in di am e ter, col lected from the grey “Laevigites” lime −stone (Nodosoclymenia bed?) at Ostrówka. Its su ture is char ac ter is ti cally simple but growth lines are notrecognizable.

An other ques tion is whether the North Amer i can type pop u la tion of T. crebriseptum is truly conspecificwith the Holy Cross Moun tains ma te rial. Spec i mens from the Three Forks Shale are py ritic in ter nal mouldswith open um bi li cus, which may ex press an in creased thick ness of the shell form ing a kind of umbonal cal lus (see Fig. 144A). This char ac ter was used by Korn and Ti tus (2006) to sub stan ti ate trans fer of the spe cies toPernoceras Schindewolf, 1922, the type spe cies of which is Protornoceras kochi (Wedekind, 1908. TheMontana spec i mens show a con i cal de pres sion around umbo, re sem bling the Pol ish spe cies of Protornocerasfrom Kowala in this re spect but dif fi cult to com pare with spec i mens from Łagów and Jabłonna hav ing theorig i nal shell pre served. The spe cies may truly be re lated to P. kochi.

Dis tri bu tion. — The C. marginifera (sam ples Ł−5, 28, 29, 30, 31, and 33) and pos si bly the P. trachyterazones at Łagów−Dule and Jabłonna (bed 24), per haps L. styriacus Zone at Ostrówka; re worked at Sieklucki’sbrickpit in Kielce.

196 JERZY DZIK

Fig. 144. Late spe cies of Tornoceras from the mid Famennian of the Holy Cross Moun tains. A–D. T. crebriseptum Ray mond,1909; growth lines traced from im prints on the conch in ter nal mould, su ture, and views of spec i men ZPAL AmVII/642 from theC. marginifera Zone at Łagów−Dule (A, sam ple Ł−5), growth lines of spec i men ZPAL AmVII/646 from the P. trachytera Zone at Jabłonna (B, bed 24) with nu mer ous ven tral shell thick en ings, but do not show ing su ture, pos si bly be long ing to the same spe cies; and lar val (ammonitella) conchs AmVII/1916 and 1614 from the same zone (C, D, sam ple Ł−38). E, F. T. pseudobilobatumDybczyński, 1913 from the mid C. marginifera Zone at Kowala; growth lines traced from im prints on the conch py ritic in ter nal

mould, su ture, and views of spec i mens ZPAL AmVII/1004 (E) and ZPAL AmVII/112 (F).

Tornoceras pseudobilobatum Dybczyński, 1913(Figs 144E, F and 159)


Ma te rial. — Six spec i mens.Di ag no sis. — Su ture with in cip i ent dorsolateral lobe and rel a tively straight course of the ventrolateral

sad dle, oth er wise gen er al ized tornoceratid conch.Re marks. — In the two best pre served rel a tively large spec i mens, the siphuncle po si tion is some what

asym met ric. Ju ve nile whorls are gently rounded, any re la tion ship to Falcitornoceras bilobatum is thus un −likely, con trary to at tri bu tion of this spe cies to F. falcatum by Becker (1993b). This is prob a bly rather an ad −vanced mem ber of the Tornoceras branch.

Dis tri bu tion. — The mid C. marginifera Zone at Kowala; re worked at Sieklucki’s brickpit in Kielce.

Genus Polonoceras Dybczyński, 1913

Type spe cies: Polonoceras planum Dybczyński from the Famennian of Kielce, Holy Cross Moun tains.

Di ag no sis. — Acute au ri cles of the ap er ture, in vo lute discoidal conch with small um bi li cus, flat sides and more or less tab u late venter; weak ventro−lat eral vault ings may de velop in the septum.

Re marks. — The in cip i ent vault ings of the sep tum seems to be an ex pres sion of lo cally de layed with −drawal of the vis ceral sac, at tached to the shell with muscles.

Polonoceras sandbergeri (Foord et Crick, 1897)(Figs 145A and 159)

Type ho ri zon and lo cal ity: Prob a bly early Famennian at Nehden, Rhenish Slate Moun tains (Becker 1993b).

Ma te rial. — Two spec i mens.Di ag no sis. — Au ri cles of the ap er ture with rel a tively blunt, par a bolic profile.Re marks. — The com plex no men cla tor ial his tory of the spe cies was clar i fied by Becker (1993b) who

clas si fied it as Truyolsoceras sandbergeri. It seems that it rep re sents an early part of the P. planum lin eageand I find such ge neric dis tinc tion re dun dant. Per haps Gomi−monomeroceras (Tornoceras) dorsoplanumavaricatum Sobolew, 1914 from the Lower Łagów Beds rep re sent this spe cies, al though its some what widerumbo may sug gest af fin i ties to the Protornoceras lineage.

Dis tri bu tion. — The early K. crepida Zone (bed 6) at Jabłonna.

Polonoceras bashkiricum Bogoslovsky, 1971(Figs 145B–D and 159)

Type ho ri zon and lo cal ity: Up per part of the Cheiloceras Stufe, 200 m from the mouth of the Ishikay River, South Urals(Bogoslovsky 1971).

Ma te rial. — Three spec i mens.Di ag no sis. — Acute tips of au ri cles of the ap er ture; dis tinctly tab u late venter of ma ture spec i mens,

rounded in juveniles.Re marks. — The Frasnian−style su ture and the subtrapezoidal cross sec tion of the whorl make this spec i −

men sim i lar to the holotype of Linguatornoceras linguum (see Becker 1993b, pl. 3: 13, 14). Conchs of theFamennian spe cies at trib uted to the ge nus by this au thor are, how ever, dif fer ent from the Łagów spec i men inhav ing a very nar row flank lobe of the su ture and rounded mar gins of the um bi li cus. In the lat ter as pect, thespec i men dis cussed re sem bles the stratigraphically older spe cies at trib uted here to Gundolficeras korni(Becker, 1993) and Protornoceras simplicius, be ing dif fer ent in the subtrapezoidal whorl sec tion and deepflank lobe of the su ture. Spec i mens from sam ple Ł−14 ten ta tively at trib uted to the lat ter spe cies may also be −long here, as well as the spec i men from the Up per Łagów Beds il lus trated by Sobolew (1914) as Tornocerasescoti Frech, 1902. The spe cies seems both mor pho log i cally and stratigraphically tran si tional be tween P.sandbergeri and P. planum.

Dis tri bu tion. — The late K. crepida (bed 15) and prob a bly the C. marginifera (bed 17) zones atJabłonna; the C. marginifera Zone at Łagów−Dule; pos si bly also re worked at Sieklucki’s brickpit in Kielce.


Polonoceras planum Dybczyński, 1913(Figs 145E–H and 159)


Ma te rial. — 12 spec i mens.Di ag no sis. — Dis tinctly tab u late venter even in ju ve nile spec i mens, nar row um bi li cus.Re marks. — Tips of au ri cles are lo cated some what dor sally to the tab u late venter, where they some times

leave a shal low fur row on the phragmocone in ter nal mould. In this re spect the spe cies is dif fer ent from P.dorsoplanum, its prob a ble successor.

198 JERZY DZIK

DDD

Fig. 145. Un de rived spe cies of Polonoceras from the mid Famennian of the Holy Cross Moun tains. A. P. sandbergeri (Foord etCrick, 1897); growth lines, su ture, and views of spec i men ZPAL AmVII/340 from the K. crepida Zone at Jabłonna (bed 6). B–D. P.bash kiricum Bogoslovsky, 1971 from the C. marginifera Zone; growth lines and views of spec i men ZPAL AmVII/354 fromJabłonna (B, bed 17), su ture of ZPAL AmVII/264 (C, bed 15), and views of ZPAL AmVII/1093 from Łagów (D). E–H. Polono −ceras planum Dybczyński, 1913 from the mid C. marginifera Zone at Kowala; su ture and view of spec i men ZPAL AmVII/1009

(E), views of ZPAL AmVII/951 and 952 (F, G), su ture and sep tum of ZPAL AmVII/1010 (H).


Polonoceras dorsoplanum (Sobolew, 1912)(Figs 146A–E and 159)

Type ho ri zon and lo cal ity: Black clymeniid lime stone at Łagów−Dule, Holy Cross Moun tains (Sobolew 1912b).

Ma te rial. — 21 spec i mens.Di ag no sis. — Deep con i cal um bi li cus with raised mar gins, acute tips of au ri cles de mar cate the tab u late

venter.Re marks. — Some spec i mens show pe ri odic con stric tions on the conch flanks. Dis tri bu tion. — The C. quadrantinodosa (sam ples Ł−28 and 33), C. marginifera, and P. trachytera

zones at Łagów−Dule.

Polonoceras sudeticum (Renz, 1914)(Figs 146G, H and 159)

Type ho ri zon and lo cal ity: Clymeniid lime stone at Dzikowiec, the Sudetes.


Fig. 146. Ad vanced spe cies of Polonoceras from the Famennian of Po land. A–E. P. dorsoplanum (Sobolew, 1912) from the C.marginifera Zone at Łagów; growth lines, su ture, and views of spec i men ZPAL AmVII/343 (A), su ture and sep tum of ZPALAmVII/350 (B), views of spec i mens ZPAL AmVII/359, 358, and 351, re spec tively (C–E). F. Spe cif i cally un de ter min able lar val(or em bry onic) spec i men ZPAL AmVII/1837 from the L styriacus Zone at Kowala (sam ple Ko−174) in the Holy Cross Moun −tains. G, H. P. sudeticum (Renz, 1914) from the P. jugosus (or D. trigonica) Zone at Dzikowiec in the Sudetes; con stric tion of the

conch and views of ZPAL AmVII/151 (G) and su ture of ZPAL AmVII/090 (H).

Ma te rial. — Three spec i mens.Di ag no sis. — Closed um bi li cus of con i cal ap pear ance demarkated by a spi ral ridge; roundly tab u late

venter; se rial con stric tions on conch flanks par al lel to the ap er ture, which bears acute au ri cles; su ture with adeep, narrow flank lobe.

200 JERZY DZIK

Fig. 147. Spe cies of Armatites from the mid Famennian of Po land. A, B. Early A. lateroconcavus (Dybczyński, 1913) from theearly P. trachytera Zone at Jabłonna (bed 25); growth lines, su ture, and views of spec i men ZPAL AmVII/341 (A, bed 21), andviews of prob a bly ma ture liv ing cham ber ZPAL AmVII/166 (B). C–E. Late A. lateroconcavus (Dybczyński, 1913) from the L.styriacus Zone at Jabłonna (A, bed 26) and the late P. trachytera Zone at Łagów−Dule (D, E); growth lines, su ture, and view (C)of spec i men ZPAL AmVII/342, views of frag men tary liv ing cham bers ZPAL AmVII/356 and 349 (D, E). F. Type spec i men ofA. lewinskii (Dybczyński, 1913) from Qua ter nary de pos its at Sieklucki’s brickpit in Kielce (re pro duced af ter Dybczyński 1913,

pl. 2: 10).

Re marks. — Renz (1914) iden ti fied his only spec i men from Dzikowiec lack ing any su ture as a new va ri −ety of Oxyclymenia ornata. The con stric tions (not in ter nal thick en ings!) in its conch, ex pressed both on nu −clei and the conch sur face, are so char ac ter is tic that there is lit tle doubt that the newly col lected topotypespec i mens are conspecific. The acute au ri cles and deep lobe of the su ture point to Polonoceras dorsoplanumas the ancestor of the species.

Dis tri bu tion. — The P. jugosus (or D. trigonica) Zone at Dzikowiec.

Genus Armatites Becker, 1993Type spe cies: Goniatites planidorsatus Münster, 1839 from the Famennian of Gattendorf in Franconia (Becker 1993b).

Di ag no sis. — Conch with tab u lar or bicarinate venter and flat sides, open um bi li cus, gen er al izedtornoceratid su ture and ap er ture with wide, gently rounded auricles.

Re marks. — Among spec i mens clas si fied in var i ous spe cies of the ge nus by Becker (1993b) only thelectotype fits the spe cific mor phol ogy of the mem bers of the lin eage rep re sented in the Holy Cross Moun −tains, at trib uted to Armatites here. He later cre ated the new ge nus Planitornoceras for spe cies with a sim i larap er tural pro file but a Protornoceras−like suture.

Armatites lewinskii (Dybczyński, 1913) (Figs 147F and 159)


Di ag no sis. — Conch with rounded venter and small um bi li cus.Re marks. — The de scrip tion by Dybczyński (1913) sug gests that this may be an early mem ber of the

Armatites lin eage, as in di cated by flat flanks and semi cir cu lar ap pear ance of flank lobes. Un for tu nately, hisde scrip tion and il lus tra tions re main the only source of data on this spe cies. The il lus trated spec i men is a ju ve −nile and its gen er al ized ap pear ance (es pe cially the rounded venter) may be partly due to the early ontogenetic stage. Both its mor phol ogy and prob a ble strati graphic or i gin (C. marginifera Zone) sug gest its an ces tral po si −tion in re spect to other spe cies of Armatites.

Dis tri bu tion. — Re worked at Sieklucki’s brickpit in Kielce.

Armatites lateroconcavus (Dybczyński, 1913) (Figs 147A–E and 159)


Ma te rial. — Seven spec i mens.Di ag no sis. — Flanks of the conch sub di vided into two zones by a spi ral fur row or es carp ment.Re marks. — Un like the stud ied spec i mens, in the lectotype of A. planidorsatus (Becker 1993b, pl. 11:

10) the spi ral el e va tion is re stricted to near the um bi li cus. All of them show rounded tips of ap er tural au ri cles, which makes them dif fer ent from other spec i mens at trib uted to the ge nus by Becker (1993b). Most of thespec i mens from Łagów are rep re sented only by ven tral parts of the body cham ber, only one of them pre −served the whole conch flank, with the lat eral fur row. This sit u a tion seems to be sim i lar to that in theholotype, as de scribed by Dybczyński (1913). The spec i men of Petter (1959, pl. 15: 30) from Gourara in Al −ge ria, reillustrated by Becker (1993b) and clas si fied by him in his A. nudus, shows a sim i lar ap er ture to thePol ish A. lateroconcavus, but narrower umbilicus.

There is an ap par ent evo lu tion ary change in the lin eage. The um bi li cus is rel a tively wide in the geo log i −cally old est spec i mens from Jabłonna (beds 21–25), with gently rounded mar gin and flat sides of the conch.In the some what youn ger ma te rial from Łagów (sam ple Ł−14), a shal low fur row in the mid dle of the flanksde vel oped, achiev ing the form of a prom i nent es carp ment in the youn gest spec i men from Jabłonna (bed 26).Closely sim i lar form oc curs also in the Prolobites ho ri zon in vi cin i ties of the vil lage Spasskij in the SouthUrals (Perna 1914, pl. 2: 9).

Dis tri bu tion. — Prob a bly the late C. marginifera Zone at Jabłonna (bed 21), the P. trachytera Zone atŁagów−Dule (e.g., sam ple Ł−14) and Jabłonna (beds 23–25); and the L. styriacus Zone at Jabłonna (bed 26);re worked at Sieklucki’s brickpit in Kielce.

Genus Protornoceras Dybczyński, 1913Type spe cies: Protornoceras polonicum Dybczyński, 1913 from the Famennian of Kielce.


Di ag no sis. — Sim pli fied su ture with a very shal low flank lobe and usu ally al most straight course of itsventrolateral part; weak ventro−lat eral vault ings may de velop in the sep tum; open umbilicus.

Re marks. — This seems to be a sep a rate branch of the tornoceratids evolv ing to wards the clymenoid ap −pear ance of the conch and su ture. Var i ous forms oc cur to gether in the black shale with pyritized ammonoidsat Kowala, which sug gests their im mi gra tion to the Holy Cross Moun tains area with the sea trans gres sion ofthe C. marginifera Zone (per haps at its later stage). Prob a bly the least de rived of them is P. aphyllitiforme,with al most closed umbo, the most de rived is P. mi ra bile. House (1970) pro posed the ge nus Tornia for thelat ter. Per haps it would be truly rea son able to put the protornoceratids with nar row and wide um bi li cus intodif fer ent gen era, but the type spe cies of both Protornoceras and Tornia are widely umbilicate. Be cause of the high pop u la tion vari abil ity, their sep a ra tion seems su per flu ous. The phylo gen eti cally clos est Kirsoceras dif −fers in the subventral position of the siphuncle (Bogoslovsky 1971).

Protornoceras aphyllitiforme Dybczyński, 1913(Figs 148A–C and 159)


Ma te rial. — 14 spec i mens.Di ag no sis. — Nar row um bi li cus with the flat ad join ing wall of the conch giv ing it a con i cal ap pear ance,

prom i nent rounded au ri cles of the aperture.Re marks. — The holotype su ture is sim i lar to that in ju ve nile spec i mens from Kowala. Adult su ture (in

spec i mens up to 34 mm in di am e ter) is more tornoceratid, but it is highly vari able. The sam ple is ev i dentlyho mog e nous, be ing taken from a sin gle black clay in ter ca la tion in the stratigraphically uncondensed, thick

202 JERZY DZIK

Fig. 148. Spe cies of Protornoceras with nar row um bi li cus from the Famennian of Po land. A–C. P. aphyllitiforme Dybczyński,1913 from the mid C. marginifera Zone at Kowala; growth lines in ferred from wrin kles on the sur face of py ritic in ter nal mould,su ture, and views of spec i men ZPAL AmVII/173 (A), views of spec i men ZPAL AmVII/1007 (B), and sep tum of ZPALAmVII/956 (C). D. Protornoceras simplicius (Sobolew, 1914) growth lines, su ture, and views of spec i men ZPAL AmVII/508from the P. trachytera Zone at Łagów−Dule (sam ple Ł−9). E. Protornoceras simplificatum (Sobolew, 1914) from Sieklucki’s

brickpit in Kielce (re pro duced from Sobolew 1914, pl. 9: 27, 28).

rock se quence. An angulation at the dor sal slope of the flank lobe makes some of the Kowala spec i mens sim −i lar to the holotype of Polonoceras latum Dybczyński, 1913, which may ap pear conspecific. Other spec i −mens show a dor sally lo cated in den ta tion in the lobe, hav ing its oblique ven tral slope al most straight. This in −den ta tion cor re sponds to a small vaulted area of the sep tum, a sit u a tion closely sim i lar to that in theopen−umbilicate Protornoceras siemiradzkii Dybczyński, 1913. How ever, these are dis tinct spe cies dif fer ent in the di am e ter of umbo, the trait less vari able than the course of su ture in the Kowala ma te rial. Tornocerasobliquum Perna, 1914 from the Prolobites ho ri zon of the Urals shows a sim i lar conch form and suture butunsimilar, pointed auricles of the aperture.


Protornoceras simplicius (Sobolew, 1914)(Figs 148D and 159)

Type ho ri zon and lo cal ity: Black clymeniid lime stone at Łagów−Dule, Holy Cross Moun tains.

Ma te rial. — 13 spec i mens.Di ag no sis. — Weakly de vel oped au ri cles of the ap er ture; closed con i cal um bi li cus.Re marks. — Spec i mens from sam ple Ł−14, prob a bly rep re sent ing the type ho ri zon, do not show su ture

and may be clymenias. In the holotype, the su ture is more Tornoceras−like than in spec i mens from the C.marginifera Zone (sam ple Ł−9) but it fits well within the range of vari abil ity shown by the stratigraphicallysome what older spe cies from Kowala.

Dis tri bu tion. — The C. marginifera (sam ple Ł−9) and P. trachytera (sam ple Ł−14) zones at Łagów−Dule, the L. styriacus Zone (trench rIVc of Żakowa et al. 1986) at Jabłonna.

Protornoceras simplificatum (Sobolew, 1914)(Figs 148E and 159)


Di ag no sis. — Evolute conch with umbo about one fifth of its di am e ter; oval cross sec tion of the whorl;more or less sim pli fied suture.

Re marks. — No spec i men of this mor phol ogy has been found by my self in the topotype stra tum. Pos si −bly some other taxa by Sobolew (1914) based on Sieklucki’s brickpit ma te rial be long here: Gomi−re− monomeroceras (Tornoceras) flexuosum and G. (T.) simplicius rotundatum.

Dis tri bu tion. — The P. trachytera Zone at Łagów−Dule; pos si bly also re worked at Sieklucki’s brickpit in Kielce.

Protornoceras polonicum Dybczyński, 1913(Figs 149A–G, 150A, and 159)


Ma te rial. — 33 spec i mens.Di ag no sis. — Open um bi li cus of di am e ter about one tenth of that of ma ture conch; subtrapezoidal cross

sec tion of the whorl; rounded au ri cles of the aperture.Re marks. — Sim i larly to other spe cies of Protornoceras, the su ture is very vari able, es pe cially in re spect

to con cav ity of the flank lobe, even within the same spec i men. Width of the umbo seems to be of a higher tax −o nomic value, al though it is vari able, too (note that all ear lier il lus trated spec i mens are py ritic in ter nalmoulds). Sev eral spe cies pro posed for spec i mens from Sieklucki’s brickpit by Dybczyński (1913), in clud ingP. siemiradzkii, P. bilobatiforme, P. kielcense, P. ornatum, and P. zuberi, and by Sobolew (1914), such asGomi−re−monomeroceras (Tornoceras) genulobatum planum, G. (T.) planilobum, G. (T.) planilobum angu −lato lobatum, G. (T.) planilobum avaricatum, and G. (T.) dorsatum, may be long here. The holotype of P.polonicum is a ju ve nile spec i men with not fully developed suture.

Dis tri bu tion. — The mid C. marginifera Zone at Kowala and Łagów Słupecka 73 (sam ple ŁSł73−4); re −worked at Sieklucki’s brickpit in Kielce. Spec i mens with a sim i lar conch form but the su ture not ex posed oc −cur in the clymeniid lime stone at Łagów−Dule.


204 JERZY DZIK

Protornoceras curvidorsatum (Sobolew, 1914)(Figs 149H−M and 159)


Ma te rial. — Ten spec i mens.Di ag no sis. — Evolute conch with umbo width ap proach ing one third of its di am e ter; subtrapezoidal cross

sec tion of the whorl; more or less sim pli fied suture.Re marks. — In my ma te rial only ju ve nile spec i mens ex hibit su ture, which ap pears to be ex tremely sim −

pli fied. Growth lines are prom i nent, in one spec i men nu mer ous ap er tural thick en ings de vel oped along itsventer. From the geo log i cally older P. mi ra bile the spe cies dif fers in the nar rower umbo and higher whorls.Pos si bly some other taxa based by Sobolew (1914) on the Łagów−Dule clymeniid lime stone be long here:Gomi−re−monomeroceras (Tornoceras) planilobum arcuatolobatum, G. (T.) sinuvaricatum, and G. (T.)umbilicatoides.

Dis tri bu tion. — The P. trachytera Zone (sam ple Ł−14) at Łagów−Dule; pos si bly also re worked atSieklucki’s brickpit in Kielce.

Protornoceras mirabile Dybczyński, 1913(Figs 149N–P, 150B, and 159)


Ma te rial. — Nine spec i mens.Di ag no sis. — Evolute conch with the umbo width ap proach ing half of its di am e ter; subtrapezoidal cross

sec tion of the whorl; more or less sim pli fied suture.Re marks. — This is the type spe cies of Tornia House, 1970. The holotype shows a mod er ately evolute

conch, al most straight course of the su ture and marks of prom i nent rounded au ri cles. In my ma te rial theshape of ap er ture is dif fi cult to trace. How ever, sim i larly to other spe cies of the ge nus, P. mi ra bile pop u la tion from Kowala shows a tre men dous vari abil ity in the su ture and also, al though not so wide, in um bil i cal width.The whorl cross sec tion shows a re mark able vari abil ity rang ing from al most tab u late venter and flat flanks toa quite weak angulation. This makes likely conspecificity of some other taxa by Sobolew (1914) based on


Fig. 149. Spe cies of Protornoceras with wide um bi li cus from the Famennian of the Holy Cross Moun tains. A–G. P. polonicumDybczyński, 1913 from the C. marginifera Zone at Łagów lo cal ity Słupecka 73 (A, sam ple ŁSł73−4; with pre served shell wall) andKowala (B–G; py ritic in ter nal moulds); growth lines, su ture, and views of spec i men ZPAL AmVII/971 (A), su ture and views ofZPAL AmVII/081 (B), su tures of spec i mens ZPAL AmVII/158 and 909 (C, D), and views of ZPAL AmVII/1006, 1001 and 158(E–G), re spec tively. H–M. P. curvidorsatum (Sobolew, 1914) from the P. trachytera Zone at Łagów−Dule (H–J, L, M, sam ple Ł−14;K, loose spec i men), growth lines of spec i men ZPAL AmVII/565 lack ing su ture and of ques tion able af fin ity (H), su tures of ZPALAmVII/705 and 543 (I, J), views of spec i mens ZPAL AmVII/ 558 and 541 (L, M, sam ple Ł−14). N–P. P. mi ra bile Dybczyński, 1913from the mid C. marginifera Zone at Kowala; growth lines in ferred from wrin kles on the sur face of py ritic in ter nal mould, su ture, and views of spec i men ZPAL AmVII/146 (N), su ture and views of spec i men ZPAL AmVII/114 (O), views of spec i men ZPAL

AmVII/114 (P), and su ture and views of ju ve nile spec i men ZPAL AmVII/120 (O).

®

Fig. 150. Vari abil ity of conch di men sions (based on mea sure ments of py ritic in ter nal moulds) of Protornoceras from the mid C.marginifera Zone at Kowala.A. P. polonicum Dybczyński, 1913. B. P. mi ra bile Dybczyński, 1913.

Sieklucki’s brickpit ma te rial: Gomi−re−protomeroceras alobatum, Gomi−re−monomeroceras umbilicatum, G.umbilicatoides, G. (Tornoceras) simplificatum rotundatum, and G. (T.) s. subacutum.


Genus Pseudoclymenia Frech, 1897

Type spe cies: Clymenia pseudogoniatites Sandberger, 1856 from the Famennian of the Rhenisch Slate Moun tains.

Di ag no sis. — Aturia−like flank lobe and in cip i ent dorsolateral lobe of the su ture, evolute conch.

Pseudoclymenia dillensis (Drevermann, 1901)(Figs 151A, B and 159)

Type ho ri zon and lo cal ity: Block of clymeniid lime stone from a vol ca nic brec cia at Langenaubach near Haiger, RhenishSlate Moun tains.

Ma te rial. — One spec i men.Di ag no sis. — Some what tab u late venter of evolute conch with umbo reach ing more than one third of the

conch di am e ter.Re marks. — The su ture in the only avail able spec i men is in com plete, but pointed tip of the flank lobe in −

di cates that this is a spe cies of Pseudoclymenia. Its flat tened venter sug gests spe cies iden tity with the ma te rial of Bogoslovsky (1971; but not Petersen 1975). The orig i nals of Drevermann (1901, pl. 14: 4a, b) are in ter nalmoulds and the venter of the spec i men il lus trated by him is rounded. The rel a tively nar row umbo (11.5 mm at the conch di am e ter 30 mm) fits the spec i men from Łagów. Lar val (or em bry onic) and early postlarvalphosphatized conchs from the same sam ple (Ł−38) may also rep re sent this spe cies. Gomi−monomeroceras(Tornoceras) evolutum of Sobolew (1914) from Sieklucki’s brickpit may also rep re sent this or a less de rivedspe cies of the ge nus, as sum ing that the min ute orig i nal is a juvenile with the suture somewhat worn out.

206 JERZY DZIK

Fig. 151. Spe cies of Pseudoclymenia from the Famennian of the Holy Cross Moun tains. A, B. Pseudoclymenia dillensis(Drevermann, 1901) from the C. marginifera Zone at Łagów−Dule (sam ple Ł−38); growth lines, su ture, and views (A) of spec i −men ZPAL AmVII/1801 and lar val (or em bry onic) conch ZPAL AmVII/1818. Note the dis po si tion of wrin kle layer near the ap −er ture and the sud den change in ap er tural mor phol ogy (emer gence of au ri cles) in di cat ing a meta mor pho sis from the lar val conchto teleoconch. The lack of growth in cre ments on the lar val conch sug gests that the lar val de vel op ment took place com pletelywithin the egg cov ers and that the meta mor pho sis may have been si mul ta neous with hatch ing. C. Pseudoclymenia fundifera

(Perna, 1914) from the P. trachytera Zone at Ostrówka, growth lines, su ture, and views of spec i men IG 284.II.56.

Dis tri bu tion. — The C. marginifera Zone (sam ple Ł−38) at Łagów−Dule; pos si bly also re worked atSieklucki’s brickpit in Kielce.

Pseudoclymenia fundifera (Perna, 1914)(Figs 151C and 159)

Type ho ri zon and lo cal ity: Prolobites ho ri zon, Ural River shore 6 km north of the vil lage Kirsy, south ern Urals.

Ma te rial. — One spec i men.Di ag no sis. — Con vex ventrolateral lobe of the su ture; in dis tinct au ri cles of the ap er ture; oval whorl cross

section.Dis tri bu tion. — The P. trachytera Zone at Ostrówka.

Genus Ostrovkites gen. n.Type spe cies: Ostrovkites numismalis sp. n.

Der i va tion of name: From Ostrówka hill, the type lo cal ity of the type spe cies.Di ag no sis. — Su ture with pointed flank lobe and small sec ond ary ven tral lobes; discoidal conch with a

ten dency to de velop lon gi tu di nal (spi ral) striation.Re marks. — Su ture of the type spe cies is sim i lar to that of the praeglyphioceratid Lagowites nivae

(Sobolew, 1914) but the au ri cles of the conch ap er ture are a tornoceratid fea ture. The strati graphic po si tionsug gests that the subventral lo ba tion of the su ture orig i nated in de pend ently of that in ad vanced Maenecerasspecies.

Ostrovkites numismalis sp. n.(Figs 152 and 159)

Holotype: Spec i men IG 284.II.33 (Fig. 152C).Type ho ri zon and lo cal ity: Black clymeniid lime stone of the P. trachytera Zone at Ostrówka, Holy Cross Moun tains.Der i va tion of name: From Latin numisma – coin, re fer ring to small size and flat shape of the conch.

Ma te rial. — Eight spec i mens.Di ag no sis. — Min ute conch with se rial in ter nal thick en ings fol low ing the ap er ture on its whole ex tend,

su ture with an ad di tional nar row ven tral lobe de vel oped on the slope of ventrolateral saddle. Re marks. — There is a pos si bil ity that Sobolew’s (1914) Oma−monomeroceras (Cheiloceras) sinu vari −

catum from the Lower Łagów Beds rep re sents the same lin eage. The course of in ter nal thick en ings in the sin −gle known spec i men (Sobolew 1914: pl. 7: 10) is sim i lar to that in spec i mens from Ostrówka, al though they


Fig. 152. Lagowites−like tornoceratid Ostrovkites numismalis sp. n. from the P. trachytera Zone at Ostrówka, Holy Cross Moun −tains; growth lines, su ture, and views of spec i men ZPAL AmVII/1116 (A, sam ple Ost−16), growth lines and su ture of spec i men

ZPAL AmVII/016 (A, sam ple Ost−10), views of the holotype IG 284.II.33 (C).

are less sin u ous. Un for tu nately, the su ture was poorly pre served in the Sobolew’s spec i men, which isprobably lost.

Dis tri bu tion. — The P. trachytera Zone at Ostrówka (sam ples Ost−10 and Ost−16).

Family Posttornoceratidae Bogoslovsky, 1962Di ag no sis. — Su ture line with sec ond ary umbonal lobe and a ten dency to de velop ad di tional ventrolateral

lobes (VLU1:ID ® V L1LU1UID); conch ap er ture with au ri cles; rel a tively high whorl ex pan sion rate.Re marks. — Early ontogenetic stages of Discoclymenia iden ti fied by Becker (1995, fig. 8c, d) show that

the dorsolateral lobe in ad vanced forms is ho mol o gous to that in Gundolficeras, thus U1 in Ruzhentcev’s no −ta tion.

Genus Gundolficeras Schindewolf, 1936Type spe cies: Lobotornoceras bicaniculatum Petter, 1959 from the Famennian of Al ge ria.

Di ag no sis. — Gen er al ized tornoceratid conch with ventrolateral fur rows at early de vel op men tal stages;su ture with a deep, sym met ri cal flank lobe and a ten dency to de vel oped dorsolateral lobe.

Re marks. — House and Price (1985) ques tioned re la tion ships be tween the Famennian tornoceratids with com plex su ture tra di tion ally clas si fied in Lobotornoceras Schindewolf, 1936 and the type spe cies of the ge −nus, Goniatites ausavensis Steininger, 1856 from the mid Frasnian Büdesheim Shale of the Eifel Moun tains(House 1978). Fol low ing this sug ges tion, Becker (1993b) trans ferred the Famennian spe cies with ventro −lateral fur rows at early ontogenetic stages to Falcitornoceras. They dif fer rather sub stan tially from the typespe cies of that ge nus in hav ing rounded au ri cles and lack ing the char ac ter is tic ju ve nile or na men ta tion. Later(Becker 1995) in tro duced the ge nus Gundolficeras for some mem bers of this branch. It seems to be the firstlink in the tornoceratid lin eage lead ing to more and more com plex su ture, ter mi nat ing in the acutely discoidal Discoclymenia.

Gundolficeras korni (Becker, 1993)(Figs 153A and 159)

Type ho ri zon and lo cal ity: Early Famennian Nehden Shale at Nehden−Schurbusch, Rhenish Slate Moun tains.

Ma te rial. — Five spec i mens.Di ag no sis. — Closed um bi li cus with the flat ad join ing wall of the conch giv ing it a con i cal ap pear ance,

rounded flank lobe of the su ture of oth er wise gen er al ized tornoceratid type.Re marks. — The su ture, well vis i ble in two ju ve nile spec i mens, fits well that of the type spec i men. In the

small est one, in dis tinct ventrolateral fur rows are rep re sented. An adult in com plete spec i men with bodycham ber, of es ti mated di am e ter about 55 mm, shows in dis tinct growth lines with typ i cally tornoceratid rather rounded prom i nent au ri cles of the ap er ture. The conch ge om e try makes this spe cies sim i lar to Protornocerassimplicius, which has a rather sim pli fied and vari able su ture. Becker (1993b) in cluded it into Falcitornoceras al though its ju ve nile conch or na men ta tion re mains un known. Be cause of the shape of um bi li cus and a ratherad vanced tornoceratid su ture I sug gest that this is not Falcitornoceras, but is rather re lated to the Polono −ceras branch at the stage preceding development of acute auricles.

Dis tri bu tion. — Prob a bly the K. crepida Zone at Jabłonna (wells w150a, b, c dug by Żakowa et al.1984).

Gundolficeras bilobatum (Wedekind, 1908)(Figs 153B–D and 159)

Type ho ri zon and lo cal ity: Early Famennian at Enkeberg, Rhenish Slate Moun tains.

Ma te rial. — Eight spec i mens.Di ag no sis. — Nar row ventrolateral sad dle, shal low dorsolateral lobe, and very deep and nar row dor sal

lobe of the su ture; closed um bi li cus.Re marks. — Becker (1993b) in cluded this spe cies into Falcitornoceras, re fer ring to fur rows on py ritic

in ter nal moulds, al though its ju ve nile conch or na men ta tion re mains un known. Such fur rows, pre served alsoon a ju ve nile conch from Łagów, are typ i cal of sev eral ear li est Famennian tornoceratids and may just be aprim i tive (plesiomorphic) fea ture. Gundolficeras rotersi Korn, 2002 from the Platyclymenia annulata Event

208 JERZY DZIK

stra tum at Kattensiepen in the Rhenish Slate Moun tains does not dif fer from the Holy Cross Moun tains ma te −rial in conch form an su ture. Growth lines are not pre served in my ma te rial and this precludes reliablecomparison.

Dis tri bu tion. — The mid C. marginifera Zone at Kowala; prob a bly the C. quadrantinodosa Zone (sam −ples Ł−29, 30, and 34) at Łagów−Dule; re worked at Sieklucki’s brickpit in Kielce (Sobolew 1914).

Gundolficeras sp. n. aff. G. delepinei Petter, 1959(Figs 153E–I and 159)

Ma te rial. — Two spec i mens.Re marks. — Two spec i men from the Czarnocki’s col lec tion are sim i lar to Gundolficeras from the much

older strata at the same lo cal ity but dif fer in hav ing a rather wide ventrolateral sad dle and shal low dor sal lobe. The first fea ture is even more ap par ent in G. delepinei from the late Famennian of Al ge ria (Petter 1959),which may rep re sent the end mem ber of the same lin eage (see Becker 1995).

Dis tri bu tion. — Prob a bly the P. jugosus (or the late L. styriacus) Zone at Kowala.


Fig. 153. Lobotornoceras−like tornoceratid Gundolficeras from the Holy Cross Moun tains. A. Gundolficeras korni (Becker,1993) from the early Famennian at Jabłonna (prob a bly the K. crepida Zone); su ture of spec i men ZPAL AmVII/659 from wellw150c dug by Żakowa et al. (1984). B–D. Gundolficeras bilobatum (Wedekind, 1908) from the mid C. marginifera Zone atKowala (B, C) and prob a ble C. quadrantinodosa Zone at Łagów−Dule (D); su ture and views of spec i men ZPAL AmVII/169 (B),view of spec i men ZPAL AmVII/078 (C), su ture and sep tum of spec i men ZPAL AmVII/1864 (D, sam ple Ł−30). E–I.Gundolficeras sp. n. aff. G. delepinei Petter, 1959 from the late Famennian at Kowala (prob a bly the P. jugosus Zone), su ture andviews of spec i men IG 284.II.860 (E), su ture of unnambered spec i men from the same col lec tion (F) and lar val (or em bry onic)conchs ZPAL AmVII/1833, 1834 (G, H, sam ple Ko−187, the P. jugosus Zone), and 1836 (I, sam ple Ko−174, the L. styriacus

Zone) pos si bly rep re sent ing the same lin eage.

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