Evolution of the Goniatitaceae and Vis6an-Namurian Biogeography

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Evolution of the Goniatitaceae and Vis6an-Namurian biogeography

DIETERKORN

Korn, D. 1997. Evolution of the Goniatitaceaeand Vis6an-Namurian biogeography.- Acta Pal eontologica P olonica 42, 2, 177 -199.

Evolutionary lineages within the Carboniferous ammonoid superfamily Goniatitaceae can be recognized using cladistic and stratopheneticanalyses, showing that both ap- proaches lead to coinciding results. In the late Vis6an and Namurian A, ammonoid provinces can be defined by the distribution of lineages within the goniatite superfamily Goniatitaceae. The first province corresponds to the Subvariscan Realm (where the superfamily became extinct near the Vis6an-Namurian boundary), and the second em- braces the majority of the occurrences,e.g. the south urals, central Asia, and North America (where the superfamily with different independent lieages continued up into the late Namurian A). In the Vis6an, the superfamily was, in two short epochs, globally distributed with major transgressions,which probably led to migration events. The first is at the end of the late Vis6an A(G. fi.mbrians and G. spirifer Zones, when the genus Goniatites had a world-wide distribution with various species),and the second at the beginning of the late Vis6an C (L. poststriatumZone, when Lusitanoceras is globally distributed).

K e y w o r d s : Ammonoidea, Goniatitaceae, Early Carboniferous, phylogeny, palaeo- biogeography.

Dieter Korn [dieter.korn@uni+uebingen.de], Institut und Museum fiir Geologie und Palciontologieder Eberhard-Karls-(Jniversitrit, SigwartstraJJe10, 72076 Tiibingen, Ger- many.

Introduction

Late Vis6an and early Namurian goniatite faunashave been reported fiom numerous localities in the northernhemisphere, from Alaska andCanada, the Westernand Central United States,various regions in Europeand North Africa, Novaya Zemlya and the South Urals, Central Asia, as well as from China. They have beendescribed in several monographsand articles,mostly publishedduring the last 30 years.These data allow a comparisonof the faunalcompositions in the differentregions, and provide a tool for r78 Evolution of the Goniatitaceae: KORN

palaeogeographicalreconstructions of the plate configuration during the late Early Carboniferous. Ammonoid provincialism in the late Vis6an and early Namurian has already been postulatedseveral decades ago. Kullmann (1962) distinguishedbetween an epicon- tinental fauna (to which he referred the SubvariscanRealm), and a geosynclinalfauna (= Tethyal fauna), in which he placed the faunas collected from the Cantabrian Mountainsof Spain,North Africa, SouthlJrals, etc., and accountedfor dissimilarities in the faunal spectramostly to differencesin water depth. Kullm ann(1962) suggested that the two provinces, during the Namurian, contained different types of goniatites: the epicontinentalfauna being characterizedby goniatiteswith various ornamenttypes, while typical for the geosynclinal fauna were goniatites with modified internal struc- tures,such as advancedsutural development, and evolutecoiling of the inner whorls. The Goniatitaceaeis one of the major late Vis6an and early Namurian goniatite superfamilieswith an almost world-wide geographicaldistribution. It is recordedfrom numerous regions in Europe, from North Africa, North America, as well as from several localities between the Ural Mountains and Japan.The Goniatitaceaeis also a superfamilycontaining numerous species, of which most occur in the late Vis6an. Conchsof Goniatitaceaespecies display a ratherhigh numberof distinctivecharacters. Sutural features,conch geometry,and ornamentalfeatures allow cladistic analysesof these forms, and the densesampling enablesstratophenetic approach. Thus relation- shipsbetween the variousgenera can probably be uncovered. Discrimination of faunal provinces will be achievedherei by pointing out differen- ces in the distribution of independentevolutionary lineages. The data on which the cladistic and stratopheneticanalyses are based are taken from the literature (Ruzhencev & BogoslovskayaI97 L; Korn 1988),and the dataof the geographicaldistribution from the DatabaseGONIAI, Tiibingen(Korn & Kullmann 1996).

Conch morphology characters of the Goniatitaceae

Apomorphies. - The membersof the superfamily Goniatitaceae,like many Car- boniferousgoniatites, display a sutureline containingthe elementsEm E1 A L U I (terminology of Wedekind 1918), of which during phylogeny the El lobe (family Delepinoceratidae)or A lobe (family Agathiceratidae)became subdivided. The majority of the forms possessan E lobe with a generalv-shaped outline, and an acuteor subacuteventrolateral saddle.Characteristic for all the membersof the superfamily is the courseof ontogeneticdevelopment of the externallobe. In juveniles,it is Y-shaped, but during ontogeny it changedto a v-shaped outline. without exception among the Vis6an and Namurian species,all forms have pointed lobes (in contrast,the latest Carboniferousand Permianagathiceratids had roundedlobes and saddles).The conch is usually pachycdnicor globular with an extremelylow whorl expansionrate, and a very nalrow or closed umbilicus. Many forms bear crenulatedgrowth lines and/or spiral lines,which alwaysresult from crenulation. The phylogenetic origin of the superfamily Goniatitaceaeis unclear (Ruzhencev& Bogoslovskaya1970).In Centraland western Europe,the first speciesof the superfamily appearsnear the mid-vis6an boundary and can be classified in the genus ACTA PALAEONTOLOGICA POLONICA (42) (2) 179

Goniatites(Korn 1996),possessing aconch morphology dissimilarto any earlyVis6an goniatites. The pointed ventrolateral saddle of the suture line, maintained in the Jubsequentlin"ug"r, is unique amongthe early late Vis6angoniatites. Additionally, the naffow umbilicated, globose conch form and the strongly crenulatedgrowth lines on an otherwise unornamentedshell are novelties (apomorphies)in this group. Juvenile stages of species of Goniatiter seem to recapitulate an adult sutural outline of its ancestor,which probablypossessed a Y-shapedexternal lbbe'

Morphotogical trends in the evolution. - Afransformation of severalcharacters can be observedwithin the evolutionary history of the Goniatitaceae(Ruzhencev & Bogoslovskayalg70). Primarily, the suturaldevelopment displays a significantpat- tern, and allows separationof severalindependent evolutionary lineages, chnacteized by different modei of sutural subdivision (Figs 1, 2). Trends of sutural and conch transformationswithin the Goniatitaceaeare: (1) The median saddleheight increase(relative to the ventrolateral saddle). 0.40 in the earliestlate visean (e.g.,Goniatites hudsoni, G. crenistria), 0.50 in the late Vis6anB (e.g.,Arnsbergites, Hibemicoceras, Paraglyphioceras), 0.60 to 0.70 near the Vis6an-Namurianboundary (e.g., Neogoniatites, Dombarites), and more than 0.80 in the early Namurian(e.g., Platygoniatites, Delepinoceros, Proshu- mardites). This tendency is shown by almost all the evolutinary lineages, and a reversal apparentlydid not occur.The height increaseof the median saddleis a common feature u*ong many Carboniferousgoniatites (Ruzhencev & Bogoslovskaya1971, 1978)' henceit should not be used as a criterion for the definition ofhigher taxa. The height inctease,however, is important for the separationof generaand species (2) The E lobe widening (the width measuredat its half depth, relative to its total depth = E lobe index). On the basis of this character,four independentlineages can be recognized,these are: Goni.atites-+ Hypergoniatites, with no remarkable widening (E lobe index always between0.50 and 0.60), Paraglyphioceras -+ Neogonintites,with a continuouswidening rate during phylogeny (E lobe index 0.65 to 0.95), Lusitanoceras -+ Proshumnrdites,with a moderate widening of the E lobe (E lobe index 0.65 to 0.95,but with secondarydissection of the Alobe), and Lusitanoceras--> Delepinoceras,with a prominentwidening during phylogeny(E lobe index 0.65 to 1.50),leading to a secondarilyfiidentate El lobe' (3) The evolute coiling of the inner whorls. Derived specieswith evolute inner whorls evolve from completely involute forms, which stratigraphically occur at the beginning of the evolutionary history for example: the inner whorls are involute in Goniatites,Goniatitella, andin Hypergoniatites,but evolutein Hibernicoceras,Ams- bergites,Paraglyp,llisceras, Lusitanoceras, and Dombarjfes. Cross sections of stratigraphically later generaremain unknown, but, according to published photographs,it Ian^beassumed that in derivedforms suchas Delepinoceras and Proshumardites only very few initial whorls are evolute' 180 Evolution of the Goniatitaceae: KORN

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Fig. 1. Diagram showing the height increase ofthe median saddlein the evolution ofthe late Vis6an and early Namurian speciesof the superfamily Goniatitaceae.MS/h refers to the ratio betweenthe height of fhe median saddle(MS) and the total height of the ventrolateralsaddle (h). Note that there is a continuousheight increase (except for the late Vis6an genera Goniatitella and Hypergoniatites), and that no independent evolutionary lineages can be traced in the remaining species.In the box: Sutural parametersused in the diagrams and for the cladistic analysis. Seealso explanation to Fig. 2. ACTA PALAEONTOLOGICAPOLONICA (42) (2) 181

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0.40 0.50 0.60 0.70 0.80 0.90 1.00 1.10 1.20 1.30 1.40 1.50 ELIh

Fig. 2. Diagram showing the widening of the external lobe in the evolution of the late Vis6an and early Namurian species of the superfamily Goniatitaceae.El-/h refers to the ratio between the width of the external lobe (EL; measuredat half the height of the ventrolateral saddle) and the total height of the ventrolateral saddle (h). Note that a continuous widening takes place (except for the late Vis6an genera Goniatitella andHypergoniatites).ln the early Namurian,two independentevolutionary lineages can be seen,one leadin gto Proshumardires(family Agathiceratidae;with moderatewidening of the external lobe), and one leading to Delepinoceras (family Delepinoceratidae; with prominent widening of the extemal lobe).See insert in Fig. l. r82 Evolution of the Goniatitaceae: KORII

Table 1. Charactermatrix for the late Vis6an and Namurian speciesof the Goniatitaceae.Characters (18) and (20) are weighted as being of double value. Abbreviation: orn. - omament.

suture line conch orn.

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Eoglypiocerastruncatum 0 0 0 0 0 0 0 0 0 0 0 0 0 00000 t rtr; ri Goniatitescrenistria 1 0 0 0 0 0 0 0 0 0 0 0 0 00000 0 0 0 010 0 Amsbergitesfalcatus L I 0 I 0 0 0 0 0 0 0 1 0 0 00100 1 0 0 0 I 0i Hibernicocerashibemicum I 0 I 0 0 0 0 0 1 0 I 0 0 00100 100010 Paraglyphiocerasrotundurn)I 0 I 0 0 0 0 0 0 0 1 0 0 00000 r o o o]ool Lusitanoceraspoststriatum i I 0 I 0 0 0 0 0 0 I 1 0 0 00100 100110 Goniatitellaagricola 1 0 0 0 0 0 0 I 0 0 0 0 0 00100 011001 Neogoniatitesmilleri i 1 0 I 1 0 0 0 0 0 0 1 I 0 00000 1 0 0 0 0 0L Hypergoniatitesexiguus I 1 0 0 0 0 0 0 0 0 0 0 0 00000 00r000 Dombaritestectus 1 0 1 11 0'0 0 0 I 1 0 0 00110 100110 Proshumarditesdelepinei I 0 I I I 1 1 0 0 1 1 0 0 00111 1 0 0 1 1 oi Platygoniatitesmolaris 1 0 1 1 I 1 0 0 0 1 I 1 I 00100 10 0 1110 oyllltnocemybressony t O r I 1 I 1 l 0 0 I 1 I I t t t l o1I 0 0 1 I 0L

(4) The apertureshape in juveniles. Within the late Vis6an,two aperturaltypes in immature specimensare present: the rectiradiatetype, where the growthlines run in an almostradial direction (Goniatites,Hypergoniatites, Paraglyphioceras, and Goniati- tella wfth linear growth lines), and the prorsiradiate type, where the growth lines prominently arch forward on the flanks to form a broad ventral projection Q.4msber- gites, Hib ernicoc e ras, Lusitanoceras). ACTA PALAEONTOLOGICAPOLONICA (42) (2) 183

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Fig. 3. Cladogramrepresenting one of two most parsinomoustrees using PALIP3.0 (Swofford 1989) of the Goniatitaceae.Numbers of autapomorphiesand homoplasiesin both figures refer to condition (1) in the character matrix shown in Table 1. Note that the superfamily Goniatitaceae is characterized by one synapomorphy, the ontogenetic development from a juvenile Y-shaped external lobe towards an adult V-shapedexternal lobe (character0). Gonintites, which is regardedas the ancestorof all the other genera of the superfamily, does not possessan autapomorphy,but the other two independentlineages are defiled by (20) the higher whorl espansiori',rate(Goniatitella, Hypergonintites), and by (18) the evolution of the inner whorls, (2) the height increase of the median saddle, and (10) the widening of the external lobe (occuring in all the other genera). The latter clade is shown as an unresolved trichotomy. The ancestral Paraglyphioceras cannotbe definedby an autapomorphy.The sameis truefor Neogoniatifes,but this genus sharestwo homoplasieswith generaof the families Agathiceratidaeand Delepinoceratidae,the continuing heightincreaseofthemedian saddle (3), andthewidening ofthe externallobe (11).Neogoniatites,however, is distinguished by lacking some of the features from these genera, and is thus be interpreted as an independentlineage. The sutural developmentis then the main characterfor further hierarchized subdivi- sionoftheclade. The beginningsinuosity ofthe flanksofthe adventivelobe(15) inArnsbergitesisfollowed by the tectiform outline (9) in Lz sitanoceras.Fromthe latter genusderive the two families Agathiceratidae, defined by the angularity of the adventive lobe (1.6) and finally its trifurcation (1.7), and the family Delepinoceratidae,defined by the rapid widening of the extemal lobe (12, 13), leading to tridentation of its prongs (14) in the terminal stageof this lineage.Abbreviations : F. Agath. - family Agathiceratidae,F. Delep. - family Delepinoceratidae. 184 Evolution of the Goniatitaceae: KORN

Cladistic analysis. - Phylogeneticrelationships within the family Goniatitidaeand their immediatedescendants Delepinoceratidae and Agathiceratidae have been discussed byBogoslovskaya(1966)andRuzhencev&Bogoslovskaya(1970,1971).Theseauthors, however, acceptedonly four late Vis6an genera within the Goniatitidae (Goniatites, Hypergonintites,Neogoniatites, andHibernicoceras), and treated Goniatites as a very voluminousgenus unifying more than 30 different species.This conceptwas not followed by Korn (1988),who distinguishednine generawithinthe Goniatitidae: Goniatitesde Haan, 1825, HibernicocerasMoore & Hodson.1958. LusitanocerasPereira de Sousa.1923. Parag lyp hio c e ra s Bridrning,1923, ArnsbergitesKorn, 1988, Goniatitelb Korn, 1988, Ne o g o niatit es Ruzhencev& Bogoslovskaya, 197O, HypergoniatilesRuzhencev & Bogoslovskaya,1970, and JunggaritesLiang& W*9, 1991(problematic genus). Acladogram(Fig. 3), basedmainlyon suturalcharacters, is presentedhere showing proposedrelationships of the Early Carboniferousgenera of the Goniatitidae,Agathi- ceratidae,and Delepinoceratidae. For the cladisticanalysis, well known representative specieshave been chosen (Table 1). Included charactersare only those, which can be regarded as significant on the genus-level,such as the coiling of the innerwhorls, the sutureline (especiallythe ratios of dimensionsin sutural elements,although these are sometimesonly suitable for discriminationof species),and the aperturaloutline. Of these,the coiling of the inner whorls and the increaseof the later whorl expansionrate are relatively weighted as being of double value, since this has been shown to be a rather stable feature in the evolutionof late Vis6anammonoids (Ruzhencev & BogoslovskayalgTl; Korn 1988). Characterssuch as the conch ornamenttype, which are proved to be less stable,and often can only be usedfor discriminationon the species-levelwithin the genera,are largely omittedfrom the analysis. From the cladogram,the paraphyleticnature of the familiy Goniatitidae is obvious. This is especiallybecause the advancedtaxa in the phylogeneticfiee, which show anadvancedsutural development, have been cut-offto form the monophyleticfamilies Delepinoceratidaeand Agathiceratidae, characteized by different sutural develop- ments, but with conch shapeswhich do not vary much throughout their phylogeny. The transformationfrom Goniatites, which is regardedas ancestralto the succeeding generaHypergoniatites and Goniatitella, is unresolvedbecause of the lack of intermediateforms.

Evolutionary lineages within the Goniatitaceae

The cladogram(Fig. 3) can easilybe transformedinto an evolutionarytree (Fig. 4), in which the stratigraphicaldistribution of the generaand possibleancestor-descendant sequencesare presented. This indicatesthe rathergradual evolution ofthe superfamily ACTA PALAEONTOLOGICAPOLONICA (42) (.2) 185

Goniatitaceaewith severalindependent lineages, which will be describedbelow. Some of the proposedphylogenetic lineages are well documented,such as Lusitanoceras -) Platygoniatites-+ Delepinocerasby continuouswidening of the externallobe and subdivisionof its branches,and Lusitanoceras-+ Dombarites-+ Proshumarditesby successivewidening andtrifurcation of the adventivelobe. Gonintites. - In the late Vis6an A, the genusis representedby at least four successivespecies in Central Europe (Korn 1988). A morphological trend of sutural developmentcan be observed,which includes the following features: (1) widening of the E lobe: from imndex value 0.50 rn Goniatites hudsoni to 0.60 in Goniatites spirifer, (2) increaseof median saddle heighl from index value less than 0.40 tn Gonintites hudsoni to 0.50 in Goniatites spirifer. It is is uncefiain if the stratigraphically succeeding species form a single phyletic lineage. Ornamentaldifferences, e.g. the strongly biconvex courseof the growth lines in Goniatites crenistria and the presenceof spiral lines already tn Goniatites(?) globostriatus suggest that these species belong to side-branches.It is still unclear, however, if Goniatites(?) globostiatus in fact belongs to Goniatites, or rather to a beyrichoceratid genus.From the Rhenish Massif, no three-dimensionally preservedmaterial which could have been dissectedhas beentraced so far, and specimensfrom other regions do not display the suture line either. It can be stated with certainty, however, that at least Goniatites maximus var. saourensisPareyn, 1961 is, according to its suture line, a beyrichoceratid. This is confirmed by the recent collection of well preserved new material from near Erfoud, Tafilalt, Morocco. Conch and ofirament development lead to the interpretation that there was a phyletic -+ lineage Goniatites hudsoni -+ Goniatitesfimbriatus Goniatites spiifer. Goniatites -+ Hypergoniatiles. - Hypergoniatites is the latest Vis6an goniatite genus with an archaic conch morphology, resembling that of species from the late Vis6an A, but with a remmkable increaseof the whorl expansion rate (seecross sectionsin Ruzhencev& Bogoslovskaya 1971). It is characteized by a suture line that displays features similar to Goniatites crenistia, with a narrow extemal lobe, pointed ventrolateral saddle, and a V-shaped adventive lobe. The E lobe index is only 0.50 to 0.55) low ratios known elsewhere only in much older representativesof the superfamily Goniatitaceae(Fig. a). No speciesof Goniatites, however, has an external lobe that is, at its base,as narrow as in Hypergoniatites. There are no intermediate forms known which stratigraphically ltrnk Goniatites and Hypergoni- atites.The only genus with an intermediate stratigraphicaloccurrence is Goniatitella, which shares similar conch and sutural relations with Goniatites and Hypergoniatites, but markedly differs in ornament with coarse spiral lines, and a suture line with Y-shaped external lobe and rounded ventrolateral saddle.For thesereasons, it cannot be regardedas a direct ancestorof Hypergonirttites. The small size of Goniatitella specimens and their suture line that in the adult stage still displays a juvenile outline suggeststhat it is a paedomorphic (progenetic) form. Goniatites --> Pmaglyphioceras -> Neogoniatites. - After the abrupt decline of species of the family Goniatitidae with involute inner whorls at the end of the late Vis6an A, its representatives with open umbilicate juveniles dominate the goniatite faunas in Central and Westem Europe. The first genusof this kindis Paraglyphioceras,which already co-occurswithGoniatitesintheG. fimbrians Zone, representedthereby Paraglyphioceras(?)semistriatum (Nicolaus, 1963),known from crushed material, in which inner whorls and suture lines are not preserved. From the G. spiifer Zone, Paraglyphioceras radiatum (Hodson & Moore, 1959) is known, of which well preservedmaterial is available from heland (Hodson & Moore 1959) aswell asthe Harz Mountains (Korn 1990b; Gischler & Korn 1992).kr these specimens,the inner whorls and the suture line can be examined, allowing comparisonwith the ancestralGoniatites and descendantArnsbergites. Paraglyphioceras radiatum possessesa morphology which is advancedin comparison with the stratigraphically youngest species of Goniatites by the following set of features: (1) the juvenile whorls, up to 4 mm conch diameter, are evolute, (2) the external lobe depth is 1.20 of the width of the adventive lobe (E lobe index = 0.65), (3) the median saddleheight exceedshalfofthe ventrolateral saddle,and (4) the ventrolateral saddleis subacute,rather than acute as tnGoniatites. 186 Evolution of the Goniatitaceae: KORN

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Fig. 4. Phylogram displaying the proposedrelationships between the late Vis6an and early Namurian genera of the superfamily Goniatitaceae.The same apomorphiesand homoplasies as in Fig. 3 are used, but the genera are here arranged according their stratigraphical occurrence. Stratigraphical scheme of the early Namurianafter Ruzhencev& Bogoslovskaya(1971), and the late Vis6anafter Korn (1996).Open boxes show generawithout remmkable morphological trends, grey boxes show generawith striking morphological transformation during phylogeny. Cross sectionsare to the samemagnification, sutwe lines not to scale. ACTA PALAEONTOLOGICAPOLONICA (42) (2) 187

Fig.5. Palaeogeographicalmap ofEurope during the late visdan (modified afterTiegler 1989).

Additionally, the adult conch of Paraglyphioceras radiatumbears a falcate ornament, consisting mainly of densely spacedbiconvex riblets. Paraglyphioceras extends up into the P rotundum Zone. Only in this zone is the genus the most frequent, with the dominant species P rotundum Briiming, l923.In this species, the suture line displays more derived charactersin comparisonwith P radiatum,for examplethe extemal lobe is 1.5 wider than the adventive lobe (E lobe index = 0.75). A speciesthat closely resemblesP. rotundum is P. striatum (Sowerby, 1813) although the flanks of the lobes are much more sinuous. The sutural ratios of the two species, however, are very similar and suggesta similar stratigraphicalorigin of the species. The main trend in the development of the suture line of Gonintites continued within the genus Paraglyphioceras. A character in common with Goniatites is the shape of the aperture that remains stable during the Paraglyphioceras lineage. Neogoniatites has a similar conch form to Paraglyphioceras,blut differs in the outline of the suture line, which can be regarded as derived, the extemal lobe is wider, its width is two times greater than that of the adventive lobe (E lobe index = 0.90), and the median saddleis higher (0.60 of the ventrolateral saddle). It is especially the lack of some characters (lack of angularity of the flanks of the adventive lobe, lack of a tectiform ventrolateral saddle, lack of prorsiradiate juvenile aperhrre) that characterizes the gefirs Neogoniatites and separates it clearly from other genera such asDombarites and Platygoni- atites (Ftgs 4, 5). The two advancedsutural featureslisted above me thus intelpreted as homoplasies. Apparently, the evolutionary lineage leading to Neogoniatites became extinct near the Vis6an-Na- murian boundary. 188 Evolution of the Goniatitaceae: KORN

Paraglyphioceras -->Arwbergites. - In the shape of the juvenile aperture, all the other Vis6an genera within the family Goniatitidae differ remarkably from Goniatites and'Paraglyphioceras, for instance, species of Arnsbergites, Hibernicoceras, and Lusitanocelas possessin preadult stages a prorsiradiate aperture that is characteizedby a prominent ventrolateral projection. and a very shal.lowor absentventral sinus. The stratigraphicallyoldestof theseforms isArnsbergitesfalcatzs (Roemer, 1850), a speciesthat occurs immediately above Paraglyphi.ocerasradiatum and Goniatites spirtftr, but differs from the latter in the evolution of the inner whorls, as well as the much wider external lobe, higher median saddle,and subacuteventrolateral saddle.These modifications are so prominent that they cannot be explained by a gradual development from one of the youngest Gonfuttitesspecies. Arnsbergitesfalcatus sharesconspicuous ornamental features with Paraglyphioceras radiatum, but with the following differencesin intemal morphology: (l) n A. falcatu.r, more inner whorls are (2) evolute, thejuvenile apertural shapeis very dissimilar, prorsiradiatein A. falcatus,but rectiradiate in P. radiatum, (3) the flanks of the adventive lobe are more sinuous tn Arnsbergites, and (4) the external lobe is wider in A. falcatus (E lobe index = 0.72 in comparisonwith 0.65 in P radiatum). These modifications agreewith generaltrends in the evolution of the superfamily Goniatitaceae, and hence a close relationship between the two speciesis postulatedhere. Stratigraphically younger species of Arnsbergites are characterized by a weakening of the ornament and by the loss of the falcate pattem. Furthermore, the apertural outline changes: In A. falcatus, the growth lines run rectiradiate over flanks and venter, but in the younger A. gracilis and A. amsbergensls,they run in a rursiradiatedirection. [n this character,Hibernicoceras is very similar to Arnsbergites, but differs in the more rounded ventrolateral saddle and the loss of spiral lines on flanksandventer.Forthesereasons,Hibemicocerasisregardedasadescendantof Arnsbergites. Arnsbergites -->Lusitanoceras. - The phylogenetic positionof Lusitanoceraswithin the superfamily Goniatitaceaeis clear, although a direct ancestoris still unknown. The genus most probably derived from the spiratly omamented Amsbergites by developing an advanced sutural outline with sinuous flanks of the adventive lobe and a tectiform ventrolateral saddle.Conch morphology of the inner whorls is similar with open umbilicus and a prorsiradiate aperture, but a novelty is the triangularity, caused by deep constrictions, of the immature whorls. This character is maintained in almost all the descendantsof Lusitanoceras.

Lusitanoceras --> Proshumardites (Agathiceratidae). - Lusitanoceras is ancestral to Dom- barites. Species of Lusitanoceras are morphologically and stratigraphically close to Dombarites, much more so than to the genus Goniatites. Although Lusitanoceras and Dombarites have hardly been recordedfrom the samelocalities in succeedingstrata, it can be assumed,that Dombarites more or less directly succeededlasitanoceras. In their article on the relationships between the genusGoniatites and phylogenetically subsequent genera,Ruzhencev & Bogoslovskaya (1970) proposed a lineage initiating with Goniatites (in their interpretation; pxtTy Lusitanoceras as interpreted here), and leading to the PermianAgathicerasby passingthroughDombarites andProshumardites.Intheirmorphology, severaltrends are observable: (1) Increaseofmedian saddleheight: from 0.50 to 0.55 rtr Lusitanoceras, from 0.55 to 0.'75in Dombarites, and from 0.65 to 0.85 tn Proshumardites. (2) Development of tridentition of the adventive lobe sinuous flanks in Lu sit ano c e r as, angular flanks tn Dombarites, and tident in P ro shumardit es. (3) Loss of angularity ofthe ventuolateralsaddle. The concept ofRuzhencev & Bogoslovskaya (1970) in separatingbetween Pericleites (in their definition) and Proshumardites is not followed here. Apart from the difficulties in interpretation of the original Pericleites atticus (Renz, 1910) (Schindewolf 1939), the early Namurian forms possess ACTA PALAEONTOLOGICAPOLONICA (42) (2) 189

a moryhology which can be regardedas intermediate between Dombaites andProshumardites of lhe P. karpinskii group. In their conch geometry and ornament, the generaLusitanoceras and Dombarites are extremely similar, and in some casesinseparable. Most of the speciesof this lineage display an ornament with coarse spiral lirae, and rather often a falcate ornament is developed in adults. Triangular coiling of juvenile conchs is coflrmon in many species.There are also remarkable shifts in the maximum size ofthe conchs: 65 to 105 mmin Lusitanoceras,usually less than 80 mm, up to 150 mmin Dombarites, up to 65 mmin Proshumardites,but usually smaller, and up to 45 mminAgathiceras. Lusitanoceras -->Delepinoceras @elepinoceratidae). - Ruzhencev& Bogoslovskaya(1970) also proposed this lineage, which developed parallel to the one describedabove. The main transfor- ming feature is the extemal lobe, whose prongs becametridentate during phylogeny. This is already seenrn Platygoniatites, where they are already mammiform. The width of the external lobe [E lobe index) increasesremarkably, for example: 1.1 0 in the stratigraphically oldest form P. omniliratus, 1.30 in the stratigraphically younger forms P. molaris and P. superior, and 1.50in Delepinocerasbressoni. The height of the median saddle increased from 0.80 in Platygoniatites to more than 0.90 in Delepinoceras. In the other characters, the two genera are basically similar, both show spirally ornamented conchs which can be triangularly coiled in juveniles. Platygoniatites, however, is a giant among the late Early Carboniferous goniatites with a diameter attaining almost 200 mm, whereas Delepinoceras usually only reaches about 70 mm in diameter. Juveniles of the two genera are very similm to those of Lusitanoceras andDombarites,hence a close relationship can be proposed.The slight arching of the flanks of the adventive lobe in Platygoniatites may be an indication for an origin of Platygoni- atites from Lusitanoceras, rather than from Dombarites.

Palaeogeographical distribution of the Goniatitaceae

Late Vis6an and early Namrnian arnmonoids are known fiom numerous regions in Europe, North Africa, Asia, and North America. Knowledge of the ammonoid faunas from the variousregions is higtrly variable,comparisons are sometimes based on very incomplete information. Furthermore, the lack of any modern syntheseson the ammonoid assemblagesprevents exact correlation. Only from few regions(Tables 2, 3), more or less completefaunal successionsare known, allowing applicationof the stratigraphicalscheme introduced by Korn (1996) for the Rhenish Massif. Geographicaldistribution of the index specieson which the stratigraphicalscheme is basedis very irregular.Some of the species,.suchas Lusitanoceraspoststriatum, occur in almostall the regions;whereas others, such as Lyrogoniatites eisenbergensis, aregreatly restricted. Goniatiles. - Goniatites is frequently found in late Vis6an A sediments of Central and Westem Europe. Speciesfrom other regions can,by their suturalratios, be comparedwith the Europeanforms. G. americanus Gordon, 1971 from the lower Chainman Shale of the Confusion Range of Utah, for instance, displays a suture that in its ratios resemblesthat of G. hudsoni, and may have a similar shatigraphical age.Higher in the succession,G. multiliratus Gordon, 1962 occurs, with suturalratios such as in G. spirifer, indicating a late late Vis6an A age. r90 Evolution of the Goniatitaceae: KORN

Table 2. Documentationof the different ammonoidzones of the late Vis6an in the variousregions within the SubvariscanRealm. Shading indicates the presence ofammonoid-bearing strata (light shadingindicates questionableoccurrences), black dotsindicate the presenceof speciesof the Goniatitaceae,and opendots refer to questionableoccunences. Stratigraphical scheme after Korn (1996).Note that the distributionof faunasin particularregions is ratheruniform, differencesmay expressthe stateof knowledge.

E E;X - ; ? lE !r E ? ;= r i i5t a tEy .* e _'= d 3-s__ :+, -,:a: n =d _a; # r; E^: {7 i=:iS-fii ii $ Esd::E:=E{ipjk6_iS:Etga g"E;?-t:i ;' €u;;E 2^E';;;i i; ,? ;r !i siEt;;fli =:iX€;#: ss g;;? riv.;i1ii E,z;ir:; ;?=;E' s;H" pseudocoronulaZone a novalisZone ,E :!S!,""*2" .$ tiethensis Zone g eisenbergensisZone 6 ) poststriatumZone liiili!:o aao i::;=}: .f:;1.: f, e iiti.,: ,O. :;f::,:1;l!.j,; , , , P. suerlandenseZone l=, tr, :,,.:r,,.r,.: ,. =-.=:,+i:: .3 rotundum Lone 1,,;qr- .a::!:l ' =*=. :r trr::tr:.tl ,'e=:t.= -$ == spiraleZorc 1|-:;r;l r;1l:;r a:: = ,l.-: g grctcilisZone *;-l:'| r:,;:le ':: :: .:= :'tl' d ,1:r :::r:: : : ) JAlcatusZone ,-i-;,iiC .==,! t:...:,,'a:.,, tloo I spiriferZone :*:,::a:"4 l:.r:,r.a' o -,::',1': ra; ;ri:,r::'"' .r:a:.::14:a::.:.; ::. :: .F fmbriatus Zone *;r 1l '.1i t t:.: : t:: I :a l:::ll;::::rll:r.: -=. ra .a,. (ren8fno Lone i::;;:|i* i,.'::l: J a.. t 1::r .::!::1:;1., 5 .aialtatt!: :i.:; 9* globostriatusZone t::.: lraa:l:r!a!l : ::r::r::: .- :::i:ii:i::l!!tll:r'::rl::: ) ltudsoniZone ,* t1,;1 *:rr: -,:=;11L, ,, r.,4.:!:::t l: : grimmeri Zone

Occurrences of Goniatites in other regions are probably in time-equivalent strata. Kullmann (1961) described two spirally ornamented species from the Alba Formation of the Cantabrian Mountains, G. stenumbilicatus and G. globformis, and Bogoslovskaya (1966) describedtwo fatcate omamented speciesfrom the South Urals as G. crenfolcatus and G. sphaeroides.It is striking that in both occurencescompressed and globular morphotypes are represented,as in G. hudsoni, in which they are interpreted as an intraspecific variability. The four speciesdescribed by Kullmann (1961) and Bogoslovskaya (1966) display sutureswith dimensional ratios resembling thoseof G.fimbriatus and G. spiriftr, and hence, are probably of the same age. T|ne Goniatites -+ Hypergoniatites lineage. - Hypergoniatites is a frequent fossil in the latest Vis6an sedimentsof the South Urals (Ruzhencev & Bogoslovskaya 7970, 1971), Novaya Zemlya (Kusina & Yatskov 1988),central Asia (Pitinova 1975; Nikolaeva 1994, 1995),china (Liang & Wang 1991), and the CantabrianMountains (Kullmann 1961;Wagner-Gentis1980). Several iso- chronous species have been discriminated from these regions, but there is no indication of an occuffence of the genusin the SubvariscanRealm. Afler the decline of the genus Goniatites in the SubvariscanRealm at the end of the late Vis6an A, members of the family Goniatitidae with irivolute inner whorls occur only in a much higher stratigraphicallevel. In the N. suerlandenseZone of the Rhenish Massif, Goniatitella agricolaKom, ACTA PALAEONTOLOGICAPOLOMCA (42) (2) 191

rTiTI.:N active fold belts [ ::i:i:i:i:i:i:, I lr=j:i::--r:l inactive fold belts

FIII- cratons l::::::::::::itt:lcontinental

tl oceanicbasins

Fig. 6. Palaeogeographicalmap for the Namurian (after Tiegler 1989).

1988 is very cofilmon, a tiny goniatite with coarsespiral lines and a sutureline with juvenile outline. This specieshas not yet been reported from other regions. The Paraglyphioceras + Neogoniatites lineage. - The oldest genus of this lineage, Paragly- phioceras, is representedby several successive species in the Subvariscan Realm, known from southwestPortugal to the Lublin Coal Basin. Unambiguous reports of this ggnus from other regions do not exist. In contrastto this, the subsequentgen*S Neogoniatites is completely absentin Central Europe' It is a common faunal element in the latest Vis6an and earliest Namurian rocks. In North America the form is narned Goniatites kentuckiensrsMiller, 1889, in the Cantabrian Mountains Goniatites (Girtyoceras) palentinus Kullmann, 1961. In the Urals (Ruzhencev& Bogoslovskaya 1971) several species occur, and in South China Xainzalites xainzaensis Sheng, 1983, Goniatites platyformis t92 Evolution of the Goniatitaceae.' KORN

Table 3. Documentationof the differentammonoid zones of the late Vis6anand earlyNamurian in various palaeogeographicalregions. Shading indicates the ocurenceof ammonoid-bearingstrata (light shading indicatesquestionable occurrences), black dots indicatethe presenceof speciesof the Goniatitaceae,and open dots refer to questionableoccunences. Stratigraphical scheme after Ruzhencev& Bogoslovskaya (1971)and Korn (1996).Note that the distributionoffaunas containingspecies ofthe Goniatitaceaeshows a striking difference.In the SubvariscanRealm, the superfamilybecomes extinct before the end of the Vis6an,but in almostall the otherregions it continuesup into theNamurian. Note alsothat twice in the late Vis6an,atthe endof the lateVis6an A aswell asthe base of thelate Vis6an C. the Goniatitaceaehad a elobal distribution.

av .EE rQ r . .=5 xtrd '50 €a : i Ja: s -?. sa s G : Xa ; .aa H se !- < il K: H: 10 :, * i: si os ; oi >! :oe E t^": e - S; E: E' @9 : a: i! s-,iii ^" XF {: ct D;.9i- ;, de i6r .4,; .=i :_ :^ 9?ci i: 'F4 Hi s=*1 €; =q si i; Sa Q. 6. OE -- N: .!, 4i S: 6i {E X1 ;: &; zE 6i OA ZE za 6i >; X.: x; qi

Nm2a a f::,::.: a t:1;:r:,:: +: -::a: Nmlc .,..=i' a: ,..= .::-::: :::a: ::r:r:11::r::::: ::rl:: l:l :r: 14.1'l::al .e'.1=- -r: ::: :: :::::l:::: Nmib t-:1: ::::': l:::.: ::: l:.:l ::,il: i it t' t:::a at:r:::1.1 ra.::; ir:t ,,..' i:;.li:11: r:::l::r::: l::r,l::, l.l:!",.,. +,ta:t= :'a:::t:l' It :: :t r]14 -+:=,::'a r .:::: .::::l :: i::i::'. Late Vis6anC f ':::=' ta:aaa-- r- )::.:::: .:: r-::ra r::' it;l :ii ;:::l: a 'i:: -:::: iar::t ' ,::;:i;:l'i.i:ii:i' ::+ :'::: ::::]: ],4::riaa ::r::r :: r:l :::r::l if;=i-:-1,:1;:c Late Vis6anB Lliriillill::]r :il :i: :::r :- C= Jiriririr':::r:i a:,:,.,;-:ii:!::.i iEirl;'ir#::!;rC +:::::a.a: :l::l ra ra":;;l::: l r::, .:: l:: LateVis6an A ,iaa ataa::::.!. :.1 :+ :!::- :l :,t-:. :..: r:r;: i ':.+:" ::::l*:;:::; a:;*: :'::,t 3:::;.::t Early Vis6an

Sheng, 1983 and Neogoniatites xainzaensis Ruan, 1984 may belong to this genus. They occur in rather small numbers of individuals. In none of the regions, however, does the genus play any important role. Lusitanoceras. - Lusitanoceras is a genus with an almost global distribution. By separating different isochroneous species, palaeogeographical relationships may be discovered. Different speciesof lr.ritanoceras occur with in the various regions as fo11ows. ( 1) Central Europe. A handful of speciesfrom localities in the SubvariscanRealm, which can be assignedto Lusitanocera^r,have been introduced by several authors, and most of them are clearly jnnior synonyms of only one species,Lusitanoceras poststriatum (Briining, 1923).L. poststriatumis known from a large suite of individuals from the type area in the Rhenish Massif, and is represented in all growth stagesby three-dimensionally preservedspecimens (Korn 1988). Hence intraspecific variability is well known in this species,and formerly erectedspecies and subspeciesnames such as Neoglyphioceras cuboides Brtining, 1923, Glyphioceras granosum spiraloides Schmidt, 1925, and Glyphioceras granosum fotma jasterdoffiensis Patteisky, 1930 must be treatedas junior synonyms. The meaning of this is that in Central Europe L. poststriatum is the only speciesof that genus. It is unclear,however, as to whether I . poststriatum canbe separatedfrom L. granosaru(Portlock, 1843), a species based on one single deformed specimen from County Tyrone in keland. The single ACTA PALAEONTOLOGICA POLONICA (42) (2) r93

distinguishing characteris the number of spirals,which is lowerinL. granosuru.Unless new material of L. granosum is found, this problem will remain unsolved. (2) Eastem Central Europe. In the papers of Patteisky (1930), Knopp (1931, 1935: North Moravia), Korejwo & Teller (1967: Lower Silesia), arrdZakowa (1971: Holy Cross Mountains), mostly deformed material-has been described, not always allowing exact determination of these individuals. The best material derives from drills near Galgzice (Zakowa 1971: pl. 10: 1-3), which representsseveral morphotypes ofjuvenile Lusitanoceraspoststrintum (Briining, 1923), comparable with those known from the Rhenish Massif. It is unclear, however, whether all these specimens belongto L. poststriatum,becausesome of thempossessonly about 80 spirals (Zakowa 1971: pl. 10: 2), hencethey show affinities to L. granosum (Portlock, 1843). (3) South Central Europe. A report of poorly preservedgoniatites, which may belong to Lusita- noceras, from the Palaeozoicof Ndtsch can be found in Aigner & Heritsch (1929), but the presented figure allows no closer assignment. (4) British Isles. Although Lusitanoceras occurs quite frequently in localities in North England and heland, representation in the literature is rather scarce. Figured specimens have normally been assignedto L. granosum (Portlock, 1843). The high number of spiral lines on the specimenfigured by Moore (1936: pl. 3: 1, 10), however, suggests that L. poststriatum is also representedthere. Records of Lusitanoceras from northwestem keland (Brandon & Hodson 1984) have not been figured yet, so their specific assignmentmust remain uncertain. The type of L granosum (Portlock, 1843), refigured by Moore (1936: pl. 3: 6) and Gordon (1965: pl. 18: 7), only shows 80 spiral lines from umbilicus to umbilicus (in contrast to L. poststriatum which always possessesmore than 100), and an almost punctiform umbilicus at 36 mm diameter (unlike l. poststrintum with an umbilicus that amounts 0.15 of the conch diameter). (5) Southwest Portugal. In this region, two sympatric speciesof Lusitanoceras occur, L. postria- tum and L. alganiense Pereira de Sousa, 1923, the type species of the genus. Both are clearly separatedby ornament and conch characters,and are not connectedby intermediates(Kom 1997).I. algarviense is unique becauseof its falcate adult omament and by its narrow umbilicus. A similar form is not known, the only one comparable is t. granofalcatum (Kullmann, 1961) from the CantabrianMountains, with a much wider umbilicus. (6) Cantabrian Mountains. Species which can be attributed to l;usitanoceras were desribed and figured by Kullmann (1961) and Wagner-Gentis (1963, 1980). The first of these species is L granofolcatum (Kullmann, 1961), characterizedby a strong falcate ornament in the adult stageover 70 mm diameter, and by about 80 coarse spiral lines. The rather widely umbilicate holotype of the species displays fragments of the suture line, with an adventive lobe lacking angulm sinuosity ofits flanks and a median saddle with approximately half of the height of the ventrolateral saddle, hence an attribution to the genus Lusitanoceras is much more likely thanto Dombarites. A number of forms with strong longitudinal ornament have been described by Wagner-Gentis (1963, 1980), who referred them to Goniatites granosus Portlock, 1843, Dombarites acicularis (Pareyn, 196l), Dombarites acicularis (Pareyn, 1961) early form, Dombarites granofalcatzs (Kull- mann, 1961), Dombarites canitna Wagner-Gentis,1980, Revilloceras barruelense (Wagner-Gentis, 1963), andRevilloceras globosum Wagner-Gentis,1980. They have been collected in the section near Revilta, mostly in stratigraphical otder. Dombarites acicularis (Pareyn, 1961) early form andDombarites canitnaWagner-Gentis,1980 arehere included in Lusitanoceras granofalcaturu (Kullmann, 1961). (7) North Africa. Pareyn (1961) describedrather small specimenswhich he classified as Goni- atites-Mesoglyphioceras-granosasPortlock, G oniatites-Mesoglyphioceras-granosusvar. dilatalta, and Goniatit es-Meso glyphioceras-grano susv ar. aciculare. Of these, the two variants introduced by hirn are based on specimens smaller that 14 mm diameter, hence very difficult to compare with the material from other regions. The sutural drawings suggest that var. acicularehas to be placed in Dombarites, var. dilatalta tn Platygoniatites, and only the first in Lusitanoceras. Unfortunately, the conch and ornament characteristicsofthese specimensdo not allow a real comparison with speciesknown from other regions. Besidesthis, Pareyn (1961) figured 194 Evolution of the Goniatitaceae.. KORN

rather well preserved adult specimens of his Goniatites striatus kentuckiensis Miller, which have since been regarded by Ruzhencev & Bogoslovskaya (1970,1971) as Dombarites falcatoides Ruzhencev & Bogoslovskay a, 7970. (8) South Urals. Lusitanoceras orientalls (Librovitch, l94O) was described from the Salcnara River (Bogoslovskaya 1966).These well preservedspecimens can be compmed with specimensfrom Central Europe, showing that the extemal lobe in the material from the Uials is narrower. (9) Kazakhstan.Librovitch (1940) reported the two new speciesLusitanoceras oientalis and Lusitanoceras irtyshensis fromthe vicinity of the Lake Taran-Kul in the Priirtysh region. Both species are, in their morphologies, very close to L. poststriatun, but differ in their niurower umbilicus and more shong curved growth lines. (10) Novaya Zemlya. Goniatites pola;rls Kusina, 1987 is, according to its suture line, a Lusita_ noceras speciesthat differs in the absenceof triangular inner whorls from most of the other species of the genus. ( I 1) China. Specimensof Lusitanocera^shave been describedfrom severalregions of China, but most of the material is too poorly preserved for exact determination. They derive from Xinjiang (Liang & Wang 1991) and Xizang (Sheng 1983). (12) American Midcontinent. Lusitanoceras granosum (Portlock, 1843) has been reported from the Batesville Sandstoneand Ruddel Shaleof severallocalities in Arkansas (Gordon 1965).Revision of this material is required to confirm the identifications given by Gordon (1965). -+ The Dombariles Poshumardites lineage. - In contrast to the global distribution of Lusita- noceras, the subsequent genera of this lineage are restricted in their occurrence, and not a single record from the SubvariscanRealm exists. Dombarites is rather widely distributed, and is known from many different regions, e.g. North America (Gordon 1965; Drahovzal & euinn 1972), the cantabrian Mountains ('wagner-Gentis1963), North Africa @areyn 1961), Novaya Zemlya (Libro- vitch et al. 1993); the south Urals (Ruzhencev & Bogoslovskaya 1970, r97l), central Asia (Nikolaeva 1994,1995), and china (Ruan 1981; Sheng 1983; Gao 19g3; yang 1986; Liang & wang 1991). In some of theregions Dombaiter occurs in tremendousnumbers of individuals, forming thi major component of the faunas. The distribution of Proshumardites is very similar to that of Dombarites (see references for Dombarites), but the species does not occur in the Namurian of North America. Advanced species areknown fromthe BloydianandAtokan of Texas,Arkansas, andOklahoma(Plummer& Sc ottl937: Gordon 1965). -+ The Platygoniatiles Dehpinoceras lineage. - Platygoniatites and,Delepinoceras arc, in contrast to Dombarites and Proshumardites, fw more restricted in their distribution (see references for that gents). Platygoniatites is known from the Cantabrian Mountains, North Africa (most probably), the South Urals, central Asia, china (Xinjiang, Guangxi, and Ningxia) and Nevada, but absent in the American Midcont;nent. Delepinoceras isknown from the regions listed for platygoniatites,but also occurs in Arkansas and Oklahoma. There is no evidence ofthese genera from the Subvariscan.

Sea-level changes and goniatite biogeography

The distribution pattern of the speciesof the Goniatitaceaeis remarkablefor of several reasons: (1) The continuoussuccession of speciesofthe late Vis6anA andlate Vis6anB in the SubvariscanRealm is in contrastto the discontinuousdistribution of the superfamily in otherregions. (2) Twice in the late Vis6an,the superfamilyis globally distributed(at the end of the late Vis6anA and at the baseof the late Vis6anC). At the endof the late Vis6anA, ACTA PALAEONTOLOGICA POLONICA (42) (2) 195 the genus Goniatitesis, in the various regions,represented by well distinguishable species,and at the baseof the late Vis6anC, Lusitanocerashas a world-wide distribution with morphologically extremely similar species. (3) The superfamilybecomes, towards the end of the Vis6an,suddenly extinct in the Subvariscan,but continuesup to the late Namurianin other regions,especially in the SouthUrals andCentral Asia. The constant presenceof the superfamily in the SubvariscanKulm series may reflect a stability of the enviroffnent over a long duration. The suddendecline of the Goniatitaceaein the latest Vis6an sediments of the Subvariscan is a mystery. No environmentalchange which canbe regardedas responsible for this extinctioncan be seen in the sediments. Similarly, no plausible explanation for the absenceof the superfamilyin otherregions can be proposed. It seems significant that there were two main transgressiveepochs in the late Vis6an. The first one resulted in the widely distributed Crenistria Limestone (Herbig 1994),and'the second one led to depositionof theActinopteria Shales.Both immediately precedethe intervals from which goniatitids are known in world-wide disfibu- tion. These transgressionsprobably led to mixing of former$ allopatric populations, resulting in co-occurrencesof morphologically very similar specieswith an equal state of sutural development,distinguishable only becauseof slightly different ornaments.

Conclusions

Results of cladistic and stratopheneticanalyses of a gtoup of Palaeozoicammonoids are closelysimilar. The superfamily Goniatitaceaeis representedby a few independentevolutionary lineages.According to the distribution of conch shape,ornament, and sutural characters,it canbe statedthat: (1) The superfamily is a monophyletic taxon, characteize'dby one synapomorphy, the ontogenetic change of the external lobe in the suture line from a Y-shaped to a V-shapedoutline. (2) Of the three traditional families within the superfamily, the Goniatitidae is a paraphylum,but the Agathiceratidaeand Delepinoceratidaeare monophyla. (3) From the results of a biometrical analysis of sutural elements, tends in the sutural evolution of the superfamily can be observed,allowing a comparisonwith the modification of other characterssuch as conch parameters.Such a comparisonallows a discriminationof derivedhomologies from homoplasies. (4) According to the relatively low number of informative characters, several unresolvedtrichotomies are expressed in the cladogram. During the late Vis6an and early Namurian, two faunal provinces are easily distinguishableon the basisof the distributionof the Goniatitaceae. (1) The first provincespans from the Lublin Coal Basin in the southeastover the RhenishMassif towardskeland in the northwest,and also includesSouth Portugal. Typical elementsofthe late Vis6anfaunas are a successionofgenera and species ofthe Goniatitidae.In this province,Lusitanoceras is the youngestgenus of the Goniatitaceae at the baseof the late Vis6anC. 196 Evolution of the Goniatiraceaz.'KORN

(2) The secondprovince includes the South Urals as well as the Cantabrian Mountainsof Europe,but alsothe occurrencesin North America,Noth Africa, Central Asia and China can be regardedas belongingto this province.From theseregions, Paraglyphioceras,Arnsbergites, and Hibernicoceras have not beenreported. Of the Goniatitaceae,Hypergoniatites, Neogoniatites, and Dombarites occtt in bedswhich areyounger than the Lusitanoceraslevel, and clearly dominatethe fauna. In the late Vis6anand Namurian,there are only very few speciesof the Goniatita- ceae,which occurin both provinces. The degreeof difference betweentwo faunal provinces is variable during the late .Vis6an.In two short epochs,almost cosmopolitanfaunas occur after major trans- gressiveperiods: (1) Towardsthe end of the late Vis6anA with a global distributionof the genus Goniatiteswith clearly distinguishablespecies, and (2) atrhe baseof the late Vis6anC with globally distributed,morphologically very similar speciesof Lusitanoceras. The two phasesof faunal homogenizationof the two provincesis explainedby global transgressions,which led to mixing of formerly allopatric faunas.

Acknowledgements

This study is part of a Ph.D. thesis submitted to the Faculty of the tnstitut und Museum flir Ggologie und Paliiontologie der Eberhard-Karls-Universitiit Tiibingen. It was achieved using the Database System GONIAI, founded by the Deutsche Forschungsgemeinschaft. I am indebted to Prof. Jerzy Dzik (Warsaw), as well as Dr. David Gower, Prof. Wolf-Ernst Reif and Rainer Schoch (all Tiibingen) for many helpful discussions and for reviewing the typescript.

References

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Ewolucja goniatyt6w Goniatitaceae oraz biogeogralia wizenu i namuru

DIETERKORN

Streszczenie

Przy zastosowaniumetod analizy kladystycznej i stratofenetycznejrozpoznano linie ewolucyjne w obrgbiekarboriskiej nadrodziny goniat5rt6wGoniatitaceae. Obie metody daty podobne rezultaty. W p6Znym wizenie i w namurze A prowincje goniatytowe mog4 byi zdefiniowane na podstawierozmieszczeniaposzczeg6lnychlinii Goniatita- ceae.Pierwsza prowincja odpowiada obszarowi subwaryscyjskiemu, gdzie goniatyty tej nadrodziny wymarly w poblizu granicy wizen-namur. Druga prowincja obejmuje resztgobszar6w wystgpowania nadrodziny, a wigc np. potudniowy Ural, Srodkowa Azjg i Amerykg P6lnocn4,gdzie Goniatitaceaeprzetrwaty do Namuru A. Goniatitaceae mialy Swiatowerozprzestzenienie podczas dw6ch tr6tkich epok wielkich transgresji w wizenie,kt6re ulatwialy migracje.Pierwsza z tych epokodpowiada p6inemawize- nowi A, kiedy rodzaj Goniatites(Zony G. fimbriatus i G. spiriftr) mial Swiatowe rozprzestrzenienie.Druga epoka przypadla na pocz4tek p6lnego wizenu C, kiedy to Swiatowerozprzestrzenienie mialrodzaj Lustinoceras(ZonaL. poststriatum).