Acta Geologica Polonica, Vol. 49 (1999), No.2, pp. 159-173 406

/U S UNES 0 I

Permian corals of the Cordilleran-Arctic-Uralian Realm

JERZY FEDOROWSKIl, E. WAYNE BAMBER2 & CALVIN H. STEVENS3

1 Institute of Geology, Adam Mickiewicz University, Makow Polnych 16, PL-61-606 Poznan, Poland E-mail: [email protected] rd 2 Geological Survey of Canada (Calgary), 3303-33 Street N.W., Calgary, Alberta T2L 2A7, Canada E-mail: [email protected] 3 Department of Geology, San Jose University, San Jose, California 95192, USA E-mail: [email protected]

ABSTRACT:

FEDOROWSKI, J., BAMBER, E.W. & STEVENS, C.H. corals of the Cordilleran-Arctic-Uralian Realm. Acta Geologica Polonica, 49 (2),159-173. Warszawa.

Permian rugose corals of the Cordilleran-Arctic-Uralian Realm are abundant in shallow-water carbon­ ates along the northwestern and western margin of , from the area in , through the Svalbard Archipelago and arctic and western North America, to Bolivia and Peru. The colo­ nial forms are of particular interest for the biostratigraphy and reconstruction of the paleogeography of this extensive region. A revision of the systematics of these corals has shown that, although important differences exist between the assemblages in the areas listcd, the faunas are recognizable throughout the entire realm. Almost all of the faunas in the realm, on cratonal Pangaea, are ( to ) in age although younger faunas occur in rocks of terranes subsequently accreted to North America. The cratonal faunas show a general trend from a predominance of relatively simple, fasciculate species with open axial areas or weakly developed axial structures in the lowest Permian, to younger, more complex fasciculate and massive species with a variety of morphological elements in their axial structures and dissepimentaria. Suppression of the walls occurs commonly in the youngest faunas. As a result of the northward movement of Pangaea into cooler watcrs, colonial rugose corals wcre extermi­ nated from the more northerly arcas by early to mid-Artinskian time, but persisted throughout the Artinskian and possibly into the and carly in the cratonic successions of the western U.S.A. Similarities betwccn the coral faunas of cratonal North America and the western allochthonous terranes indicate that faunal interchange occurred between these various coral faunas dur­ ing the Cisuralian. The only colonial corals recovered from to rocks in this region are waagenophyllid corals of tethyan affinity from the Cache Crcek and Quesnellia terranes of British Columbia and the Hayfork and Eastern Klamath terranes of California. Key words: Permian, Rugosa, Western Pangea, Biogeography.

INTRODUCTION distribution of marine faunas and affected their subsequent phylogeny. Prior to its closing in Changes in world geography during Late time, there surely was free communica­ time had profound effects on the tion through a -Moscovian seaway 160 JERZY FEDOROWSKI, E. WAYNE BAMBER & CALVIN H. STEVENS connecting the Prae-Tethys to the Uralian Ocean The effect of this factor was proportional to the (FEDOROWSKI 1981, RODRIQUEZ & ai. 1986, level of sensitivity of a given taxon to environ­ GOLONKA & ai. 1994). The possibility that such a mental conditions, its genetic potential for seaway leading to the southern Urals Ocean per­ change, and its ability to disperse. Thus, the sisted through the Asselian or even into the stenotypic, sessile benthic faunas were most has been postulated by some authors strongly influenced, and these certainly include of palaeogeographic maps (e.g., GOLONKA & ai. the rugose corals. The response of the latter to 1994), but distribution of facies-sensitive rugose geographic and environmental changes was gov­ corals indicates a pre-Permian closure. erned partly by differences between the sensitivi­ Regardless of the exact timing of the closure, ty of individual morphotypes - a factor often there is general acceptance that, beginning in Late omitted from palaeoenvironmental analyses. In Carboniferous time, Pangaea existed as a solid general, the environmentally most resistant soli­ landmass with substantial differences in the tary, non dissepimental rugose corals may be dis­ marine faunas living on its eastern and western tinguished from those with other growth forms. shelves. The differences between the rugose coral Normal salinity was the only general restricting faunas in these regions are so striking that factor for the former, whereas other factors gov­ FEDOROWSKI (1986) introduced the idea of two erned the second group. Within the second group, independent realms: the Tethys Realm and the dissepimental solitary corals and fasciculate colo­ Cordilleran-Arctic-Uralian (CAU) Realm. (Text­ nial taxa apparently were less sensitive to their fig. 1) Most of the latter realm was earlier referred environment than were the massive colonial to as the Thysanophyllum Belt by STEVENS forms. In this paper, all of these morphological (1982). The existence of two realms is accepted in groups will be briefly analysed to provide a com­ the present paper because it has since been sub­ prehensive historical overview of rugose corals in stantiated by new data and interpretations of both the CAU Realm. the palaeogeography and the systematics and dis­ In our analysis we use two Carboniferous sub­ tribution of rugose corals. systems and the three Permian (Cisuralian, The unique geography of the world during the Guadalupian, Lopingian) that have been recently " , Cisuralian was a primary factor influencing rela­ recognized by the Subcommission on Permian tionships between invertebrate faunas of that age. Stratigraphy (e.g. lIN YUGAN 1996). We omit

------.---. ------...... , "",.c:.. ... so-----__ ------...... / ...... /' c:::::J / y --"::"Cp\S " ~30------BC' -~ (?~------"" " JR I ~ ~ '\ 1- 0------ ---, \ D I \ / , , 30 ------,1 , / ...... /' ...... ,/ "'<:-...... 60 - ---;;I"""'" ------I Fig. 1. Palaeogeographic map of Pangea based on ZIEGLER & al. 1996, Early Permian (Sakmarian). BC - British Columbia, ,j CA - Canada, CC - Central Cordillera, CR - Canadian Rocky Mountains, S - Spitsbergen, SA - South America, T - SW , Y - Yukon Territory PERMIAN CORALS 161 from our discussion all Lower Carboniferous and sented family Bothrophyllidae FOMICHEV, 1953. most of the Upper Carboniferous strata and coral Representatives of this family dominate in the faunas, except where the latter have a direct bear­ CAU Realm, but some are present in the Tethys ing on the history and distribution of Permian fau­ Realm as well, where they persisted longer than in nas. Unfortunately, there are difficulties in corre­ the CAU Realm. For instance Arctophyllum is lating upper Cisuralian (Leonardian/Kungurian) known to occur in Vietnam into the Guadalupian strata of southern North America with the inter­ and Wordian Stages. In the CAU Realm, nationally accepted standard in the Ural Permian representatives of this family were most Mountains. For the remainder of the Cisuralian abundant in the Asselian. Arctophyllum, succession, however, the story of rugose coral Bothrophyllum, Siedleckia, Timania, and development and extinction in the CAU Realm is Caninophyllum?, which originated in the reasonably clear. Carboniferous, continued to occur throughout most of the realm and Fedorowskiphyllum appeared and spread from the Northern Urals to ROOTS OF PERMIAN RUGOSE CORALS California. Some of these genera continued OF THE CORDILLERAN-ARCTIC-URALIAN through the Sakmarian, but none survived beyond REALM the earliest Artinskian Burtsevsky Horizon (= Pseudofusulina concavuta Biozone). They never Almost all main stocks of Permian rugose created their own communities in the Cisuralian corals contain three categories of taxa: long-last­ and did not dominate over colonial morphotypes ing taxa that are rooted in Carboniferous or older - a character clearly distinguishing Asselian coral strata and are common to both Permian realms; faunas from those of the Gzhelian. taxa clearly restricted to one of the realms, lranophyllum, which is a dissepimental, soli­ although not necessarily restricted to Permian tary representative of the Wentzelellinae', is one strata; and taxa morphologically similar in both of the most diversified, widespread and abundant realms, but probably homeomorphic. The latter genera in the Tethys Realm. It has never been category includes mostly species and genera, found on the western shelves of Pangaea and whereas the former two groups are at the family therefore provides a reliable marker for distin­ or suborder level. This division often crosses gen­ guishing between the two realms. eral taxonomic boundaries and corresponds only Permian remnants of the families Petalaxidae partly to growth forms and/or ecotypes (i.e., the FOMICHEV, 1953 and Geyerophyllidae MINATO, so called "Faunas" of HILL 1938-1941). 1955 include solitary dissepimental and colonial Perhaps all of the "horn corals", or corals - both fasciculate and massive. Following "Cyathaxonia Fauna" of HILL (1938-1941) - i.e., tradition, we distinguish between those two fami­ nondissepimental taxa at the genus and higher lies in this paper, but they are most probably syn­ taxonomic level - are rooted in strata older than onyms. The very few representatives that occur in Permian. Also, "horn corals" constitute the most the Asselian of the CAU Realm are obvious widely distributed and longest lasting morpho­ holdovers from the Carboniferous Period. The type because of their great environmental oppor­ only genus that may be considered typical for the tunism and, perhaps, largest genetic plasticity. As Asselian-Sakmarian, rather than Gzhelian, is a result, "horn corals", as a group, are of little use Lytvophyllum, although it possibly appeared very for palaeogeographical and palaeoecological late in the Carboniferous. Representatives of analysis unless considered individually as low these families were much more abundant and also level taxa. Many of them belong to deep, cold lived longer in the Tethys Realm than in the CAU water faunas and are important environmental Realm. indicators. However, we will not consider this Among the colonial forms rooted in the group further, except in our discussion of the mid­ Carboniferous, the family Lithostrotionidae is rep­ Permian faunas. resented by the single, but geographically wide­ Almost all of the dissepimental solitary spread genus Tschussovskenia. Its taxonomic Rugosa - i.e., the "Caninia-Clisiophyllum relationships are discussed by us elsewhere Fauna" of HILL (1938-1941) - are rooted in the (FEDOROWSKI & ai., in prep.). Our analysis shows, Carboniferous. This is especially true for the most however, that it can be linked through intermedi­ widely distributed and the most frequently repre- ate genera to the Visean genus Diphyphyllum. 162 JERZY FEDOROWSKI, E. WAYNE BAMBER & CALVIN H. STEVENS

Representatives of Tschussovskenia are known to dissepimental corals, and the remaining genera are occur in Asselian-Sakmarian strata from the Ural massive colonial forms. The relative proportion of Mountains through Svalbard and the Canadian these different growth forms reflects the composi­ Arctic Archipelago to California and Nevada. We tion of the entire fauna, briefly discussed later. cannot exclude their presence in South China and Although it is possible, as mentioned above, Indochina, although those tethyan corals assigned that homeomorphic genera may be present in both to Tschussovskenia may be distinct at the genus realms, this cannot be definitely proven, because level (FONTAINE 1961, Wu 1962). current data are incomplete. It is probable, that The stratigraphically and geographically most Japanese species such as Durhamina kitakamien­ important families - the Durhaminidae and sis MINATO & KATO, 1965 are not durhaminids "Heritschioididae" of the CAU Realm and the because they possess clinotabellae (FEDOROWSKI Waagenophyllidae and Yatsengiinae of the Tethys 1997, p. 101), and the North American Realm - are rooted in the Late Carboniferous as Yabeiphyllum rossi MINATO & KATO, 1965 is well. However, unlike the lineages discussed included by us (FEDOROWSKI & aZ., in prep.) in the above, those four families were separated geo­ Durhaminidae. Also, some tethyan bothrophyllids graphically from the beginning of their develop­ and caniniids do not necessarily belong to the ment, thus creating the main basis for the distinc­ same genus as those in the CAU Realm. The same tion between realms. The solitary, dissepimental may be true for the genus Paracaninia. Abundant Amandophyllum may perhaps be the only genus Gzhelian and Kazanian hom corals from the East present in both realms. It is rooted low in the European Platform were included in that Chinese Carboniferous if its identification by FEDOROWSKI genus by WEYER (1982) and WEYER & lUNA (1971) in the Upper Visean of Poland is accepted. (1979), but our knowledge of the type material for The development and relationships of Paracaninia is only fragmentary, so we may well Durhaminidae and "Heritschioididae" will be dis­ be dealing with homeomorphs. cussed later. Further discussion of the W aagenophy llidae and Yatsengiinae is omitted because this paper deals only with the rugose corals COMPARISON OF RUGOSE CORAL of the CAU Realm. Throughout this paper, the FAUNAS FROM AUTOCHTHONOUS name Heritschioides and its derivatives appear in DEPOSITS WITH THOSE inverted commas, because our recent re-investiga­ OF ALLOCHTHONOUS TERRANES tion (FEDOROWSKI & aZ., in prep.) of H. coZumbicum SMITH, 1935, the type species of the Permian rugose coral faunas of western North genus, has shown it to differ in morphology and age America occupied the western cratonic margin of from all described Permian species currently Pangaea and the margins of more westerly island assigned to that genus in the literature. chains and platforms accreted to North America as Except for the Diffingiina FEDORowSKI, 1986 allochthonous terranes during the . The new taxa at the family or higher level have not reader is referred to MONGER & NOKLEBERG (1996) been recognized from Permian strata of the CAU for a summary of the interrelationships and tecton­ Realm, but some exclusively Permian genera such ic history of these terranes. Cisuralian colonial as Allotropiochisma, AssimuZia, Euryphyllum, rugose coral faunas occur along the margin of cra­ LeonardophyZum, Lophbillidium (subgenus), tonic Pangaea and in the following allochthonous LytvoZasma, Fedorowskiphyllum, Hornsundia, terranes: Northern Sierra, Eastern Klamath, Langenheimia, KZeopatrina, Permastraea, Grindstone, Quesnellia, Stikinia, and Wrangellia. ProtoZonsdaZeiastraea, Stikinastraea, and Guadalupian colonial corals are known from the StyZastraea, as well as many species have already Eastern Klamath, Hayfork, Quesnellia and Cache been described. In addition, several new genera Creek terranes, but are unknown from the will be introduced by FEDOROWSKI & aZ. (in prep.). Pangaean margin of North America, except for Most of the colonial genera and all of those to be new genera of possible Roadian age, recently dis­ introduced by us belong to the Durhaminidae and covered in Nevada. Lopingian corals are known "Heritschioididae", the most abundant and diversi­ only from the Hayfork Terrane, where rare, early fied Cisuralian rugose coral families in the CAU colonial waagenophyllids occur. realm. The first six genera listed above are solitary, The Permian positions of the allochthonous ter­ non-dissepimental corals, the next two are solitary, ranes relative to North America are uncertain. PERMIAN CORALS 163

Paleomagnetic data suggest that many of these ter­ tonal margin of western-central U.S.A. In the ranes have been displaced northward (MONGER & Eastern Klamath Terrane (STEVENS & al. 1987) NOKLEBERG 1996), but the longitudinal placement these Tethyan corals occur stratigraphically is more difficult. STEVENS & al. (1990) and above faunas of CAU aspect, whereas in the BELASKY (1994) used data on modem corals and Quesnellia terrane (NELSON & NELSON 1985) the Permian colonial corals in different manners to tethyan corals occur in structurally complex suc­ postulate that some of the terranes bearing cessions and are associated with fault blocks con­ Cisuralian corals originated a considerable dis­ taining older, non-tethyan faunas. The reason for tance west of the Pangaean margin, but still well the presence of tethyan corals and other tethyan within the eastern part of the Panthalassan Ocean. fossils in North America is not known. They Their conclusions are based on differences could have arrived by migration across between the allochthonous faunas and those of cra­ Panthalassa using mid-ocean islands as dispersal tonal North America, and the complete lack of sim­ points, by transport on moving ocean plates, or by ilarity between western North American Cisuralian a combination of these two methods. corals and coeval tethyan faunas of Asia. The new, comprehensive review of North American Permian colonial faunas, by F'EDOROWSKI & al. (in prep.), COMPOSITION OF THE RUGOSE CORAL has shown that the cratonic faunas are closely relat­ FAUNAS ed to each other and are similar to the faunas of the arc terranes to the west. Cisuralian The Guadalupian colonial coral faunas found in the Cache Creek, Eastern Klamath and Our knowledge of the composition of a fossil Quesnellia allochthonous terranes are entirely fauna depends, in many instances, on the state of tethyan in aspect. All other colonial rugose corals its preservation, the intensity with which it has of the CAU Realm had become extinct near the been collected and investigated, and the approach end of the Cisuralian, with the possible exception to the taxonomy, as demonstrated by the follow­ of locally occurring Roadian? species on the cra- ing three examples: 1. In almost two hundred of investiga­ tion of the central European Permian, no more ARCTIC OCEAN than two, and perhaps only a single coral species belonging to Calophyllum have been discovered. Thus, there is practically no chance of collecting an abundant coral fauna. 2. The mainly solitary, nondissepimental coral fauna from the Cisuralian and Guadalupian of southwestern Texas, which is the richest in the realm, was unknown until it was described in papers by Ross & Ross (1962, 1963) and FEDOROWSKI (1986,1987). Thus, in earlier analy­ ses that area would have been considered almost '---'- barren of corals. However, the richest, Roadian part of the fauna remains undescribed, making our analysis incomplete because we cannot include taxa from that part of the collection, although one of us (IF) has several thousand specimens at his disposal. 3. The morphology of colonial taxa of the fam­ Fig. 2. Terrane map of northwestern North America showing ilies Durhaminidae and "Heritschioididae" is terranes containing Permian rugose corals. Terranes: extremely variable, and the type material of some CC - Cache Creek, EK - Eastern Klamath, G - Grindstone, of the more common species has been inade­ H - Hayfork, NS - Northern Sierra, QS - southern Quesnellia, quately studied. This has resulted in the introduc­ including Harper Ranch subterrane, S - Stikinia, tion of many species, which we consider to be Wa - Wrangellia synonyms. Our conclusion is based on a review of 164 JERZY FEDOROWSKI, E. WAYNE BAMBER & CALVIN H. STEVENS many of the type specimens and our analysis of corals to taxa of southwestern Texas, but the docu­ the variability shown by all described colonial mentation of their occurrence by Kossov AY A species from Permian strata of the CAU Realm (1996) is a welcome addition to our knowledge of (FEDOROWSKI & al., in prep.). Thus, the actual the Russian coral fauna. number of species recognized by us is drastically The succession of colonial taxa in the Russian reduced, compared to the number of species sections, as reconstructed by KOSSOVAYA (1996, names that have been introduced into the litera­ fig. 10) follows a very regular pattern, leading from ture. For instance, re-investigation of the holotype domination by fasciculate morphotypes in the of Stylastraea inconferta LONSDALE, 1845 lower Asselian Sphaeroschwagerina vulgaris-So showed that species to be synonymous with 13 fusiformis Biozone to domination by massive ceri­ subsequently named species. A quantitative oid forms higher in the Asselian and into the analysis based on such names taken uncritically Sakmarian. These are followed by the youngest from the literature would have been considerably fauna in the area, consisting mainly of cerioid­ different from the analysis we propose. aphroid colonies, from the middle Sakmarian Although our analysis is strongly handicapped Pseudofusulina verneuilli-P. uralica Biozone to by factors such as those listed above, we have the Lower Artinskian Pseudofusulina concavuta compensated for this to some extent by the use of Biozone. Our analysis (FEDOROWSKI & al., in unpublished data from undescribed faunas in prep.) shows that several species and genera dis­ western U.S.A. and western and northern Canada. tinguished within those faunas by earlier Russian authors (e.g. STUCKENBERG 1895; DOBROLYUBOVA 1936a, b; SOSHKINA & al. 1941) and subsequently Eastern European Platform, Ural Mountains repeated by SIMAKOVA (1986) and KOSSOVAYA and northern Timan (1996, 1997) are synonyms. Without further dis­ cussion in this paper, we agree with WILSON (1998) KOSSOVAYA (1996) summarized the distribu­ that "Heritschioides" is absent from the Russian tion data for uppermost Carboniferous and part of the CAU Realm. Also, we restrict the occur­ Cisuralian rugose corals from the eastern part of rence of Thysanophyllum to Lower Carboniferous I the Eastern European Platform, the Urals and strata and consider Borealephyllum KOSSOVA YA, 1 northern Timan and proposed a coral zonation cal­ 1997 and Protowentzelella PORFIRIEV, 1941 to be ibrated with the existing fusulinacean zonal junior synonyms of Stylastraea LONSDALE, 1845. scheme for the area. Also, she presented correla­ Thus, the massive colonial corals from the Urals tions within the CAU Realm, based on coral faunas and northern Timan are in fact restricted to (KOSSOVAYA 1996, fig. 11). Our distribution data Kleopatrina, Stylastraea, Pseudocystophora, do not confirm her conclusions concerning Protolonsdaleiastraea, Permastraea and at least Vestspitsbergen, Eastern Nevada, the Klamath two new genera presently being described by us Mountains in California, and southwestern Texas, (FEDOROWSKI & al., in prep.). The fasciculate but we have adopted her biostratigraphic analysis genus Heintzella is also present. Other fasciculate for the Russian faunas. It confirms the succession colonies assigned by Russian authors to either of rugose coral faunules through the Corwenia or "Heritschioides" and to Fergano­ Carboniferous/Permian transition, previously out­ phyllum require further study to be fully under­ lined by FEDOROWSKI (1981, 1986) - i.e., the stood. Thus, we do not recognize them in this absence of colonial corals except for some paper, but we confirm the occurrence of Petalaxidae high in the Carboniferous; domination Lytvophyllum and Tschussovskenia. I by Bothrophyllidae in Gzhelian strata; rapid disap­ Artinskian strata of the Eastern European .~ pearance of that family (except for the genus Platform, northern Timan, and especially the Ural Timania) early in the Permian and their replace­ Mountains yield fairly abundant nondissepimental ment by colonial taxa. KOSSOVAYA (1996) supple­ rugose coral faunas. Three cerioid-aphroid mented the data on Permian faunules of the species, included by KOSSOVAYA (1996) in the Russian part of the CAU Realm with information genus Protolonsdaleiastraea are present in the on nondissepimental solitary corals, almost Lower Artinskian Pseudofusulina concavuta unknown from earlier studies. Following Biozone of the Timan Mountains. SOSHKINA & al. FEDOROWSKI (1997) we are not certain of the (1941) summarized earlier descriptions and assignments of most of these nondissepimental described some new Artinskian taxa. Many identi- PERMIAN CORALS 165 fications of genera included in that paper do not Solitary nondissepimental taxa have not yet been conform to modern taxonomic practices and need found. Also, the genus "Heritschioides", which is careful revision. Also, the precise stratigraphic common in North America, is absent from this positions of most species are unknown. The posi­ part of Vestspitsbergen, as it is from the Urals and tions of only a few species have been documented northern Timan. The occurrence of this fauna by SIMAKOVA (1986). These include two species of reflects a short transgressive episode in the histo­ Amplexocarinia, two species of Ujimia, and a sin­ ry of the area, and its rapid disappearance coin­ gle species of Cyathocarinia, Hexalasma and cides with a fairly abrupt regression. Soshkineophyllum, assigned to the Lower The most complete Carboniferous-Permian Artinskian Pseudofusulina concavuta-Ps. pedis­ succession occurs in Central Vestspitsbergen. An sequa Biozone, and one species of abundant coral fauna was described from that area Amplexocarinia, assigned to the Upper Artinskian by HERITSCH (1939), but unfortunately, the type Ps. Juresanensis-Parafusulina lutugini Biozone. material for that monograph appears to have been Among the Artinskian taxa described by lost, thus making taxonomic revision impractical. SOSHKINA & al. (1941) and not recognized by Also, the stratigraphic occurrence of the fauna is SIMAKOVA (1986) or KOSSOVAYA (1996), we were mostly unknown. Some introductory identifica­ able to confirm the occurrence of Calophyllum, tions by FORBES & al. (1958) added little to our Lophophyllidium (Lophbillidium) (and perhaps understanding of the Central Vestspitsbergen the nominative subgenus Lophophyllidium), fauna. More recently, however, SOMERVILLE Lophocarinophyllum, Paralleynia, Plerophyllum (1997) began a restudy of the stratigraphic distri­ and Pseudowannerophyllum. The exact strati­ bution and taxonomy of the rugose corals from graphic position of those genera is unknown. this area. He recognized fairly abundant solitary Thus, following the original descriptions, we con­ dissepimental corals low in the Asselian­ sider them to be Artinskian. Following Sakmarian succession, accompanied by fascicu­ FEDOROWSKI (1974, 1986) we do not accept the late colonies of Heintzella and Tschussovskenia reported occurrence in the Ural Mountains of the and, higher in the section (probably in Sakmarian genera Timorphyllum and Verbeekiella, both of strata), by massive colonies of Kleopatrina and which are restricted to the Tethys Realm. Stylastraea. He did not mention cerioid- aphroid taxa. Cystophora svalbardica HERITSCH, 1939 is the only cerioid-aphroid coral described from this Svalbard Archipelago area. Unfortunately, the only location given for this specimen is "east side of Green Harbour". Cisuralian rugose corals are abundant in the Two species assigned by HERITSCH (1939) to Svalbard Archipelago, but their ages are uncer­ Orionastraea were collected from the Hornsund tain. They are present on several islands of the area and belong to Stylastraea. Thus, it appears Archipelago, but are best known from the that cerioid-aphroid corals are extremely rare in Hornsund area of Vestspitsbergen (FEDOROWSKI Central Vestspitsbergen. 1997). Unfortunately, lack of associated index On Bear Island, the Carboniferous-Permian suc­ fossils has prevented a reliable age determination cession resembles that of Central Vestspitsbergen. for that abundant fauna. FEDOROWSKI (1997) con­ Unfortunately, the abundant Bear Island fauna, col­ sidered its. age to be Sakmarian by comparison lected in the late seventies by one of us UF) and with other coral faunas of the realm. It consists housed in Paleontologisk Museum in Oslo, has mostly of cerioid colonial corals of the genera never been described. Introductory field identifica­ Stylastraea, Kleopatrina and a new, undescribed tions by J. FEDOROWSKI indicate that cerioid genus. Cerioid-aphroid colonies are not abundant colonies of Kleopatrina and Stylastraea greatly and, in contrast to the succession of the Urals and outnumber other rugosan growth forms, but bothro­ northern Timan, they occur stratigraphically phyllids and Heintzella are present as well. below the cerioid genera listed above. The ceri­ oid-aphroid colonies are accompanied by abun­ dant representatives of the Bothrophyllidae and a Canadian Arctic Archipelago few species of the fasciculate genera Heintzella and Tschussovskenia, which are less common in Cisuralian rugose corals are abundant in the higher strata (FEDOROWSKI 1965, 1967, 1986). Sverdrup Basin, but most of them remain unde- 166 JERZYFEDOROWSKI, E.WAYNEBAMBER & CALVINH.STEVENS scribed. From papers by SALTER (1855), complex axial structures, at least some of which TSCHERNYSCHEV & STEPANOV (1916), MINATO belong to Fedorowskiphyllum. Such solitary (1960), HARKER (1960) and introductory identifi­ corals also occur in allochthonous terranes in cations by two of us (E.W.B and J.F.) it is obvious northwestern British Columbia (undescribed) and '"I that the rugose coral fauna comprises all growth California (WILSON 1982). The non-dissepimental forms, with fasciculate colonial corals prevailing coral taxa identified by ROWETT (1975), and slightly over massive corals and solitary nondis­ briefly re-interpreted by FEDORoWSKI (1997) as sepimental taxa remaining in the minority. The lat­ Allotropiochisma, Calophyllum, Euryphyllum ter group occurs mainly in the middle to upper and Soshkineophyllum, are typical for the Sakmarian part of the succession. The succession youngest coral-bearing strata of Vestspitsbergen, of growth forms appears to be similar to that Greenland and the Sverdrup Basin. Thus, it may described by KOSSOVAYA (1996), but this cannot be suggested that rugose coral faunas persisted in be verified at present. Colonial rugose corals are Alaska as long as in the other arctic areas listed very rare in the lowest part of the Asselian succes­ and became extinct when extrinsic conditions sion, where only a single occurrence of became unacceptable for coral growth. "Heritschioides" is known. This level contains fairly abundant solitary, dissepimental Siedleckia, Timania, and Fedorowskiphyllum. These and British Columbia related genera also occur in varying abundance throughout younger Cisuralian strata. Younger, The Cisuralian rugose corals of British upper Asselian to lower Sakmarian strata are dom­ Columbia have been only partly described (Wu & inated by fasciculate genera, including al. 1985; STEVENS & RYCERSKI 1983, 1989) or Paraheritschioides and Heintzella, accompanied preliminarily identified by one of us (EWB). ,I by Durhamina, and Tschussovskenia. Also at this They occur in the thin, cratonic succession of I level there are a few cerioid durhaminid corals east-central British Columbia and in the accreted belonging to Pararachnastraea and a new, unde­ Stikinia and Quesnellia terranes of northwestern scribed genus. Kleopatrina, Stylastraea and and south-central British Columbia. Pararachnastraea are the most abundant genera The undescribed, upper Asselian to mid­ in the overlying, mid-Sakmarian? part of the suc­ Sakmarian corals of the autochthonous succession cession. Protolonsdaleiastraea and other cerioid­ in east-central British Columbia consist almost aphroid forms dominate the upper part of the suc­ entirely of colonial forms belonging to taxa that are cession, ranging through the upper half of the widespread in correlative strata of the CAU Realm. Sakmarian and into strata of possible early Precise age ranges are not yet firmly established Artinskian age. These are accompanied by but there appears to be an older, upper Asselian to Permastraea and the new cerioid durhaminid lower Sakmarian fauna containing the fasciculate genus referred to above. genera "Heritschioides", Paraheritschioides, Heintzella, and Durhamina, as well a notable occurrence of the cerioid-aphroid genus East-central Alaska Protolonsdaleiastraea. Cerioid-aphroid forms also occur this low in the succession in the A small Asselian/Sakmarian fauna, described Svalbard Archipelago and in the Stikinia Terrane by ROWETT (1969) from the Wrangellia Terrane of northwestern British Columbia, as noted else­ in east-central Alaska, contains taxa in common where. This fauna is overlain by a younger, lower with both autochthonous and allochthonous fau­ to middle Sakmarian fauna dominated by cerioid nas elsewhere in the CAU Realm. These include colonies of Stylastraea and Kleopatrina, associat­ colonial species of "Heritschioides" and ed with fasciculate Durhamina. Durhamina, also found in allochthonous strata of A rich fauna of Asselian to lower ? Sakmarian east-central British Columbia and arctic mainland colonial and solitary, dissepimented corals (Wu & Canada. In addition, undescribed collections al. 1985; STEVENS & RYCERSKI 1989) occurs in the available to one of us (CHS) contain the genera lower part of the Permian succession of the ?Lytvophyllum, ?Stylastraea, Kleopatrina, allochthonous Stikinia Terrane of northwestern Stikineastraea and?Protolonsdaleiastraea. Also British Columbia (GUNNING & al. 1994). The ver­ present are dissepimental solitary corals with tical succession of taxa within this terrane is not PERMIAN CORALS 167 well established, but the age range of the fauna Similarly, the autochthon to the east has yielded appears to be restricted to Asselian and early the richest fauna known from the Pangaean cra­ Sakmarian, based on associated fusulinacean fau­ tonal margin. Several workers, including EASTON nas. Genera present include: Petalaxidae - (1960), STEVENS (1967), WILSON (1991, 1994), Cystolonsdaleia, ?Lytvophyllum, Petalaxis; and WILSON & LANGENHEIM (1962, 1993) have "Heritschioididae" - Heintzella, "Heritschioides", studied this fauna, with many additional unde­ Paraheritschoides, ?Protolonsdaleiastraea, scribed forms collected by one of us (CHS) and Stikineastraea; Durhaminidae - Durhamina, identified by us in 1996. Pararachnastraea, and two new cerioid-aphroid In the Eastern Klamath terrane, relatively sim­ genera. The latter constitute a third occurrence of ple fasciculate corals probably belonging to, cerioid-aphroid forms at a relatively low strati­ Paraheritschioides first appear in the Upper graphic level. The presence of Petalaxis and Carboniferous along with solitary dissepimental Cystolonsdaleia, which are absent from more rugose corals. The cerioid genus Petalaxis appears northerly successions of North America, the in the lowermost Permian strata, followed by a Svalbard Archipelago and Russia (BAMBER & cerioid durhaminid and several species of FEDOROWSKI 1998), is a feature in common with "Heritschioides" . Younger, middle Asselian corals more southerly faunas of the western U.S.A. and of fusulinacean Zone D include cerioid-aphroid Bolivia. Such complex colonial genera as genera and fasciculate "heritschioidids" with septal Langenheimia and new, unnamed genera with lamellae outnumbering major septa. Kleopatrina extra septal lamellae, found in more southerly suc­ with extensive development of lonsdaleoid dissepi­ cessions, are absent from northwestern British ments occurs in fusulinacean Zone F (lower Columbia. Solitary corals occurring in the Stikinia Sakmarian). In the youngest beds some of the Terrane fauna have not yet been studied in detail, corallites are very large, reaching diameters of 27 but appear to comprise mainly Bothrophyllidae mm and 16 mm in fasciculate and cerioid-aphroid similar to those elsewhere in CAU Realm, and also corals, respectively. Thus, the sequence of appear­ include forms with complex axial structures known ance of different coral morphotypes in this area is from east-central Alaska (ROWETT 1969) and similar to that in other parts of the CAU Realm. In Eastern Klamath Terrane in California (WILSON the faunas of the Eastern Klamath terrane we rec­ 1982). ognize the following colonial coral genera: In south-central British Columbia, the Petalaxis, Cystolonsdaleiastraea, ?Lytvophyllum, Quesnellia Terrane contains a small, undescribed Durhamina, two new genera of cerioid Asselian colonial coral fauna, which includes the durhaminids, Paraheritschioides, a new "herits­ genera Petalaxis, Cystolonsdaleia and chioidid" genus with extra septal lamellae, ?Lytvophyllum, in common with the Stikinia Stylastraea, Kleopatrina, Langenheimia, Terrane and other, more southerly allochthonous Pseudocystophora, and a new cerioid-aphroid terranes ofthe CAU Realm. Also occurring in this genus. structurally complex succession, however, is a In the autochthon of west-central U.S.A. the lower Guadalupian species of Parawentzelella, a sequence of appearance of coral morphotypes is genus with tethyan affinities, as noted below. similar to that in the Eastern Klamath terrane, but some taxa appeared later if the correlations used here are correct. Fasciculate corals, including West-central U.S.A. Paraheritschioides, appear in the Upper Carboniferous. In the lowermost Permian strata In west-central U.S.A., Cisuralian colonial cerioid corals belonging to the simple genus rugose corals are present in both the allochtho­ Stylastraea and simple durhaminid fasciculate nous terranes and the autochthonous rocks of the corals are present. These are succeeded by the Pangaean margin. The best succession of more complicated cerioid genus Kleopatrina. allochthonous corals, associated with fusuli­ Higher in the section (in the Artinskian) there are nacean zones (SKINNER & WILDE 1965), occurs in fasciculate durhaminids with extra septal lamellae the Eastern Klamath terrane. This fauna was and cerioid-aphroid genera. Interestingly, where­ described by WILSON (1982, 1985) and is appar­ as "Heritschioides" is abundant in Asselian rocks ently the richest of all those represented in the of the allochthonous terranes, the genus may not allochthonous terranes of North America. occur in the autochthon below the Artinskian. 168 JERZY FEDOROWSKI, E. WAYNE BAMBER & CALVIN H. STEVENS

Finally, cerioid forms with extra septal lamellae should be mentioned as being very common and occur in the youngest beds containing colonial abundant. Bradyphyllum and Lophotichium are corals. These colonial corals also appear to be the genera rooted in the Carboniferous and present in youngest colonial corals to originate in the CAU both the Tethys and CAU realms. The greatest Realm, probably having lasted through the . number of species and specimens in the Artinskian and possibly into the early Cisuralian fauna of the Diffingiina Province occur Guadalupian (Roadian). in the upper Wolfcampian part of this succession. Genera recognized in the autochthonous part of In undescribed collections, the richest faunas are west-central U.S.A. include Tschussovskenia, those from the Cisuralian and the Roadian. The Petalaxis, Durhamina, a new durhaminid genus youngest corals of the Diffingiina Province are with a continuous axial column, 3 new cerioid found in strata. A drastic change in durhaminid genera, Heintzella, Paraheritschioides, environmental conditions ended the history of this . a new durhaminid genus with extra septal lamellae, coral province. Stylastraea, Kleopatrina, Permastraea, ?Protolonsdaleiastraea, and "Heritschioides". Rare colonial rugose corals similar to those in GUADALUPIAN the autochthon of west-central U.S.A. are present -I in northwestern Mexico, Guatemala, Bolivia, and As mentioned above, the richest Guadalupian 1 Peru (e.g. ROWETT & WALPER 1972, WILSON coral fauna occurs in the Diffingiina Province. 1990, STEVENS 1995), but our present knowledge According to the description of rugose corals of those faunas is insufficient for a meaningful from Greenland by FUlGEL (1973), the discussion. Guadalupian fauna from the northeastern part of this Island should be considered as the second richest. However, re-description and revision of Southwestern Texas and southeastern part of that fauna by FEDOROWSKI (1982) and by New Mexico FEDOROWSKI & BAMBER (in prep.) show that an excessive number of taxa has been recognized The rugose coral fauna of southwestern Texas (for instance, FLUGEL'S five new species of and adjacent part of New Mexico (Glass Calophyllum all belong to one previously Mountains and ) can best be described species) and that some important taxa characterized as follows: paucity of massive colo­ have not been recognized. In our opinion the nial taxa; impoverishment of fasciculate colonial Greenland Guadalupian fauna is represented by and solitary dissepimental taxa and restriction of the following: Calophyllum columnare these groups mainly to the Upper Carboniferous (SCHLOTHEIM, 1813), which is common for the and lower Cisuralian; abundance of solitary entire northern part of the realm, including nondissepimental taxa, continuing into the Alaska; Paracaninia variabilis (SOSHKINA, Guadalupian, when coral faunas were generally 1941), which is almost as widespread as C. impoverished or absent from other areas of the columnare; three species of Allotropiochisma CAU Realm. Those characteristics, combined (one in common with the Sverdrup Basin fauna); with the presence of the endemic suborder and single species of Tachylasma, Ufimia (new Diffingiina, led FEDOROWSKI (1997) to propose species in common with Sverdrup Basin), proba­ the name Diffingiina Province for this area. bly Leonardophyllum and Soshkineophyllum. In addition to the Diffingiina, represented by Most of these genera have long stratigraphic two families, three genera and several species, ranges and are cosmopolitan. Only fourteen non dissepimental rugose coral species Leonardophyllum, if truly present in Greenland, have been described from this area. The most is a surprising newcomer from the south. abundant and diversified genera are Assimulia, A fauna very similar to that of northeastern Lophophyllidium and Paraduplophyllum, cach Greenland, and equally diverse, has been represented by two subgenera and several species. described by FEDOROWSKI & BAMBER (in prep.) For some of these species, very large numbers of from the Sverdrup Basin in the Canadian Arctic specimens were investigated (FEDOROWSKI 1987). Archipelago. It contains two species of Allotropiochisma, Bradyphyllum, Euryphyllum, Allotropiochisma (one unnamed), two species of Falsiamplexus, Lophotichium and Lytvolasma Euryphyllum, and single species of Calophyllum PERMIAN CORALS 169

(c. columnare), Lytvolasma, Soshkineophyllum Greenland and Vestspitsbergen and those of the (also found in correlative strata of Svalbard) and Zechstein in the Central European Basin. Thus, Ujimia. Although represented by relatively few from several lines of evidence, it would appear specimens, this fauna is surprisingly diverse. that a Wordian (=early? Kazanian) age may be Some rugosan taxa from the Kapp Starostin assigned to all occurrences of this widespread Formation of Central Vestspitsbergen are identi­ solitary coral fauna, which includes some of the cal with those of Greenland and the Sverdrup youngest corals in the CAU Realm. Basin, but further analysis of the content of the Kapp Starostin fauna is not possible because no modern, detailed investigation of these corals has THE NEWCOMERS been completed. Preliminary lists and illustrations of the Kapp Starostin corals were published by As discussed earlier in the paper, with the pos­ EZAKI & KAWAMURA (1992) and EZAKI (1997). sible exception of the youngest autochthonous The rugose coral fauna of the Central faunas of the west-central U.S A., all Cisuralian European Basin and Eastern European Platform is colonial rugose coral faunas of the CAU Realm, taxonomically monotonous, although commonly along the margins of Pangaea and in adjacent rich in specimens. Revisions by WEYER (1979, allochthonous arc terranes (Text-fig. 2), became 1997), WEYER & lUNA (1979) and FEDOROWSKI & extinct prior to the beginning of the Guadalupian. BAMBER (in prep.) indicate that no more than two During mid- Guadalupian (Wordian) to early species of Calophyllum (C. column are and C. Lopingian (early Wuchiapingian) time, however, quadrifidum) and Paracaninia variabilis occur an impoverished colonial coral fauna of obvious in this large area. Ujimia and Allotropiochisma tethyan affinities developed on the Palaeopacific could each be represented by one species as well, carbonate buildups presently preserved in but such occurrences are doubtful. allochthonous terranes of western North America Precise stratigraphic relationships and ages are (NELSON & NELSON 1985; STEVENS & al. 1987; difficult to establish for the Guadalupian occur­ BAMBER, Geological Survey of Canada collec­ rences discussed above. They have all been tions). Waagenophyllids identified to date include assigned to the Kazanian or correlated with that Waagenophyllum, Parawentzelella, and possibly , but the term Kazanian may, in some Pseudohuangia (newly discovered, not yet instances, equate to the entire Guadalupian. The described), one or more of which occurs in the age of the Sverdrup Basin fauna has been fairly following terranes: the Quesnellia and Eastern well established as Wordian (Middle Klamath terranes, which also contain Cisuralian Guadalupian) and early Kazanian on its associa­ corals of the CAU Realm; the Cache Creek ter­ tion with rare ammonoids (NASSICHUK 1995, p. rane, in which fossils as old as mid-Carboniferous 218), brachiopods (THORSTEINSSON 1974, p. 72) have strong tethyan or mixed tethyan-North and palynomorphs. In eastern Greenland, the American affinities; and in limestone blocks of youngest beds of the PermiaRsuccession contain the Hayfork terrane. As previously noted, such early Wuchiapingian (=earliest Lopingian) waagenophyllid corals are typical of the tethyan ammonoids of the genus Cyclolobus (NASSICHUK, Realm and the means by which they arrived in 1955, p. 229). Older strata in the area, from above North America has not been established. the coral-bearing limestone, have yielded con­ Unlike the colonial waagenophyllids, the odonts of late Wordian or Capitanian age Guadalupian solitary, nondissepimental corals (NASSICHUK 1995, p. 230), thus indicating a prob­ known from Texas, arctic Canada, Greenland and able upper limit of Wordian for the age of the Svalbard (see above) do not appear to be new­ Greenland coral fauna. The brachiopod fauna of comers to the CAU Realm for the following rea­ the coral-bearing Kapp Storostin Formation of sons: 1) There was a permanently developing Vestspitsbergen (see above), was equated by fauna in the Diffingiina Province to serve as a NAKAMURA & al. (1992, p. 83) with that of the potential source for the fauna of other areas. 2) Trold Fiord Formation (Wordian), which contains Most, if not all of the solitary, nondissepimental part of the solitary coral fauna of the Sverdrup genera occurring in the Guadalupian are known to Basin, and also with the brachiopod fauna associ­ occur in the Cisuralian as well, thus providing the ated with the Greenland corals. They also noted a possibility for a natural lineage to develop. 3) All correlation between the brachiopod faunas of species present are endemic to the realm, 170 JERZY FEDOROWSKI, E. WAYNE BAMBER & CALVIN H. STEVENS although the wide geographic distribution sug­ Archipelago, arctic North America and British gests that they had long-living larvae, and there­ Columbia) is consistent with a low latitude position fore the potential ability to cross Panthalassa. for those areas during much of Cisuralian time. 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Manuscript submitted: 15th October 1998 Revised version accepted: 5th December 1998