Reprint of: The Bohemo-Iberian regional chronostratigraphical scale for the Ordovician System and palaeontological correlations within South Gondwana

JUAN CARLOS GUTIERREZ-MARCO, ARTUR A. SA, DIEGO C. GARCIA-BELLIDO AND ISABEL RABANO

We are reprinting the above article in this special issue on ‘The contribution of fossils to chronostratigraphy, 150 years after Albert Oppel’ in issue 50:3 of Lethaia. The article was published in error in an earlier issue (Lethaia 50:2), and is reprinted here for ease and to complete the special issue. Furthermore, we have made a small correction to Figure 2 in this reprinted version of the article. The origi- nal published version had a missing label in the image. In the right-hand column, the text ‘Dobrotivian St’ should appear in the rectangle above the text ‘Oretanian St’; this has been corrected in this reprinted version of the article (and also in the online version of the article published in issue 50:2). Wiley would like to apologise to the Editors and authors for this error during production. For consistency, please continue to cite the article based on its original publication: Gutierrez-Marco, J.C., Sa, A.A., Garcıa-Bellido, D.C. & Rabano, I. 2017: The Bohemo-Iberian regional chronostratigraphical scale for the Ordovician System and palaeontological correlations within South Gond- wana. Lethaia 50, 258–295. doi: 10.1111/let.12197.

DOI 10.1111/let.12216 © 2017 Lethaia Foundation. Published by John Wiley & Sons Ltd The Bohemo-Iberian regional chronostratigraphical scale for the Ordovician System and palaeontological correlations within South Gondwana

JUAN CARLOS GUTIERREZ-MARCO, ARTUR A. SA, DIEGO C. GARCIA-BELLIDO AND ISABEL RABANO

Gutierrez-Marco, J.C., Sa, A.A., Garcıa-Bellido, D.C. & Rabano, I. 2017: The Bohemo- Iberian regional chronostratigraphical scale for the Ordovician System and palaeontological correlations within South Gondwana. Lethaia, Vol. 50, pp. 258–295. Abstract

The Bohemo-Iberian regional scale for South Gondwana, involving the ‘Mediter- ranean Province’, comprises five regional stages (Arenigian, Oretanian, Dobrotivian, Berounian and Kralodvorian) plus the global Tremadocian and Hirnantian. The predominance of shallow-water taxa in those high-latitude faunas imposes serious dif- ficulties for correlating the regional succession with the formal global chronostratigra- phy because of the almost total absence of the key graptolites and conodonts defining the base of the standard series, stages and stage slices. Instead, the abundant benthic faunas (trilobites, brachiopods, molluscs, echinoderms) of South Gondwanan origin largely dominated in the area from the middle Darriwilian to the late Katian. The poleward faunal migration of originally Avalonian, Baltic, Laurentian and even Asiatic taxa during the Boda Event of global warming progressively ends with the endemicity in the region, where the ensuing benthic assemblages were severely affected by the Hir- nantian glaciation. The regional scale significantly improves the precision of correla- tions between Ordovician strata from SW and central Europe, North Africa and a large part of the Middle East. An updated record of palaeontological data from areas where Mediterranean faunas remain practically unknown, or are still poorly investi- gated, is also included. Palaeobiogeographical relationships based on the distribution of faunas across South Gondwana are suggested as an improvement for positioning many territories in modern palaeogeographical reconstructions and offer a construc- tive approach to problems related to the pre-Variscan and pre-Alpine orogenic puz- zles. □ Ordovician, Palaeobiogeography, Regional chronostratigraphy, South Gondwana. Juan Carlos Gutierrez-Marco ✉ [[email protected]], Instituto de Geociencias CSIC, UCM and Departamento de Paleontologıa, Facultad de Ciencias Geologicas, Jose Antonio Novais 12, 28040 Madrid, Spain; Artur A. Sa [[email protected]], Departamento de Geologia, Universidade de Tras-os-Montes e Alto Douro, Quinta de Prados 5001-801 Vila Real, Portugal; Artur A. Sa [[email protected]], Centro de Geoci^encias, Universidade de Coimbra, Polo II 3030-790 Coimbra, Portugal; Diego C. Garcıa-Bellido [Diego.Garcia-Bellido@ adelaide.edu.au], Department of Biological Sciences, University of Adelaide, Adelaide, South Australia 5005, Australia; Isabel Rabano [[email protected]], Museo Geominero, Instituto Geologico y Minero de Espana,~ Rıos Rosas 23, 28003 Madrid, Spain; manuscript received on 13/02/2016; manuscript accepted on 16/09/2016.

In the Prague Basin, and well before the establish- century, the traditional British Ordovician scheme ment of the Ordovician System by Lapworth (1879), persisted as a kind of ‘lingua franca’ for Ordovician the French palaeontologist Joachim Barrande (1799– chronostratigraphy (Webby 1998). However, the 1883) recognized the separate character of the Bohe- high faunal endemicity that characterized this period mian Ordovician faunas by the definition of a regio- at a global scale made it difficult to achieve a detailed nal stage (‘etage D’, Barrande 1846) within the set of correlation criteria, especially those arising extent assigned by him to the ‘Silurian system’ of from both the traditional and revised British series Murchison (1839). This ‘Stage D’ was later docu- and stages. This favoured the proliferation of a num- mented (Barrande 1852) and subdivided into five ber of regional chronostratigraphical units (includ- ‘bands’, which comprise the current Lower and Mid- ing regional series, stages, substages and ‘horizons’) dle Ordovician series (‘bande d1’) and the Upper developed during the twentieth century in parallel to Ordovician one (‘bandes d2–5’), based on the distri- the British scale, for a more effective interregional bution of his so-called Second Fauna. correlation. These are the successful regional This original local Bohemian classification was schemes developed for Australia–New Zealand, not recognized internationally, and for more than a Baltoscandia (essentially based on the Estonian

DOI 10.1111/let.12197 © 2016 Lethaia Foundation. Published by John Wiley & Sons Ltd LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 259 classification), China, North America, Russia– REVISED ORDOVICIAN BRITISH-AVALONIAN Kazakhstan (with at least six scales for different CHRONOSTRATIGRAPHY REGIONAL SCALE Ordovician basins or structural provinces) and the (Williamset al. 1972, (Forteyet al. 1995, peri-Gondwanan Mediterranean area (Webby 1998 Harlandet al. 1982) Cockset al. 2010) and references therein). At the same time a consider- able effort by British workers to produce a more Hirnantian Hirnantian standardized scheme based on traditional stratotypes Rawtheyan Rawtheyan led to the discovery of significant stratigraphical gaps ASHGILL or overlaps of the widely used units (Finney 2005). Cautleyan Cautleyan This also generated a conceptual evolution of the ASHGILL former scale (Fig. 1) into a substantially modified Pusgillian Pusgillian et al. scheme (Gutierrez-Marco 2016; with previous Onnian Onnian references). The refined British scheme (Fortey et al. Streffordian 1995, 2000) did not satisfy the international correla- Actonian Actonian Marshbrookian Marshbrookian tion requirements to become the global standard Cheneyan Woolstonian and is today considered a regional scale applicable CARADOC Longvillian for East Avalonia (including the Anglo-Welsh area, Longvillian southern Ireland and Belgium) and parts of northern SoudleyanBurrellian Soudleyan Germany and NW Poland (‘Far Eastern Avalonia’).

Harnagian CARADOC Harnagian To avoid these widespread problems for interconti- nental correlation, the International Commission on Costonian Costonian Stratigraphy (IUGS) first completed in 2007 the for- Upper Aurelucian Velfreyan mal chronostratigraphical classification for the LLANDEILO Middle Ordovician System, introducing a threefold division into Lower, Middle and Upper series and seven new Lower Llandeilian global stages. Among these, only the Tremadocian and Hirnantian stages were adopted from the existing Upper LLANVIRN British names, but their basal global stratotypes were Abereiddian located in Canada and China, respectively (Harper Lower LLANVIRN 2011; Cooper & Sadler 2012). The ‘golden spikes’ for Upper the new series and stage boundaries were placed in Fennian deep-water facies within continuous and fossiliferous

sections, coinciding with the first appearance datum no stages defined (FAD) of particular conodont and graptolite species ARENIG Whitlandian Lower of worldwide distribution. However, the majority of

ARENIG these cosmopolitan taxa (or their FADs) can hardly be recognized in the strata of the extensive shallow- Moridunian water, inner-shelf environments and epicontinental seas that surrounded Gondwana, Baltica and Siberia palaeocontinents. Here, the poor applicability of the Migneintian standard scale is compensated by the capacity of regional scales to obtain more precise local to interre- TREMADOC gional correlations. This is the reason why, in agree- Cressagian ment with the Ordovician Subcommission, an TREMADOC increasing refinement of the regional scales could provide a fundamental contribution to the develop- Fig. 1. Correlation between the former ‘modern’ view for British Ordovician chronostratigraphy, and the present-day British– ment of a better global chronostratigraphy Avalonian regional scale. (Bergstrom€ et al. 2009; Harper 2011). This paper examines the present status of the Bohemo-Iberian (= ‘Mediterranean’) regional Palaeogeographical setting chronostratigraphical scale and its suitability for cor- relating the Ordovician strata of the south polar The Ordovician sedimentary record extending from Gondwanan and peri-Gondwanan areas, as well as the Middle East to north Africa and central and its potential biostratigraphical ties with both the southern Europe is characterized by the develop- global and British regional scales. ment of thick siliciclastic successions, deposited in 260 Gutierrez-Marco et al. LETHAIA 50 (2017) widespread shallow-marine environments generally (Havlıcek et al. 1994; Fatka & Mergl 2009). Accord- ranging from inshore to lower offshore settings. ing to Servais & Sintubin (2009), the available Black shale intercalations are extremely rare and palaeontological and structural evidence suggests proper limestone deposition was mainly restricted to that Perunica was a part of the ATA, and the the short interval of global warming (Boda Event) endemicity of some faunas within Bohemian assem- that preceded the latest Ordovician glaciation (Villas blages did not necessarily derive from a different et al. 2002; Boucot et al. 2003; Fortey & Cocks palaeogeographical setting. 2005). The main regional stratigraphical gaps are The terranes integrated in the ATA mostly have a related to the extensional processes associated with Cadomian basement, indicating pre-Ordovician the splitting and drift of Avalonia from Gondwana affinity with the margin of Gondwana (Winchester in the Early Ordovician (Murphy et al. 2006; Nance et al. 2002). This is consistent with the palaeontolog- et al. 2010); later with the uplift and subsequent rift- ical record for the early Palaeozoic, which confirms ing of the ‘Sardinian–Taurian rise’ in the early Late that continental separation from Gondwana was Ordovician (Leone et al. 1991; Hammann 1992; never significant (Robardet 2003). However, Servais Alvaro et al. 2016) and finally with the glacio- & Sintubin (2009) point out that the separate drift eustatic unconformities produced by the Hirnantian history suggested for some of the peri-Gondwanan glaciation, which had a special impact over the large terranes would indicate that the ATA was possibly continental and marine areas located around the part of a separate plate, albeit palaeobiogeographi- South Pole at that time (Ghienne et al. 2007a,b, cally continuous with Gondwana. This could resem- 2014). ble a ‘Californian scenario’, where both sides of the There is considerable palaeontological, stratigraph- San Andreas Fault share fauna and flora, although ical and palaeomagnetic evidence of the high latitude they are situated on different tectonic plates, as may position of this region in the Southern Hemisphere be the case for the ATA and southernmost Gond- even before the glacial influence over the entire area wana. The existence of transform faults affecting the became evident (Spjeldnaes 1981; Cocks & Fortey peri-Gondwanan Cadomian belt was already sug- 1988; Ghienne et al. 2007b; Torsvik & Cocks 2013a, gested, among others, by Fernandez-Suarez et al. b). The palaeogeographical relationships among the (2002), who interpreted the change of provenance present African and Middle East part of Gondwana sources of detrital zircons from late Neoproterozoic with peri-Gondwanan Europe are, however, very rocks of Ibero-Armorica to record along-margin ter- difficult to assess due to the complex geodynamic rane transport before the opening of the Rheic evolution associated with the eo-Variscan tectonics Ocean. However, Meinhold et al. (2013) proposed and the Variscan closure of the Rheic ocean, and also an alternative interpretation, discarding the separate to the strong Alpine overprint in some important plate hypothesis. In any case, independent evidence areas (von Raumer et al. 2013). In particular, the that the Gondwana ice sheet reached southern Tur- peri-Gondwanan ‘Armorican Terrane Assemblage’ key (Monod et al. 2003) and northern Spain (ATA), or Armorica sensu lato, was considered dur- (Gutierrez-Marco et al. 2010) also argues in favour ing many years as an independent ‘microcontinent’ of such physical continuity of these regions with the or ‘microplate’ between the continents of Gondwana, Gondwana continental mass during the latest Laurentia and Baltica on the basis of flawed or incon- Ordovician. sistent palaeomagnetic data (Tait et al. 1997; Matte The fact that certain Ordovician faunas from 2001; Winchester et al. 2002). However, this isola- Ibero-Armorica show close similarities to those tion was never supported by palaeontological recorded from NE Algeria, Libya and Saudi Arabia evidence, and thus, such a ‘microcontinent’ is con- led Gutierrez-Marco et al. (2002a) to propose that sidered fiction with the ATA having been attached to during Ordovician times Iberia was probably adja- Gondwana all along (Robardet 2003). cent to Libya instead of Morocco, as previously The ATA originally included all the Variscan areas believed. The location of central Iberia probably of southern Europe plus the Saxo-Thuringian and north of present-day Libya or Egypt and not very far Moldanuvian crustal blocks, which supposedly rifted from the Ordovician South Pole was later supported and migrated together towards Baltica from their by isotopic and inherited studies of magmatic rocks former peri-Gondwanan position (Winchester et al. (Bea et al. 2010; Meinhold et al. 2013). These indi- 2002). Later, the Bohemian Massif and its adjacent cate a close proximity of Iberia with the northern- areas were considered as a microcontinental plate most part of the Neoproterozoic Eastern African (Perunica), also independent from Gondwana, Orogen, instead of NW Africa near Morocco (Nance although sharing many palaeontological features in et al. 2008). U–Pb detrital zircon ages for the Early common with Ibero-Armorica and NW Africa Ordovician sandstones of the Iberian Peninsula LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 261

(Shaw et al. 2014) also indicate a central to east Afri- eastern branch of the Rheic Ocean (Hunia terranes). can depositional realm. The latter proposes that the In other recent papers (Arenas et al. 2016; Dıez 2,300-km-long Cantabrian–Central Iberian portion Fernandez & Arenas 2015), the proposal of the of the Early Palaeozoic Gondwana margin stretched new Galicia–Ossa-Morena Zone in western Iberia east–west along the northern limits of the then low- involves a Late Devonian Variscan suture that dupli- lying Saharan Metacraton and Arabian–Nubian cates the Gondwanan platform, with the former out- Shield. board Ossa-Morena Zone being juxtaposed on top A more advanced study using hafnium isotopic of the Central Iberian Zone and/or the ‘SW Iberia fingerprints in detrital zircons over a wide European Autochthon’, which were located closer to the main- area (Henderson et al. 2016) supports the interpre- land before the Variscan amalgamation. tation that autochthonous Iberian terranes occupied Regardless of the above-mentioned advances and a palaeoposition adjacent to the Sahara Metacraton problems for understanding pre-Variscan palaeogeo- during the late Neoproterozoic and early Palaeozoic, graphical reconstructions, Torsvik & Cocks (2011, whereas the Armorican Massif, the French Massif 2013a,b) tend to simplify the correlation between Central and the Bohemian Massif derive from the the European and African Gondwana, using Bug- West African Craton. However, U–Pb ages of detri- Plates because ‘no palinspastic reconstructions are tal zircons from the Bohemian Massif also support a agreed’. They show Iberia as a single unit, rotated North African derivation of parts of this area from 180° and placed in the vicinity of Morocco and Alge- metamorphic and magmatic sources on the Ara- ria. The Ossa-Morena Zone is represented already bian–Nubian Shield (Bahlburg et al. 2010) or the attached to the Central Iberian Zone in the early Trans-Saharan belt (Drost et al. 2011). Palaeozoic, in spite of its Ordovician succession Focusing on SW Europe, and according to the containing deep-water Ordovician faunas (nileid– stratigraphical and palaeontological evidence, the cyclopygid biofacies), in evident contrast to the Mid-North Armorican Domain should be consid- remaining Iberian areas. Recent studies have pro- ered a direct extension of the Central Iberian Zone posed that Ossa-Morena was deformed and stacked of the Iberian Peninsula (Paris & Robardet 1977, on the central shelf as part of a huge rootless nappe 1990; Robardet et al. 1990). The Armorican Massif by the Late Devonian, as a result of an eo-Variscan would be inverted to its present-day position (Ham- continental subduction prior to the Rheic closure mann 1992) and originally located towards the west (Dıez Fernandez & Arenas 2015). After this process, of Iberia. This Mid-North Armorican–Central Ibe- the Ossa-Morena Complex would have obliquely rian domain appears to be a key element for the pre- moved to its present-day position by long strike-slip Variscan reconstructions of southern Europe faults and narrow shear belts during the main Varis- (Robardet 2002), despite frequently being over- can phase (see discussion in Gutierrez-Marco et al. looked in palaeogeographical studies and Ibero- 2014b). The palaeogeographical reconstructions of Armorican structural correlations. Many of these Torsvik & Cocks (2011, 2013a,b) support the stabil- placed the North Armorican Domain as being ity of the Mid-North Armorican–Central Iberian related to the Ossa-Morena Zone based on their domain, but place the Armorican Massif to the east strong Cadomian imprint (e.g. Franke 2000, 2006; of Iberia, at a palaeolatitude of about 80° South for Matte 2001; Martınez Catalan 2011), in obvious the Early Ordovician. However, this orientation contradiction with their sedimentary and palaeobio- ignores the deepening of the extensive inshore shelf geographical record for the Early Palaeozoic (Robar- that occurred southwards in Mid-North Armorica det 2002). and northwards at the Central Iberian Zone – both The pre-Variscan European scenario is even more in present-day coordinates – as demonstrated, complex when considering other plate-tectonic among others, by the distribution of the Middle reconstructions proposed for the southern Gond- Ordovician sandy tempestites (Brenchley et al. wana margin. Some authors (von Raumer & Stamp- 1986) and the trilobite biofacies (Henry 1989; fli 2008; von Raumer et al. 2013) have proposed a Rabano 1989). Likewise, the French Massif Central ribbonlike continent, more than 10,000 km long, and the Italian territories of Sardinia and Calabria spreading along this margin since the Cambrian, were incorrectly placed on each side of Ibero- calling it the Greater Galatian or ‘Hun’ superterrane. Armorica. Nonetheless, those reconstructions admit- This superterrane would comprise some of the ted that all these territories did not split from Gond- future Avalonian blocks, a number of the Palaeozoic wana until the opening of the Palaeotethys Ocean in south European intra-Alpine blocks (‘Galatian ter- the latest Silurian or even later. rane’), as well as some terranes possibly accreted to Despite the great difficulties in elucidating the the North China Block after the opening of the relationships among the pre-Variscan European 262 Gutierrez-Marco et al. LETHAIA 50 (2017) areas, the following general consideration must be latitude shallow-shelf faunas as being composed of a added, in line with the important observations made typical ‘calymenacean–dalmanitacean’ assemblage by Servais & Sintubin (2009). It is well known that with calymenoids (Colpocoryphe, Salterocoryphe, Pra- the supercontinent Gondwana stretched from South doella, Calymenella, Iberocoryphe, Eohomalonotus, Gondwana (today’s Southern Europe and North Kerfornella and Plaesiacomia), dalmanitoids (Zelisz- Africa), then at high latitudes, to tropical North kella, Toletanaspis, Kloucekia, Morgatia, Crozonaspis, Gondwana (Australia and North China), lying in the Eodalmanitina, Retamaspis, Ormathops, Dalman- Ordovician Northern Hemisphere. South America, itina, Eudolatites and Dreyfussina), as well as repre- Middle East and a number of south Asian peri- sentatives of other trilobite groups (e.g. Isabelinia, Gondwanan terranes were located in intermediate Nobiliasaphus, Merlinia, Ogyginus, Parabarrandia, palaeolatitudes (e.g. Fortey & Cocks 2003; Torsvik & Barrandia, Ectillaenus, Placoparia, Eccoptochile, Ura- Cocks 2013a). Most palaeogeographical maps coin- lichas and Selenopeltis). Also very typical of these cide in placing present-day North Africa and Gondwanan inshore-shelf fauna is a group of large Mediterranean Europe near the Ordovician South linguliform brachiopods such as Lingulobolus, Ecte- Pole, although part of this southern Gondwanan noglossa, Pseudobolus, Lingulepis and Monobolina. margin was facing north. This is the reason why this Among the rhynchonelliform brachiopods wide- area has being erroneously labelled as ‘North Gond- spread endemic genera as Crozonortis, Cacemia, wana’ and the term should be avoided and replaced Apollonorthis, Tazzarinia, Eodalmanella, Euorthisina, by one that reflects either their appropriate palaeo- Nocturniella, Ranorthis and Prantlina can be recog- geographical position, such as South Gondwana (= nized, as well as the typical Aegiromena–Drabovia ‘south polar Gondwana’ of Servais & Sintubin 2009; fauna that dominated from Bohemia to Africa in Gutierrez-Marco et al. 2016), or the present-day ‘Caradocian’ (=Berounian) times. extent, with Ordovician rocks occurring in many cir- As discussed above, the East and ‘Far Eastern’ cum-Mediterranean countries. In the last case, the Avalonia were part of the Gondwanan margin until old name ‘Mediterranean area’ that is still being used the earliest Middle Ordovician, when the microcon- by many palaeobiogeographers may be rescued. tinent split from Gondwana and drifted towards Bal- tica, carrying slightly different faunas. Gondwanan trilobites from Wales and Shropshire comprise, Palaeobiogeographical notes among others, Neseuretus, Colpocoryphe, Caly- menella, Ormathops, Placoparia, Ogyginus and The existence of distinct assemblages of trilobites, Merlinia (Cocks & Fortey 1988), exhibit greater brachiopods, echinoderms, molluscs, bryozoans, and affinities with the deep-water cyclopygid faunas that even ostracods over a wide area of North Africa, rarely occur in the marginal facies of the Baltica con- peri-Gondwanan Europe and eastern Avalonia tinent. A number of Early and Middle Ordovician allowed the early differentiation of the so-called trilobite and brachiopod species were shared Mediterranean fauna (Spjeldnaes 1961), Selenopeltis between Avalonia and South Gondwana, before fauna (Whittington 1966) or – proto – Tethyan significant broadening of the Rheic Ocean: a good fauna (Dean 1967a). This was followed by the pro- example is the pliomerid trilobite Placoparia (P.) posal for a Mediterranean Province (Spjeldnaes cambriensis, recorded from Wales, Shropshire, Lake 1967), approximately equivalent to the Selenopeltis District, Belgium, Ibero-Armorica, Bohemia, north Province of trilobites (Whittington & Hughes 1972), Germany and Morocco, together with species of typically recognized in Bohemia and Ibero-Armorica, Selenopeltis or Pricyclopyge, some of them remark- Morocco, most of Germany, Bulgaria, Belgium, the ably extending into the Balkans. Anglo-Welsh region and Florida. Transitional areas The faunal list for South Gondwana has increased with faunas from intermediate palaeolatitudes in considerably in the last two decades with the Turkey, Middle East and South America were also advancement of a detailed knowledge of many fossil recognized. According to Spjeldnaes (1967), among groups, their stratigraphical distribution and the the most distinctive taxa of this province are the environmental patterns controlling the development trilobite genera Dalmanitina, Kloucekia, Colpocoryphe, of some particular biofacies and ‘communities’ in Calymenella and the constituents of the ‘Neseuretus both the main, stable core of the Mediterranean Pro- tristani fauna’, brachiopods such as Nicolella, Svobo- vince and adjacent areas with intermediate palaeolat- daina and Porambonites, cystoids and bryozoans itude. Periods of faunal migrations favoured by (Chasmatoporella, Hallopora, Homotrypa and Praso- oceanic and marine currents, significant transgres- pora). Cocks & Fortey (1988), Cocks et al. (1990) sions and short-lived episodes of global climatic and Fortey & Cocks (2003) characterized the high- amelioration such as the Boda Event led to the LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 263 arrival in cool-water shelf palaeoenvironments of dramatic impoverishment as previously believed South Gondwana of taxa previously known from (Havlıcek 1989). intermediate palaeolatitudes of South America and Havlıcek (1982, 1989) initiated the study of Baltica (Gutierrez-Marco et al. 1999) and also from Ordovician benthic communities in Africa and peri- tropical areas of East Asia and Laurentia (Fortey & Gondwanan Europe, with an essentially palaeoeco- Cocks 2005). Proper non-cosmopolitan but pan- logical and palaeoclimatic approach. Subsequent Gondwanan taxa, which could cross a wide range of studies have considerably increased the number of climatic zones, are a minority but are represented by benthic communities (presently described as ‘assem- some trilobite genera (Neseuretus, Hungioides), blages’) defined by particular taxa, showing that the molluscs (Peelerophon), echinoderms (Lingulocystis, changing biofacies are also the expression of the ?Ascocystites, ?Ramseyocrinus) rare conodont species development of a rift-basin structure for the Barran- (Panderodus nogamii) and arandaspid fish dian area. This favoured the differentiation of ben- (Sacabambaspis). thic atheloptic assemblages with common record of Focusing on the Mediterranean area, Servais & nileids and mesopelagic trilobites (cyclopygids) Sintubin (2009) considered the possibility that the towards the centre of the basin, but far from the Bohemian Massif was the site of a distinctive palaeo- oceanic depths reached by the typical olenid biofa- biogeographical province within peri-Gondwanan cies. Limited development of shallow inshore bio- Europe. As already noted, the presumed microconti- topes under steep shelf gradients and roughly nent ‘Perunica’ was proposed based on a combina- coinciding with intervals of tectonic instability did tion of Ordovician faunal lists (Havlıcek et al. 1994; not facilitate the dispersal into Bohemia of some Fatka & Mergl 2009) and palaeomagnetic data, now widespread shallower assemblages (for instance, the disregarded as contradictory (Krs & Pruner 1999). Neseuretus biofacies) that characterize the north The former show the influx of Anglo-Baltic faunas African–SW European platform in the Middle and in the Lower Ordovician, the exclusive relationships Upper Ordovician. with South Gondwana for most of the Middle and The South Gondwana biogeographical region Upper Ordovician, and the re-establishment of the would be considered as broadly equivalent to pre- faunal connection with Baltica and with more cos- sent-day Antarctic and Subantarctic regions and mopolitan faunas by the late Katian and Hirnantian. essentially limited by water temperature and The independent plate-tectonic scenario for ‘Peru- palaeotopography (Servais et al. 2013), but with a nica’ was not supported by the provenance studies relatively small diversity of neritic palaeoecosystems. based on detrital zircons (Bahlburg et al. 2010; Drost Geographical continuity and circumpolar and subpo- et al. 2011). Besides the record of many endemic lar currents must have favoured the dispersal of par- forms among the Ordovician taxa from Bohemia, ticular species and genera that became broadly but probably influenced by the remarkable persis- distributed and thus provide good biostratigraphical tence and quality of palaeontological studies in the tools. However, some conceptual problems with the small Prague Basin for more than 160 years, the use nomenclature of Gondwanan/Bohemian, Baltic, of the broad term ‘Bohemian faunas’ has been Avalonian or Laurentian taxa persist over the entire criticized since Dean (1967a), who already indicated area. These terms usually derive from places where that ‘the shelly faunas of Bohemia, varied and well- certain genera or species are abundant and were first developed though they are, inhabited only part of a described in the palaeontological literature, yet do much larger Tethyan region with broadly similar not coincide with their area of origination. For faunal characteristics’. The same view was shared by, instance, the disparid crinoid Iocrinus was usually among others, Havlıcek (1989) with regard to pre- regarded as a Laurentian genus, but due to its first Variscan palaeontology and palaeogeography of chronostratigraphical occurrences in Oman, Mor- the South Gondwana margin. Within this biochore, occo, Spain and Great Britain (Donovan et al. 2011; intraregional faunal differences have been strongly Zamora et al. 2015) it should be regarded as a Gond- conditioned by the development of different wanan genus that later migrated into lower palaeolat- biofacies defined by the type of substrate, inshore– itudes. Gutierrez-Marco et al. (1999a) added offshore gradients, climate–palaeolatitude, palaeocur- examples of echinoderm and trilobite species of rents and active tectonics coeval with Ordovician Ibero-Armorican origin that later migrated to Bohe- sedimentation. Furthermore, the increasing number mia, where they received their first description. The of modern palaeontological studies from areas out- same happened with a number of conodonts, grapto- side the Prague Basin shows that the palaeodiversity lites and ostracods that are commonly regarded as of benthic assemblages estimated in a poleward sense ‘Baltic’ elements in Gondwana, although they range starting from Bohemia did not record such a over more than one palaeobiogeographical province 264 Gutierrez-Marco et al. LETHAIA 50 (2017) or even can be considered almost cosmopolitan taxa. Algerian Sahara after finding correlation difficulties Inversely, some typical benthic elements (i.e. the between the allegedly endemic North African grap- bivalve Hemioprionodonta lusitanica) in Southern tolite and brachiopod faunas and the generalized Gondwana made their first appearance in the Andean faunal succession derived from British stratotypes. Basin (Sanchez & Babin 2005) from where they They introduced the regional stages ‘n1’ (roughly migrated along the Gondwanan margin during the equivalent to the Tremadocian to lower Arenigian – Early Ordovician to reach higher palaeolatitudes in sic – interval), ‘n2’(~Llanvirnian), ‘n3’(~Llandeilian the Middle Ordovician. to middle Ashgillian) and ‘n4’ (upper Ashgillian). The palaeobiogeographical puzzle is even more Each corresponded to a major sedimentary cycle, complex when migration routes are considered. The and two of them also subdivided into substages first one went eastwards (in present coordinates), (‘n1a–c’, ‘n3a–c’). Although these informal units probably favoured by oceanic currents similar to were used by Legrand (1985), they were never reused today’s Antarctic Circumpolar current, dispersing or revised by other authors. A recent review by macrobenthos larvae from South America to south Gutierrez-Marco & Martin (2016) shows that the polar Gondwana and the Middle East. A second Lower Ordovician graptolite succession from North migration route is suggested by faunal interchange Africa, in spite of some endemisms seems to be per- with Asian faunas dispersing from temperate areas fectly correlatable with the generalized zonal located to the modern-day East, which could have sequence for Avalonia–Baltica and Andean Gond- been accelerated by currents similar to today’s sub- wana, and the same applies to Middle Ordovician polar currents (‘east wind drift’) and eventually records of this fossil group. On the other hand, the reaching the Andean area of South America in the brachiopod and trilobite faunas from Algeria also Upper Ordovician, mixing those faunas with the show the same ‘Mediterranean’ character as their ones from the ‘Celtic’ Province. Additionally, there equivalents in high Ordovician palaeolatitudes. is the poleward invasion across Gondwana of In a general palaeogeographical analysis of numerous hitherto Laurentian, Anglo-Baltic and Ordovician strata across the ‘Tethyan’ region, South China–Kazakhstanian endemic taxa of trilo- Spjeldnaes (1967) proposed a new chronostrati- bites, brachiopods and bryozoans (Hammann 1992; graphical scheme for the Mediterranean region of Hammann & Leone 1997, 2007; Jimenez-Sanchez & SW Europe and North Africa, due to the general Villas 2010; Colmenar 2015) as a consequence of the correlation problems with the highly endemic ben- faunal shift brought on by the global warming of the thic faunas of the British Ordovician. He introduced Boda Event (Fortey & Cocks 2005). Nevertheless, the the series names ‘Zagorian’ – named after the city of migrating warm-water taxa that reached their maxi- Zagora in the Moroccan Anti-Atlas; ‘Morgatian’ – mum diversity in carbonate facies did not spread after the Armorican port of Morgat, westernmost uniformly across the region. This was the case of France; and ‘Tafilaltian’ – after the Tafilalt region in Bohemia and the inshore siliciclastic shelf areas near the eastern Anti-Atlas. The Zagorian series com- the South Pole, which retained their typical poorly prised all the strata known to that date assigned to diversified brachiopod and calymenacean–dalmani- the Tremadoc–Arenig series in Morocco and SE tacean trilobite assemblages with very little input France. It was characterized by endemic species of from cosmopolitan faunas (Destombes et al. 1985; trilobites and echinoderms, showing some relation- Havlıcek 1989). ships with Bohemia, and graptolites of Atlantic In any case, the attempts to reach a consensus in affinities. The Morgatian series included the wide- the reconstruction for the puzzling South Gondwana spread fossiliferous shales bearing the ‘Neseuretus palaeobiogeography will extend well into the future, tristani fauna’ plus its predecessors. It ranges from requiring additional detailed studies for which the the upper Arenig to lower Caradoc, being essentially Bohemo-Iberian regional scheme represents a more an equivalent to the Llanvirn–Llandeilo series of precise chronostratigraphical frame for regional cor- Great Britain. Spjeldnaes (1967) observed that, in relation that presently provided by the Global spite of problems with the definition of its lower and Ordovician standard scale. upper boundaries, Morgatian fossils have a distinc- tive character, consisting of low diversity assem- blages of trilobites (mainly calymenids and Historical perspective on regional phacopids), bivalves, gastropods, brachiopods and chronostratigraphy ostracods, which occur not only in Europe and Africa but also in Syria and presumably also Florida Legrand & Nabos (1962) were the first authors to and South America (Spjeldnaes 1961). The same propose a regional scale for the Ordovician of the author added the record of some Ordovician LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 265 trilobites from China that, in his opinion, revealed the only ones that locally present good international undoubted Morgatian affinities at temperate to correlation potential. The global Tremadocian stage tropical palaeolatitudes (Spjeldnaes 1967). The is succeeded by a first regional stage, named as Tafilaltian series was tentatively correlated with Arenigian after the original Arenig series of Britain. lower Caradocian to middle Ashgillian strata. Its This secured the widespread use of the Arenigian typical fauna is dominated by bryozoans (e.g. (sic) in publications over the Mediterranean area Chasmatoporella), brachiopods (Svobodaina, Poram- since the 1960s, and also follows the opinion (Cocks bonites and Nicolella), cystoids and some trilobites et al. 2010, p. 174) that the former British ‘series’ (Calymenella, Kloucekia and trinucleids), and is might be better used now as regional stages for Aval- recognized in North Africa, SW Europe and Turkey. onian and Gondwanan stratigraphy. However, the The conceptual value of the regional series main reason to justify the use of this British-adapted proposed by Spjeldnaes (1967) was questioned by stage for the rocks spanning the Lower and Middle Destombes (1971), who pointed out the imprecise Ordovician boundary in the south-Gondwanan definition of its boundaries and the better possibili- regional scheme is the generalized absence of valuable ties of correlation with the Bohemian faunas. As a correlation criteria to identify the Floian, Dapingian consequence, the names for these ‘Mediterranean’ and lower Darriwilian stages of the global scale. This series were regarded as superfluous, because the cor- regional and pragmatic use of the Arenigian stage for responding strata were roughly equal to the so-called the pre-Didymograptus artus graptolite biozone strata lower, middle and upper divisions of the Ordovician has been justified thus far by the shared palaeobio- already used for a large area of SW Europe and geographical links prior to the complete splitting of North Africa. However, this anomalous triple divi- Avalonia from southern Gondwana, probably fin- sion corresponds to never-defined units – of subsys- ished by late Arenigian times (=Dapingian to early tem or series rank? – absent in the British Darriwilian), but persisting for some time in the dis- Ordovician scale. The ‘local adaptations’ of the Bri- tal shelf environments of Avalonia before the sub- tish terminology in Mediterranean countries are also stantial broadening of the Rheic Ocean. evident in the use of the original British series as The Avalonian–Gondwanan Arenigian stage is fol- ‘stages’ (Llanvirnian, Caradocian, Ashgillian, etc.). lowed by the four Bohemo-Iberian regional stages The misinterpretation reached a state where wide- (Oretanian, Dobrotivian, Berounian and Kralodvo- spread facies such as the Armorican Quartzite were rian) established for the remaining Middle to Upper dated as ‘Skiddawian’, a name commonly recorded Ordovician strata. The regional Ordovician scale in the Ibero-Armorican – plus some Moroccan and ends with the global Hirnantian stage, equivalent to Bohemian – literature and maps, in spite of the fact the former Kosov regional series (Havlıcek & Marek that this alleged British stage was apparently never 1973) or Kosovian regional stage (Fatka et al. 1995) applied to the Skiddaw Group (Williams et al. 1972; of Bohemia, as well as to the British Hirnantian ? Cooper et al. 2004). Even today, the chronostrati- substage if the regional Ashgill series is given the graphical terminology based on the historical type rank of stage (Cocks et al. 2010). areas of the British Ordovician is being followed in Spain and Portugal by some authors and university groups and in many current geological maps; see Sa The Bohemo-Iberian regional scale et al. (2010) for a Portuguese illustration of this practice. Not only by misusing informal series did This regional scale is based on the broad biostrati- these turn into ‘stages’ and non-existent units, but graphical distribution of the Ordovician shelly fau- also many local authors still followed the first stan- nas of ‘Mediterranean type’ in the extensive shelfal dardizations of the scale (e.g. Williams et al. 1972; strata of South Gondwana, combined with a good Harland et al. 1982) instead of the modern British palynological record. Sporadic occurrences of con- chronostratigraphy adopted by British authors odonts and graptolites, as well as the faunal inter- (Fig. 1). change observed with Baltic or Avalonian regions, A more accurate Mediterranean (or south-Gond- allow some correlation with those other regional wanan) chronostratigraphical scheme was later schemes and also provide indirect ties for correlation developed in Bohemia (Havlıcek & Marek 1973) and with the global scale. completed in Iberia (Gutierrez-Marco et al. 1995, Modern concepts of a Mediterranean scale began 2002a, 2008, 2016). Its historical development is with Havlıcek & Marek (1973) who proposed new summarized in Fig. 2. The resulting regional scale chronostratigraphical names as partial replacement (Fig. 3) extends from the top of the Tremadocian to of some traditional British series to improve their the base of the Hirnantian. These global stages are potential for more accurate and precise regional 266 Gutierrez-Marco et al. LETHAIA 50 (2017)

Lipold & Kettner (1916) Havlíč ek & Krejčí (1860) Kettner & Kody m (1919, pars) Bouč ek (1937, pars) Marek (1973 ) Barrande Fatkaet al . MODERN Krejčí Bouč ek (1928, pars) Havlíč ek (1961) [Př9 ibyl (197 )] (1852) (1995) USAGE (1860-1863, Šuf & Prantl (1946, pars) HavlíčVan ek & ek (1966) Havlíč ek & 1877) Kettner & Prantl (1948) Fatka (1992)

Kosov Q B dζ Hirnantian St Kosov Q 2 Kosov Fm. Kosov Sr Kosovian St (GS) d5 Králův Dvů r ζ Králův Dvů r Kralodvorian Kralodvorian Králův DvůS r d 1 Králův DvůFm. r (Köningshofer) S Zdice (Králodvor) Sr St St Bohdalec ? Bohdalec S dε Bohdalec Fm. upper 2c Substage d4

Zahoř any S ice B ε Chlustina B d 2b ZahořFm. any č Loděnice middle Vinice S Nu ε Substage

d3 řB d Vinice Fm. =Trubín S Černín B 2a Berounian Beroun Sr Berounian ε Letná Fm. St St Zaho any Letná B d 1b ? Chrustenice Libeň S

nice B ε d2 Libeň S d 1a lower Chruste- Substage Libeň Fm. Drabov Q Drabov Q dδ Rěvni- ce Q γ Dobrotivá S Svatá Dobrotivá d 2b

tage

? ň Dobrotivá Fm. Dobrotivian Dobrotivian Dobrotivá Sr St St Skalka Q dγ Skalka Rokycany S 2a Q = Osek and Llanvirn Sr ilurian S γ Llanvirnian Oretanian St Šárka Bd 1 Šárka Fm.

S (Second D fauna) Kváň S Osek-Kvá B [St]Šárka St d1 Komárov B dβ Komárov B Arenig Sr Arenigian Arenigian St Klabava Fm. (As) α [Klabava St] St Olešná B d 3 Krušna ora B Mílina B dα Mílina Fm. hora G 2 Tremadoc Tremadocian St Tremadoc Sr α St (GS) Tř enice G d 1 TřFm. enice

Krušná H

Fig. 2. Chronological development of the Bohemian chronostratigraphical and lithostratigraphical terminology, and correlation with the modified Bohemo-Iberian regional scale for South Gondwana (right column). Lithostratigraphical correlation mainly based on reviews by Havlıcek & Marek (1973) and Chlupac (1999). Abbreviations: AS, Avalonian stage; Fm, formation; GS, Global Stage; B, beds; G, grey- wacke; Q, quartzite; S, Shale; Ser, Series; St, Stage. [Note: Figure 2 has been corrected after initial online publication.] correlation. The selected area for the new chronos- they are mainly exemplified by the more complete tratigraphical units was the Prague Basin, which has and better-explored Ordovician successions in Bohe- a long history of research (Fig. 3) and where the mia, the Iberian and Armorican Massifs, the Iberian Ordovician sedimentary sequence is exceptionally Range, Montagne Noire and Sardinia. As an exam- complete, continuous and fossiliferous, being almost ple, Fig. 4 shows a correlation chart with the local entirely developed in siliciclastic rocks ranging from biozones proposed for the Middle and Upper shallow to moderately deep-water facies. Later modi- Ordovician strata in the Ibero-Armorican area. The fications turned the regional series into regional remaining south polar or peri-Gondwanan areas stages (Fatka et al. 1995), and the incorporation of reached by the Mediterranean facies and faunas are the Oretanian stage – named after a historical region commented upon separately. in central Spain – as a replacement of the Llanvirn series of Britain in its traditional usage (Gutierrez- Tremadocian Marco et al. 1995). The former was proposed to avoid the conceptual instability of the British Llan- The true base of this global stage cannot be recog- virn from 1970 to 1990, when it changed first to nized in any section of the South Gondwanan stage (Llanvirnian) of a redefined Llandeilo series, Ordovician, where the oldest rhabdinoporinid grap- then recovered its previous status as a series but tolites recorded from North Africa, Sardinia and within a ‘Dyfed subsystem’, and finally was com- Turkey indicate an early, but not earliest, Tremado- pletely redefined as a series that embraces the Llan- cian Age (Gutierrez-Marco et al. 2015; Gutierrez- deilian stage, being the new Abereiddian stage an Marco & Martin 2016). Tremadocian strata fre- equivalent of the traditional Llanvirn series (see quently rest disconformably upon Cambrian rocks Gutierrez-Marco et al. 2016, fig. 1). and occur in thick volcanosedimentary and metavol- The main characteristics defining the stages of the canic complexes related with the opening of the Bohemo-Iberian scale are summarized below, and Rheic ocean (e.g. the ‘Ollo de Sapo’ belt), or did not LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 267

GLOBAL STAGE TS IBERO-BOHEMIAN BRITISH-AVALONIAN STAGE SLICE 2004 REGIONAL SCALE REGIONAL SCALE Hi2 HIRNANTIAN 6c HIRNANTIAN HIRNANTIAN Hi1 6b Ka4 6a RAWTHEYAN KRALODVORIAN ASHGILL Ka3 CAUTLEYAN KATIAN 5d PUSGILLIAN Ka2 u STREFFORDIAN Ka1 5c CHENEYAN BEROUNIAN m Sa2 5b SANDBIAN BURRELLIAN l Sa1 5a CARADOC AURELUCIAN u ? DOBROTIVIAN l . LLANDEILIAN Dw3 4c u ORETANIAN ABEREIDDIAN DARRIWILIAN Dw2 4b l LLANV Dw1 4a

Dp3 u FENNIAN 3b DAPINGIAN Dp2

Dp1 3a ARENIGIAN m WHITLANDIAN Fl3 2c ARENIG FLOIAN Fl2 2b l MORIDUNIAN Fl1 2a 1d Tr3 u 1c MIGNEINTIAN TREMADOCIAN Tr2 1b TREMADOCIAN m Tr1 1a l CRESSAGIAN

TREMADOC

Fig. 3. Chart showing the proposed correlation between the global stages, stage slices (Bergstrom€ et al. 2009), time-slices (TS: Webby et al. 2004), and regional chronostratigraphical units recognized in the Bohemo-Iberian scale. On the right, a tentative correlation with the present British–Avalonian regional scale. Adapted from Gutierrez-Marco et al. (2016). even get deposited on the large emerged areas that and echinoderms that later spread to Baltica, Asia extended across the Mid-North Armorican–Central and South America. Among these is the brachiopod Iberian domain (Sa et al. 2014). genus Tarfaya, supposedly endemic to Morocco, In Bohemia, Tremadocian strata start with trans- which has been recently recognized in northern Iran gressive sandstones with faunas containing large lin- and Argentina (Benedetto & Munoz~ 2016). Another gulate brachiopods (Hyperobolus and Westonisca good example of affinities, in this case at a species associations) and sparse rhynchonelliforms level, between similar faunas described from Mon- (Poramborthis assemblage). A subsequent deepening tagne Noire, is Taihungshania miqueli that also introduces the illaenid–cheirurid and nileid trilobite occurs in the upper Tremadocian of Sardinia, the biofacies, corresponding to the Hemibarrandia– Alborz Mountains of northern Iran, the Taurides in Parapilekia and Proteuloma–Ceratopyge assemblages, Turkey, and probably in younger beds of the United respectively (Fatka & Mergl 2009). The middle to Arab Emirates. Scarce records of South Gondwanan upper Tremadocian faunas of Bohemia, Bavaria, olenid trilobites occur in the Iberian Range and Montagne Noire, Iberian Range, Morocco and west- Montagne Noire, but they are not representative of ern Mauritania are complemented by old calymen- the true deep-water olenid black shale biofacies. acean and rare cyclopygid taxa and include few Conodonts are scarce with enigmatic assemblages endemic taxa. Their biogeographical affinities, such as the ‘franconicus fauna’ of Bavaria and mostly at generic level, indicate an active interchange Brittany or special genera like Hammannodus and with similar trilobite and brachiopod faunas Teridontus. The former was recorded from the Palto- recorded of Baltica, South Urals and the Central dus deltifer biozone of the Montagne Noire (Serpagli Andes, resulting in that South Gondwana is the et al. 2007), NW Argentina (Voldman et al. 2013) source of certain faunas of trilobites, brachiopods and a single Spanish locality (Sarmiento et al. 2011), 268 Gutierrez-Marco et al. LETHAIA 50 (2017)

Fig. 4. Example of schematic correlation of the main biozones for the Middle and Upper Ordovician strata in Ibero-Armorica (updated from Gutierrez-Marco et al. 2002a), with regard to the Bohemo-Iberian regional scale. DAP, Dapingian; HIRN, Hirnantian. Abbrevia- tions for generic assignments are as follows: A – Aegiromena;Ac– Acanthochitina;Ak– Akidograptus;An– Ancyrochitina;Ar– Archicli- macograptus; Arm – Armoricochitina;B– Belonechitina;C– Coplacoparia;Ca– Cacemia; Cek – Cekovia;Co– Corymbograptus;Cr– Crozonaspis; Croz – Crozonorthis;Cy– Cyathochitina;D– Didymograptus;De– Desmochitina; Dean – Deanaspis;E– Eodalmanitina;Eu – Euconochitina;Ex– Expansograptus;G– Gymnograptus; Gel – Gelidorthis;H– Hustedograptus; Het – Heterorthina; Hold – Holdenia; Hy – Hyalochitina;L– Lagenochitina;La– Laufeldochitina; Lep – Leangella (Leptestiina); Lin – Linochitina;M– Morgatia;Ma– Mar- rolithus; Met – Metabolograptus;N– Neodiplograptus;O– Oepikograptus;Ox– Oxoplecia;P– Placoparia;Ps– Pseudamplexograptus dis- tichus;S– Siphonochitina; Siv – Sivorthis;Sp– Spinachitina; Svob – Svobodaina;T– Tanuchitina; Ten – Tenuiseptorthis;Vy– Vysocania; Z – Zeliszkella. whereas the Tremadocian species Teridontus gallicus including Montagne Noire (Serpagli et al. 2008). reaches an even wider distribution in numerous Preliminary conodont data from the Fezouata Early Ordovician localities around the world, also Formation of Morocco range from the late LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 269

Tremadocian Paroistodus proteus biozone to the diverse in the Iberian Range, Montagne Noire and lower Floian Prioniodus oepiki biozone (Lehnert Moroccan Anti-Atlas, starting from the basal Euloma et al. 2016). Five consecutive graptolite biozones in filacovi trilobite biozone shared by the two-first the Fezouata Formation span from the mid-lower to areas. Several trilobite biozones have been proposed the uppermost Tremadocian (Gutierrez-Marco & for the Floian of Montagne Noire, which exhibits Martin 2016). The majority of the recognized species transgressive–regressive trends reaching the Neseuretus are cosmopolitan, but some genera were restricted to biofacies of inshore environments (Vizca€ıno & shallow-water shelf environments in high-latitude Alvaro 2003). In the Fezouata Formation of Mor- regions (Kiaerograptus, Choristograptus, Bryograptus). occo, there is a general regressive trend among the The upper Tremadocian Araneograptus murrayi Arenigian trilobite assemblages, from the raphio- biozone bears a particularly diverse assemblage that phorid to Colpocoryphe and Neseuretus biofacies, is widely distributed in South Gondwana (Morocco, with many taxa in common with the Montagne Mauritania, Algeria, Spain, SE France, Sardinia, Noire (Vidal 1998a,b; Martin et al. 2016b). The Germany) and dates the basal beds of the famous same affinities are observed among the echinoderm Gondwanan Fezouata Konservat-Lagerst€atte (Martin and bivalve faunas of Morocco and SE France (e.g. et al. 2016a). The Tremadocian trilobite faunas are Polechova 2016). rather uniform across much of Gondwana, that is in Although a general ‘Mediterranean character’ is Wales, Montagne Noire, Iberian Ranges, Anti-Atlas, recognized among numerous macrofossil and paly- China and eastern Australia. But some offshore set- nomorph groups in the South Gondwana areas, the tings (Montagne Noire, Iran) contain tropical taxa Avalonian Arenigian stage is used because of the (Carolinites, Peltabella) that possibly followed warm- presence of Anglo-Baltic taxa in a region which was water currents. still attached to Gondwana at least during the Floian, sharing the same problems of correlation with the Arenigian global scale, in particular for the upper Arenigian. The Floian graptolite sequences from the Lake This regional and loosely defined stage of the British District, the Moroccan Anti-Atlas, and the Mon- classification is represented by a widespread trace tagne Noire allow, at a local level, to estimate the fossil-rich, shallow-marine sandstone (Armorican extent of the international stage slices Fl1 to Fl3 with Quartzite facies) deposited on a wide shelf in Ibero- reference to the Baltic scheme (Gutierrez-Marco & Armorica. Commonly, this unit and its basal forma- Martin 2016). In the Ossa-Morena Complex of SW tions rest unconformably on Ediacaran to Cambrian Spain, the base of the Tetragraptus pyllograptoides rocks, but sometimes grades down or extends later- biozone can be used, as it occurs in some Baltic suc- ally into other shale/sandstone-dominated Tremado- cessions, as correlatable to the FAD of Tetragraptus cian–Arenigian sequences as in SE France, NW approximatus, defining the base of the Floian stage Iberia and the Iberian Range. In Morocco, Pyrenees, in most deeper-shelf-to-slope assemblages. But the Sardinia and Bohemia the Arenigian strata seem to scarce record of significant conodonts over the entire be continuous with the Tremadocian, being pre- area (Lehnert et al. 2016), and the predominance of dominantly shaley and very fossiliferous. Special bio- shelly faunas of limited value for a better appraisal of facies of giant lingulacean brachiopods preserved in the global stages force us to continue to use the old sandstones (Ibero-Armorica, Serbia, Algeria, Libya, ? British ‘Arenig’ as the preferred regional stage. To Saudi Arabia) are restricted to South Gondwana, but complete the Bohemian scale, promoting a replace- equivalent inner-shelf assemblages from Bohemia ment for the Arenigian, Pribyl (1979) used the terms (the Leptembolon–Thysanotos fauna) reveal palaeo- ‘Klabava stage’ and ‘Klabavian’ in a paper dealing biogeographical connections with the upper Tre- with Ordovician ostracods, but it lacks a precise madocian–lower Arenigian more temperate deposits definition. of Baltica, south Urals, northern Iran and the central The main stratigraphical problem for the Mediter- Andes. ranean and British Arenigian is the correlation of Rich trilobite assemblages from the Arenigian of those strata to the global Dapingian stage and the Bohemia display equivalents to the illaenid–cheir- lower Darriwilian stage slice. This interval is poorly urid (Pliomerops association), nileid (Asaphid and represented in South Gondwana, only recorded by Euloma associations) and cyclopygid biofacies three chitinozoan biozones from which the older (Mergl et al. 2007; Fatka & Mergl 2009), indicating ones (Desmochitina ornensis and Belonechitina henryi some connections at generic level with Baltica and biozones) lie above, or can be part of, a generalized other peri-Gondwanan terranes from temperate stratigraphical gap recognized in Ibero-Armorica, palaeolatitudes. Arenigian assemblages are also Morocco, Algerian Sahara, ?Bohemia, the Arabian 270 Gutierrez-Marco et al. LETHAIA 50 (2017)

Peninsula and extending as far as SE Iran (Videt stage/Llanvirn series (Fig. 3). The ‘Sarka series’ (or et al. 2010; Ghavidel-Syooki et al. 2014; Gutierrez- ‘Sarkian’/’Sarkan’ stage) of Pribyl (1979), although Marco et al. 2014a). The eustatic nature of this gap provisionally mentioned in a study on Bohemian is starting to be debated but according to the model ostracods, has never been substantiated as an inde- by Rasmussen et al. (2016) a possible explanation pendent regional unit. may be the onset of a sudden Mid-Ordovician Ice The Oretanian is recognized by the sudden appear- Age that presumably triggered the Great Ordovician ance of middle Darriwilian pendent Didymograptus Biodiversification Event (GOBE). Although a long on South Gondwana, Avalonia and Baltica within cooling trend through the Ordovician has been sug- transgressive graptolite shales that overlie shallow- gested as an alternative to the traditional greenhouse water facies – or temporary emerged areas – that had model interrupted by the abrupt Hirnantian glacia- been common previously. This transgression is tion (Nardin et al. 2011; Dabard et al. 2015), the recorded as diachronic and started in the late Areni- sea-level drop of about 150 m at the beginning of gian (late Dapingian–early Darriwilian) in SW Eur- the Middle Ordovician proposed by Rasmussen ope, Morocco and Bohemia, where graptolite shales et al. (2016) is not supported by the onshore to with declined, deflexed or horizontal didymograptids nearshore stratigraphical record on North African (plus Azygograptus and early biserials) are succeeded Gondwanan areas that are closest to the South Pole. by pendent Didymograptus (subgenera Didymograp- But a big enigma exists as Dabard et al. (2015) sug- tus and Jenkinsograptus). The relative immersion of gested the existence of seven rapid episodes of signifi- the SW European–north African region produced cant sea-level fall (50–80 m) in the Darriwilian shelfwards spreading of assemblages with trilobites sequential architecture of the western Armorican and brachiopods of older Avalonian origin (e.g. Massif. This marks the onset, in the late Arenigian, of Placoparia cambriensis, Paralenorthis alata), but also icehouse conditions with multiple inferred glacio- stimulated the endemic radiation of dalmanitacean eustatic cycles and distinct eustatic events related and calymenacean trilobite faunas, as well as orthid to Middle Ordovician glaciations. Previously, a and harknessellid brachiopods (Gutiorthis, Sivorthis, ‘Darriwilian Ice Age’ was also proposed for southern Almadenorthis, Cacemia), plus the earliest Jordan (Turner et al. 2012) based on the indirect evi- strophomenoids. The transgressive event continued dence provided by sequence stratigraphy. According until the late Oretanian, when the shales with Didy- to these authors, ‘the absence of glacio-terrestrial mograptus murchisoni penetrate interior platform deposits of Middle Ordovician age in Southern Gond- areas of the Arabian Peninsula and all the previously wana is most likely due to subsequent glacial erosion emerged areas of Ibero-Armorica. during repeated ice advances, up to and including The Oretanian assemblages of Bohemia are those in the Hirnantian’ (Turner et al. 2012, p. 251). dominated by atheloptic trilobites (Placoparia– After the widespread Dapingian–early Darriwilian Euorthisina assemblage) with common illaenids, sedimentary gap and condensed facies, prominent asaphids, dalmanitids and cyclopygids, and show and widespread flooding events took place on a transitional shifts towards relatively shallower or regional scale, just preceding or post-dating the Des- deeper environments, depending of the subsidence mochitina bulla chitinozoan biozone. The latter char- rates on different parts of the Prague basin. Encom- acterizes the upper Arenigian strata both in South passing the transgression, bivalves became significant Gondwana and in the type Arenig area of Wales as they did simultaneously in other parts of the (Amberg et al. 2015). Faunal links with Baltica were South Gondwana platform, and the brachiopod still present in the late Arenigian/earliest Darriwilian, Euorthisina reached a wide distribution, extending represented by the record of the conodont Barran- into Morocco and South America. degnathus bohemicus in upper Volkhhovian lime- stone of southern Scandinavia, the pre-D. bulla beds Dobrotivian of Bohemia and as reworked elements preserved in upper Pridolian rocks of the western Moroccan The base of this stage does not coincide regionally Meseta (Stouge 2005 and references therein). with any significant changeover in facies or faunal development; instead, the base of the Dobrotivian is Oretanian defined by the disappearance of pendent didymo- graptids and their stratigraphical replacement by The name of this stage was proposed originally in certain biserial forms, and its top by the arrival of central Spain (Gutierrez-Marco et al. 1995 and refer- the typical Berounian Drabovia brachiopod fauna. In ences therein), being now roughly equivalent of the Ibero-Armorica the Dobrotivian coincides with the Abereiddian? substage of the British Llanvirnian diversity peak reached by the Neseuretus biofacies LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 271 and with an active period of faunal interchange with brachiopod genera, which are also represented by Bohemia. In the latter, the shale facies with slightly endemic species. Among the most significant taxa are deeper assemblages of trilobites and brachiopods many Mediterranean species of Svobodaina, Tafilaltia, predominate. The upper Dobrotivian beds are char- Tissintia, Heterorthis, Tazzarinia, Gelidorhis, Pauci- acterized by the abundance of the trilobite Placo- crura, Drabovinella, Hirnantia, Dalmanella, Jezercia, paria (Coplacoparia) borni, also recorded from Destombesium, Reuschella, Porambonites, Saukrod- Bohemia. The species coexists in Ibero-Armorica ictya, Howellites, Iberomena, Kjaerina (subgenera with the last representatives of the genus Neseuretus, Kjaerina and Villasina), Rafinesquina (Mesogeina), in beds locally referred to the Marrolithus bureaui Hedstroemina, Longvillia, Triplesia, Bicuspina, Rostri- subzone in France and Morocco. The uppermost cellula and Irhirea. The Berounian trilobite fauna is Dobrotivian strata are already of Late Ordovician also very characteristic, with abundant genera as (early Sandbian) age as indicated by the earliest Dalmanitina, Eudolatites, Deanaspis, Marrolithus and record of Oepikograptus bekkeri, a ‘Baltic’ graptolite Declivolithus, and distinctive species of Colpocoryphe, typical from the Nemagraptus gracilis biozone, Calymenella, Eccoptochile, Flexicalymene, Pri- already known – as ‘Lasiograptus? inexpectatus’ – in onocheilus, Dreyfussina, Zeliszkella, Kloucekia, Sokretia, the upper Dobrotivian of Bohemia. Scarce and Hispaniaspis, Selenopeltis and Birmanites, among poorly preserved conodonts from upper-lower to others. Some echinoderms are widely distributed in upper Dobrotivian strata of Ibero-Armorica were the Berounian, especially cystoids such as Aristocys- loosely correlated with those from the boundary tites, Destombesia, Rhombifera, Mahrebocystis, Arach- between the Pygodus anserinus–Amorphognathus nocystites and ‘Echinosphaerites’. The Berounian stage tvaerensis Baltic biozones (Sarmiento et al. 2011). was originally subdivided in Bohemia into two or Among the Dobrotivian brachiopods, the most three ‘substages’ (Fig. 2). An informal tripartite divi- distinctive feature is the Gondwanan origin and sion into lower, middle and upper Berounian is being expansion reached by the heterorthids (Havlıcek used in Iberia and Morocco because of the correlation 1976), which occur in Morocco, Algeria, Bohemia, of some species with their stratotypes in Bohemia. Ibero-Armorica, Iran and the central Andes. The Thus, the assemblages from the Bohdalec Formation early record of the genera Aegiromena and Jezercia in (‘upper Bohdalecian’) show recognizable taxa and the Dobrotivian of Spain locally predates the onset lithologies (e.g. the Polyteichus facies) that favoured of the typical Berounian brachiopod faunas. its separation from older assemblages with long- ranging taxa, commonly persisting from the Vinice Berounian and Zahorany formations (‘middle’ Berounian). The ‘lower’ Berounian is well developed in Mor- The Berounian stage is characterized by the develop- occo, but in SW Europe it is mainly represented by ment and geographical spreading of the distinctive its lower strata, because higher beds were not depos- Aegiromena–Drabovia brachiopod fauna (Havlıcek ited or were eroded in what is known as the wide- 1982, 1989) and associated trilobites over much of spread ‘pre-Vinician gap’. It could have been related the Mediterranean area. In spite of the general pau- to the Sardic movements (Hammann 1992; city of graptolites in the ubiquitous sandy and silty Gutierrez-Marco et al. 2002a). facies that dominate in this stage, its lower and The Berounian/Kralodvorian boundary is easy to upper boundaries occur within graptolite biozones, recognize in Bohemia by a sudden replacement in making its precise international correlation very dif- benthic faunas related to a changeover in lithology. ficult. On the other hand, Oepikograptus bekkeri The Aegiromena–Drabovia association abruptly dis- (indicative of the N. gracilis zone) is still present in appeared, being replaced by more cosmopolitan taxa the basal Berounian strata of Portugal (Carregueira with many immigrants from Avalonia and Baltica. Formation), while the top of the stage (=the base of The same faunal shift is observed in many other areas the overlying Kraluv Dvur Formation) was corre- of South Gondwana, with the relative exception of lated by Kraft et al. (2015) with the Pleurograptus the Moroccan assemblages from closer to the Pole, linearis zone. According to the old regional schemes, where they retain part of their ‘Mediterranean char- the Berounian is equivalent to most of the ‘Tafilal- acter’ until the onset of the Hirnantian glaciation. tian’ stage (Spjeldnaes 1967) that also includes the Algerian substage ‘n3b’ (Legrand 1985). Kralodvorian The rich Aegiromena–Drabovia brachiopod fauna is mainly characterized by a great diversity of genuine The beginning of the Kralodvorian stage coincides Gondwanan endemic taxa that numerically predomi- with a significant deepening in the Prague Basin, nate over a small proportion of cosmopolitan which included the arrival of the globally distributed 272 Gutierrez-Marco et al. LETHAIA 50 (2017)

Foliomena Fauna in deep-water environments with evidence of the arrival of such immigrants, probably common cyclopygid and remopleuridid trilobites. because no optimal conditions for the settlement of An episode of sea-level fall recorded in the upper these trilobites, as well as of the Hirnantia Fauna, part of the Kraluv Dvur Formation favoured the were reached. The very shallow onshore shelf envi- deposition of a thin carbonate-rich horizon (the so- ronment with cool water due to its proximity to the called Pernık Bed, only 5–20 cm thick), which is South Pole was probably the main limiting factor. In immediately overlain by a bed with the first Hirnan- spite of this, the Moroccan record of Kralodvorian tian graptolites. strata is confirmed by the vertical extent of Mediter- Over extensive areas of the South Gondwana plat- ranean taxa in the Upper Ktaoua Formation (the form, the Kralodvorian strata are palaeontologically Mucronaspis termieri–Flexicalymene ouzregui assem- recognizable by the poleward faunal shift of warm- blage, not restricted to a single noduliferous ‘horizon’ water taxa of former Anglo-Baltic, Asiatic or Lauren- as suggested by Loi et al. 2010). In its topmost part, tian origins, which was caused by the Boda Event of this formation yielded Dreyfussina struvei, a wide- global warming (Fortey & Cocks 2005). It also spread Kralodvorian trilobite (Hammann & Leone brought the widespread deposition of limestones and 2007), as well as Amphitryon, a rare ?nektonic trilo- the development of carbonatic build-ups over wide bite known from the Dicellograptus laticeps biozone areas previously dominated by siliciclastic facies, as it of Bohemia and ?Sardinia (Hammann & Leone 1997; occurs in SW and central Europe, the Ghadamis Basin Budil et al. 2011) and the illaenimorph Cekovia aff. of Libya and the eastern Tafilalt region of the Moroc- loredensis (E. Villas, J. Colmenar, J.C. Gutierrez- can Anti-Atlas (Villas et al. 2002; Boucot et al. 2003). Marco, D.C. Garcıa-Bellido, S. Lorenzo, S. Pereira & Two conodont biofacies from the Amorphognathus J.J. Alvaro, unpublished data). ordovicicus biozone occur commonly in these lime- Correlation problems in recognizing the Berou- stones (Ferretti et al. 2014 with complete references) nian/Kralodvorian and Kralodvorian/Hirnantian that usually are correlated with the Ka 3–4stageslices. boundaries in some parts of South Gondwana forced Interestingly, some of the Kralodvorian limestones, Kraft et al. (2015, p. 201) to describe the Kralodvo- which in Iberia may reach up to 300 m in thickness, rian as a loosely defined regional stage, although rest disconformably on older Arenigian to upper Ber- acknowledging that it represents ‘a specific succes- ounian beds (Gutierrez-Marco et al. 2002a). sion of faunal associations that reflects distinct con- The invasion of the high-latitude Gondwanan ditions widespread in the peri-Gondwanan Europe platform by a diverse group of temperate to warm- of that time’. The Berounian/Kralodvorian boundary water brachiopods of Avalonian and Baltic origin is is sometimes difficult to recognize in the South represented by the Nicolella Fauna both in the shal- Gondwana European platform because the arrival of low-water clastic and carbonate facies from the very many brachiopod taxa occurred in several migratory beginning of the Kralodvorian. The most characteris- ‘pulses’, the first of which as early as the late Berou- tic elements of the assemblage are the genera Nico- nian. Those taxa proliferated in the Nicolella Fauna, lella, Dolerorthis, Eoplectodonta, Platystrophia, as in the case of the Gabian Formation of the Skenidioides, Leptestiina, Cremnorthis, Rhactorthis Montagne Noire (SE France), where Nicolella and Vellamo, which are often present in older strata actoniae itself and many other species apparently of other palaeocontinents. They usually mixed, bio- co-occurred in the same beds with typical upper logically or taphonomically, with Mediterranean taxa Berounian taxa. The taxonomic list compiled by Col- thriving in shallower environments, and displaced menar et al. (2013, 2014) and Colmenar (2016) them until their almost complete extinction (Colme- includes Svobodaina, Kjaerina (K.) gondwanensis, K. nar 2015; with earlier references). Also as a conse- (Villasina) pedronaensis, Aegiromena meneghiniana, quence of the Boda Event, a more cosmopolitan and and the Berounian trilobite Dalmanitina (D.) acuta diverse trilobite fauna arrived in the Gondwanan (= ‘D. socialis proaeva?’). A similar overlap has been platform, becoming abundant in the limestone and observed with Berounian taxa occurring with ele- marly offshore facies of shallow to moderate depth. ments of the Nicolella Fauna in the lower part of the The assemblage includes originally Laurentian trilo- Porto de Santa Anna Formation of Bucßaco (central bite genera such as Heliomera or Diacanthaspis,or Portugal), in the top of the Fombuena Formation of Asiatic forms as Ovalocephalus (=Hammatocnemis), NE Spain, and in the lower beds of the Portixeddu Vietnamia (=Sarrabesia), Nankinolithus, Para- Formation of Sardinia (Colmenar 2015, 2016). In the phillipsinella and Taklamakania, which have been last unit, apart from the arrival of some warm-water recorded from Kralodvorian strata in NE Spain receptaculitids and dasycladales (Hammann & Ser- (Hammann 1992) and SW Sardinia (Hammann & pagli 2003), typical assemblages of upper Berounian Leone 1997, 2007). However, in Morocco there is no brachiopods (fossil horizons BH2 and BH3 in Leone LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 273 et al. 1991) are succeeded by the Nicolella Fauna and bracketed by the typical opportunistic assem- (ibid. horizon BH4). But Hammann & Leone (2007, blages of brachiopods and trilobites of worldwide p. 114) point out that in their equivalent trilobite distribution (the Hirnantia and Mucronaspis – ex horizons (TH2a, TH2b, TH 3) ‘it is not clear whether ‘Dalmanitina’ – faunas), which have their ancestors the Berounian/Kralodvorian boundary should be in peri-Gondwanan cool-water faunas. Earliest Hir- placed either with the appearance of Nicolella actoniae nantian graptolites are represented by Metabolograptus and Dalmanitina (Thuringaspis) lamarmorai or with ojsuensis (=M. extraordinarius of Mitchell et al. the disappearance of the Eudolatites flavus-Deanaspis 2011), recorded in Bohemia in the topmost beds of goldfussi fluminensis assemblage’. In the light of the the Kraluv Dvur Formation (Kraft et al. 2015). In joint record of brachiopod and trilobite associations, Spain, a similar ?pre-glacial graptolite horizon yield- it seems clear that the latter alternative makes more ing Neodiplograptus charis has been identified at the sense, as the arrival and settlement of the Nicolella base of the Chavera Formation (Mitchell et al. Fauna appears slightly diacronic and requires special 2011). M. osjuensis (and M. extraordinarius?) also ecological conditions. On the other hand, the Berou- occur intercalated with glaciomarine strata in Sar- nian trilobite and brachiopod associations were the dinia and Niger, whereas the late Hirnantian natural inhabitants of these shelf areas and were Metabolograptus persculptus zonal graptolite occurs abruptly replaced by the new arrivals from warmer in Bohemia, ?Morocco, Algeria, Mauritania, Jordan, waters when the Boda Event had already caused irre- ?Saudi Arabia, the Carnic Alps and the Spanish Pyre- versible changes in the benthic communities. In any nees. In the latter (unpublished data) the M. per- case, defining the exact position of the Berounian/ sculptus Biozone occurs in post-glacial shales and is Kralodvorian boundary will require additional bed- succeeded by the very rare ‘pre-Akidograptus ascen- by-bed collecting in areas like the Gabian Formation sus’ graptolite fauna. This is the third or fourth of Montagne Noire. This will clarify if the last Berou- Gondwanan record of this uppermost Ordovician nian brachiopods overlap with the base of the Amor- assemblage (Underwood et al. 1998; Loydell 2007; phognathus ordovicicus conodont biozone. Storch & Schonlaub€ 2012). Probably the same ‘inter- Finally, the stratigraphical position of the Kralod- regnum’ at high palaeolatitudes may be an equiva- vorian/Hirnantian boundary has been questioned lent to the uppermost Ordovician Spinachitina with reference to the ‘Pernık Bed’, which is situated oulebsiri–Euconochitina moussegoudaensis chitino- near the top of the Kraluv Dvur Formation of the Pra- zoan assemblage, recorded from northern Chad and gue Basin. This contains a diverse and distinctive SE Libya (Thusu et al. 2013). fauna of trilobites, strophomenate brachiopods, cys- Despite a sparse record, the Hirnantian shelly fau- toids, blastoids, bryozoans and other fossils, which are nas are represented in several Mediterranean coun- related with the highly diverse, warm-water assem- tries and allow for correlation of the former regional blages known in many parts of South Gondwana. This stage ‘n4’ (from Algeria) and the Kosov series or traditional interpretation had considered this bed as stage (from Bohemia) with the global Hirnantian Kralodvorian in age due to its supposed relation with Stage. The recent discovery of a considerably diverse a late influx in Bohemia of the Boda Event. But Mergl Hirnantia–Mucronaspis fauna in the Cantabrian (2011) reinterpreted this carbonate horizon as earliest Zone of NW Spain (Bernardez et al. 2014) argues Hirnantian, and linked it to the sea-level drop associ- against a palaeoclimatic–palaeolatitudinal distinc- ated with the first stages of the Gondwanan glaciation. tion between the ‘Bani’ (=very cool polar waters) The remarkable diversity and mixed occurrence of and ‘Kosov’ (from temperate to tropical areas) pro- Kralodvorian and Hirnantian odontopleurid trilobites vinces of the Hirnantia Fauna at a global scale (Har- in the ‘Pernık Bed’ (Diacanthaspis, Bojokoralaspis, Pro- per et al. 2013), indicating that other environmental ceratocephala, Chlustinia, Eoleonaspis) suggests that factors such as depth or the type of substrate also the Kralodvorian elements accumulated in the inner played a significant role. shelf during the Boda Event were transported basin- wards as bioclastic material by mudflows during a sea-level drop in the earliest Hirnantian. The hypothe- Regional faunas and correlation sis of Mergl (2011 and later work) has not yet received wide acceptance (Kraft et al. 2015). The first attempt to stratigraphically correlate between the Ordovician succession of Bohemia, with Hirnantian the Armorican and Iberian massifs was that of Barrande (1863) who used the shared record of the This global stage is mostly represented in the so-called Second fauna. Research since then has con- Mediterranean area by glacial to glaciomarine strata firmed the close palaeobiogeographical and facies 274 Gutierrez-Marco et al. LETHAIA 50 (2017) relationships between the Prague Basin and other NE Spain, northern Chad and northern Benin (Seila- Gondwanan or peri-Gondwanan areas lying at high cher 1970; Seilacher & Alidou 1988; Alidou et al. Ordovician palaeolatitudes. Thus, a shared regional 1991). Coeval strata have also contain ‘Cruziana’ al- chronostratigraphical scale is justified, taking into madenensis, another rare trace probably made by account the unsurmountable problems associated some kind of merostome that was described in Spain with the direct adoption of either the global standard and has since been found in several African and scale or the British–East Avalonian regional scheme. Middle East localities (Seilacher 1970, and research The integrated Ordovician biostratigraphy of in progress). South Gondwana is being increasingly improved and Additional evidence of Ordovician palaeogeo- refined for some areas such as Bohemia, Sardinia, graphical relationships with distant or non-outcrop- the Carnic Alps and Ibero-Armorica (Fig. 4), but ping areas is provided by fossils reworked from older work is still to be completed in Morocco, and data formations in nearby eroded regions or those are lacking from the remaining Mediterranean and recorded in dropstones embedded in marine diamic- Middle East countries. Thus, these areas were fre- tites of the Hirnantian glaciation. As an example, the quently excluded, or vaguely considered, in the ice-rafted fossiliferous ‘Geschieben’ occurring in the modern syntheses and palaeogeographical maps for Lederschiefer Formation of Thuringia (Hirnantian, the early Palaeozoic (e.g. Harper et al. 2013; Torsvik Germany) contain Berounian and Kralodvorian & Cocks 2013a,b; Colmenar 2015). trilobites, brachiopods, cystoids and ostracods with Only an extensive palaeontological knowledge of different South Gondwanan provenances (Schall- Ordovician strata in South Gondwana allowed for reuter & Hinz-Schallreuter 1998 and references advancement of palaeogeographical reconstructions, therein). And the specimens of Acrosaccus (ex such as the interpretation proposed, among others, ‘Orbiculoidea’) massai that occur in glacial drop- by Gutierrez-Marco et al. (2002a) for the Iberian stones in the Kosov Formation of Bohemia possibly terranes with a palaeoposition adjacent to present have a North African origin (Mergl 1994). Libya by the Ordovician, instead of their usual place- The usefulness of the Bohemo-Iberian regional ment NW of Morocco. This interpretation was later scale to accommodate with reasonable detail the confirmed by provenance studies of detrital zircons. Ordovician fossil record from South Gondwana also The main reason for the change in the palaeogeo- facilitates intraregional correlations and the integra- graphical interpretation was the common record of tive calibration among biostratigraphical scales pro- the trilobite Neseuretus tristani and other typical vided by different fossil groups (Fig. 4). The more Ibero-Armorican benthic taxa, which occur in coeval precise and complete regional scheme for the whole strata from the Ghadamis Basin of Libya and Tunisia area is provided by the ‘North’ Gondwana chitino- and from Saudi Arabia and Oman, with further zoan biozonation (Paris 1981 and later refinements), records still to be confirmed from western Macedo- which has been of special value for the dating and nia, ?Iran and the Ougarta Range, Algeria (Ghienne correlation of Ordovician rocks from numerous et al. 2007a). Also the problematic fossil Hanadir- boreholes made by the oil industry in North Africa ella, originally defined from Oretanian strata in and the Middle East. In addition, macrofossils have Saudi Arabia (El-Khayal 1985), was later recovered provided precise correlations of the Bohemo-Iberian from the subsurface of Algeria, expanding its records scale with previous regional divisions (e.g. Spjeld- from slightly older beds in Ibero-Armorica (Vannier naes 1967). Thus, the ‘Zagorian series’ is equivalent & Babin 1995; Gutierrez-Marco & Rabano 1996). In to the lower (but not lowermost) Tremadocian to a similar way, rare occurrences of late Oretanian to middle Arenigian strata; the ‘Morgatian series’ to the early Berounian brachiopods from the subsurface of upper Arenigian to lowermost Berounian, and the the Algerian Sahara (Melou et al. 1999) are closely ‘Tafilaltian series’ with the lower (but not lower- related to coeval faunas from Ibero-Armorica and most) Berounian to Kralodvorian. Also the old Alge- also with the promising Oretanian and Berounian rian substages ‘n1b’, ‘n3b’ and ‘n4’ (Legrand & Nabos faunas from the Kabylia region of northern Algeria 1962; Legrand 1985) are correlated with upper Tre- (Termier & Termier 1950, pls 1–14). Many other madocian, upper Berounian and Hirnantian strata, examples can now be added, including the trace fos- respectively. sil record. Thus, the producer of Cruziana petraea Figure 5 illustrates a correlation between various was a particular arthropod with a distinct setation Ordovician sections in the Mediterranean area using (‘claw formula’) of its appendages which produced the Bohemo-Iberian regional scale. This correlation sets of three heavy, rounded and subequal scratches, is far from complete, due to the general lack of which left its traces only in the uppermost Dobroti- updated data from most of the countries that made vian–Lower Berounian strata from southern Jordan, up South Gondwana during the Ordovician. LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 275

Fig. 5. Correlation chart, using the Bohemo-Iberian regional scale, of selected Ordovician sections from some of the better-known areas in South Gondwana. A generalized section for Bohemia (Prague Basin) is shown in Figure 2. Locality numbers are as in Fig. 6.

Thus, the extent and influx of the Mediterranean collage were part of South Gondwana as corroborated faunas seem especially difficult to recognize in the by population data on detrital zircons (Henderson present North Africa and Middle East, where most et al. 2016). However, palaeontological record is palaeontological data from surface outcrops are con- scarce and remains insufficiently investigated. siderably scattered and out of date or are not The most inshore assemblages come from eastern included in recent regional stratigraphical syntheses. Serbia (Fig. 6.14), where Lower Ordovician sand- Consequently, we include below a brief account of stones have yielded giant lingulifom brachiopods the more significant data compiled only from areas (Lingulobolus hawkei, Pseudobolus? salteri) in the that are often overlooked on global palaeogeographi- Ranovac-Vlasina unit (Gutierrez-Marco et al. 1999b cal reconstructions for the Ordovician, emphasizing and references therein), Berounian trilobites and the fossil record from the Arabian plate and some brachiopods (Dalmanitina cf. proaeva, Aegiromena Eastern European countries of former Soviet influ- cf. aquila) in Upper Ordovician shales in the Kucaj ence where data were obscure and unavailable for a Mountain (Krstic 1960, Krstic et al. 1999), and bra- long time (Fig. 6). The much more abundant and chiopod coquinas with the cystoid Heliocrinites in precise data retrieved from the Ordovician rocks of the Homolje Mountains (Veselinovic 1973). Middle Ibero-Armorica, Sardinia, the Carnic Alps, Bohemia, Ordovician shales from western Macedonia Morocco and Algeria are often analysed and dis- (Fig. 6.13) contain Neseuretus cf. tristani (see Pet- cussed in international journals and are excluded kovski & Temkova 1981). In the Sofia Balkan of NW from the following summary. Bulgaria (Fig. 6.15) there are lower and upper Ore- tanian assemblages with trilobites and graptolites Carpatho-Balkanides (lower assemblage with Placoparia, Cyclopyge cf. kossleri, Didymograptus cf. artus, D. cf. spinulosus, D. Ordovician rocks from some of the Palaeozoic intra- cf. ferrugineus,?Haddingograptus; upper assemblage Alpine blocks belonging to the SE European Galatian with Pricyclopyge b. binodosa, P. binodosa prisca, 276 Gutierrez-Marco et al. LETHAIA 50 (2017)

20º 0º 20º 40º 60º 2 ATLANTIC 1 OCEAN 4 3 12 10 40º 5 9 14 40º 13 15 6 8 16 17 7 21 23 24 11 19 28 30 20 22 18 29 26 33 54 25 27 32 MEDITERRANEAN SEA 31 55 34 45 52 56 38 35 37 36 50 39 57 42 51 53 40 46 58 41 43 60 44 59 20º 20º 49 47 48 800 km

Fig. 6. Present-day distribution of most of the Ordovician successions with Mediterranean faunas cited in the text and in Figure 5. 1, Prague Basin (Bohemia, Czech Republic); 2, Saxo-Thuringia (Germany); 3, North Armorican Domain (Normandy, France); 4, Mid- Armorican Domain (Brittany, France); 5, Cantabrian Zone (NW Spain); 6, Bucßaco area (central Portugal); 7, Almaden-Sierra Morena area (southern Central Iberian Zone, Spain); 8, Iberian Range (NE Spain); 9, Pyrenees (Spain and France); 10, Montagne Noire (SE Mas- sif Central, France); 11, Iglesiente and Sarrabus areas (S Sardinia, Italy); 12, Carnic Alps (Austria–Italy); 13, Stogova mountain (W Mace- donia); 14, Homolje and Kucaj mountains (E Serbia); 15, Sofia Balkan (NW Bulgaria); 16, Istanbul area (NW Anatolia, Turkey); 17, Zonguldak terrane (NW Anatolia, Turkey); 18, Kemer area (western Taurus, Turkey); 19, Seydisßehir (western Taurus, Turkey); 20, Ova- cik area (southern Taurus, Turkey); 21, Degirmentasß–Kozan area (eastern Taurus, Turkey); 22, Amanos Mountains (western Border Folds, Turkey); 23, Derik–Mardin area (central Border Folds, Turkey); 24, Sßort Tepe (eastern Border Folds, Turkey); 25, Abba area (N Syria); 26, Sinat area (N Iraq); 27, Amadia district (N Iraq); 28, Tatavrud, SW Bandar-e Anzali (W Alborz region, NW Iran); 29, N of Teheran (central Alborz, Iran); 30, Damghan area (eastern Alborz, Iran); 31, Bojnurd area, N Esfaraen (central Iran); 32, Anarak region (central Iran); 33, Derenjal area, N Tabas (central-east Iran); 34, Kerman region (central-east Iran); 35, Kuh-e Surmeh, Zagros Range (S Iran); 36, Faraghan mountains, Zagros Range (SE Iran); 37, east-central Sinai (NE Egypt); 38, Southern Desert (S Jordan); 39, Tabuk –Sßafajah area (NW Saudi Arabia); 40, Atß-Tınıyat (Jabal Sammar)–Al Hanadir (NW Al-Qasim) areas (central Saudi Arabia); 41, subsurface of the Central Arabian Basin (Saudi Arabia); 42, Dibba Zone (NE United Arab Emirates); 43, Sait Hatat, south of Muscat (N Oman); 44, subsurface of the Ghaba Salt Basin (N Oman); 45, Anti-Atlas (Morocco); 46, Zemmour Basin (N Mauritania); 47, Hod area (S Mauritania); 48, northern Iullemeden Basin (NE Mali); 49, Djado inlier (N Niger); 50, Ougarta Range (W Algeria); 51, subsurface Ahnet Basin and Bled el Mass inlier (central Sahara, Algeria); 52, subsurface Oued Mya Basin–Hassi Messaoud area (northern Sahara, Algeria); 53, Illizi Basin (subsurface) and Tassili n’Ajjers (Algeria); 54, Kabylia region (N Algeria); 55, subsurface Ghadamis Basin (S Tunisia); 56, Tripolitanian subsurface, Ghadamis Basin (NW Libya); 57, Gargaf High, N Murzuq Basin (Libya); 58, Tihemboka High, W Murzuq Basin (SW Libya); 59, Jebel Eghei, W Kufra Basin (SE Libya); 60, Jbel Aiba, E Kufra Basin (SE Libya). Localities mentioned in the text from Benin and Eritrea are outside the map and not represented here.

Microparia, Ectillaenus and Didymograptus murchi- Ordovician strata rest disconformably on the soni), lower Dobrotivian trilobites (Pricyclopyge Cadomian basement in the Istanbul and Zonguldak b. longicephala) and middle Berounian brachiopods terranes of NW Anatolia and are subdivided in and trilobites (Dalmanitina sp., Cyclopyge cf. various formations ranging from Tremadocian to rediviva, Aegiromena aquila). A post-glacial record Hirnantian. The palaeontological record of the Tre- of the Hirnantian graptolite M. persculptus was pro- madocian to middle Darriwilian formations consists vided by Sachanski (1993). Bulgarian data were essentially of graptolites and rare trilobites of the recently reviewed by Gutierrez-Marco et al. (2002b, raphiophorid–trinucleid biofacies (Dean et al. 2000; 2003). Gonc€ uoglu€ et al. 2014). Overlying strata comprise diverse stratigraphical units, starting in places with a Turkey Sandbian limestone containing conodonts, raphio- phorid and remopleuridean trilobites and some paly- Different terranes with Ordovician rocks have been nomorphs (Ketencikdere Formation of Dean et al. recognized in Turkey. These are the northern Anato- 2000). In a different section close to Istanbul, the lia (=Pontides s.l.) area (Fig. 6.16–17), the Tauride lower Cengelkoy€ Member of the Gozda€ g Formation is Belt (Fig. 6.18–21) and the SW Anatolian Auto- a transitional sequence from Darriwilian to Sandbian chthon or Border Folds domain (Fig. 6.22–24). strata with brachiopods, trilobites and conularids of LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 277

Mediterranean affinities (Sayar & Cocks 2013). The (Fig. 6.22–23). It comprises two shaley members sep- overlying Pendik Member has rich brachiopod assem- arated by a middle sandy member. The lower shales blages of the Nicolella fauna of the Mediterranean are very fossiliferous, bearing trilobite assemblages Province accompanied by bryozoans, ostracods and representative of the middle Berounian and associ- echinoderms. The top of the Ordovician succession is ated with other Mediterranean brachiopods (Svobo- in the lower part of the Kayalidere Member, which daina havliceki, Aegiromena), molluscs (bivalves, contains a diverse Hirnantia brachiopod fauna, some rostroconchs, cephalopods, gastropods), machaeridi- possible tabulate corals (Thamnopora – which may ans, ostracods and rare graptolites (Dean 1967b, alternatively be a bryozoan), echinoderms and grap- 1983; Dean & Monod 1985). Among the trilobites tolites (M.aff.persculptus, Normalograptus). Sayar & there are dalmanitaceans (Dalmanitina proaeva, Cocks (2013) identified and in great part illustrated Kloucekia phillipsii,?Dreyfussina), calymenaceans the assemblages of these formations and other occur- (Neseuretinus turcidus – =? N. birmanicus; Colpoco- rences of Sandbian, Katian and Hirnantian bra- ryphe cf. grandis; Iberocoryphe – ex Brongniartella – chiopod assemblages in the Go€geren Formation of levis;?Huemacaspis – ex ?Platycoryphe), trinucleids the Zonguldak terrane. There, the Hirnantia fauna (Deanaspis bedinanensis, D. orthogonius, D.? laticir- co-occurs with the trilobite Mucronaspis cf. mu- rus, Dionide formosa anatolica) and other trilobite cronata and the coral Streptelasma. groups (Nobiliasaphus cf. nobilis, Selenopeltis inermis Ordovician successions from the Taurides and Bor- angusticeps, Ampyx nitidus) in an assemblage that der Folds almost invariably begin with the Seydisßehir shows some resemblance to the trilobites from the Formation, a thick sequence of shales and quartzitic Monte Orri Formation of Sardinia (Hammann & sandstones that in many places span the Cambrian/ Leone 2007), the High Zagros of southern Iran (Gho- Ordovician boundary. Trilobites and conodonts badi Pour et al. 2015b) and the Abba Formation of occur in rare carbonate intercalations in its lower Syria. Seilacher (1970) also indicates the record of part. Apart for a single occurrence of upper Tremado- ‘Cruziana’ almadenensis in the middle sandstones at cian graptolites (Sachanski et al. 2006), most of the the Amanos Mountains. Trilobites from the upper formation is referred to the Arenigian, which may green shales (Calymenella boisseli,?Dreyfussina) indi- contain, locally, thick developments of Cruziana ich- cated younger beds, tentatively assigned to the upper nofacies. In southern and eastern Taurus Berounian. The overlying Sßort Tepe Formation com- (Fig. 6.18,20,21) the sparse trilobite faunas (Taihung- prises grey shales and silty mudstones, occasionally shania cf. miqueli, Paramegalaspis, Niobella, Orometo- with thin oolitic limestones towards the base. In sev- pus) and rare molluscs (Dean & Monod 1990; Dean eral localities across the Taurides and the Border et al. 1999) show affinities with Arenigian assem- Folds (Fig. 6.18,21,24) the unit contains a diverse blages from SE France, Sardinia, Iran, United Arab trilobite assemblage of Anglo-Baltic affinities, which Emirates and SW China. At the places where the ero- becomes widespread during the Boda Event of global sional gap at the top of the Seydisßehir Formation is warming. Characteristic forms such as Ovalocephalus reduced or absent, a local development of limestones tetrasulcatus, Nankinolithus,‘Ulugtella’ mediterranea takes place towards the top of the unit (Tekmen lime- – ex Zdicella –, Prionocheilus obtusus, Ceraurinella stone) or just above it (Sobova Formation and lower and Cyclopyge (Dean & Zhou 1988; Dean & Monod beds of the ‘Kilgen Lake Formation’). Palynomorphs 1990) are representative of coeval assemblages from the uppermost Seydisßehir Formation allow for recorded from Kralodvorian strata in Sardinia and correlation to the upper Arenigian, which includes the Iberian Range (Hammann 1992; Hammann & the Darriwilian 1 stage slice (Paris et al. 2007b). The Leone 2007) and in the Alborz Mountains of Iran Sobova Formation in its type area (Fig. 6.19) has (Karim 2009). The top of the Sßort Tepe Formation trilobite assemblages with Baltic affinities (Carolinites, was correlated with the Ancyrochitina merga chitino- Metopolichas, Phillipsinella, Panderia, Illaenus, Ampyx zoan biozone by Paris et al. (2007b), but the acri- and others; Dean 1973). Conodont data from the tarch assemblages recorded from lower beds of the limestones and palynological records from the overly- same formation indicate strong discrepancies with ing shales (Fig. 6.19,21) indicate correlation to Darri- macrofossil data. Thus, its large overlap with the wilian 2–3 (Paris et al. 2007b). Bedinan Formation suggested in different areas can Fossiliferous Upper Ordovician strata from the be interpreted in terms of the extensive recycling of Taurides and Border Folds are mostly contained in palynomorphs from previously eroded strata, in a the Bedinan, ‘Kilgen Lake’ and Sßort Tepe forma- similar way as the documented reworking of Early tions, sometimes referred to as the ‘Bedinan Group’ Ordovician acritarchs (Paris et al. 2007b). in drill core studies. The Bedinan Formation Finally, the Hirnantian ice-related sediments of crops out in the Border Fold area of SE Turkey the Halevikdere Formation yielded a single element 278 Gutierrez-Marco et al. LETHAIA 50 (2017) from the Hirnantia brachiopod fauna (Mirorthis), decades, with the publication of detailed studies on recorded in the Ovacik area (Fig. 6.20). But the Hir- trilobites, brachiopods, cephalopods, ostracods, nantian age of the unit is also coincident with its echinoderms, conodonts and palynomorphs. These abundant acritarch and chitinozoan record, indica- have allowed considerable progress on the biostratig- tive of the Tanuchitina elongata biozone (Monod raphy and correlation of some important sections in et al. 2003; Paris et al. 2007b). the Alborz Mountains (Fig. 6.28–30), in central Iran (Fig. 6.31–34), and in the Zagros Mountains Syria (Fig. 6.35,36). The Ordovician sequence in the Alborz Moun- The Abba Group in a deep borehole in north Syria tains (northern Iran) occurs mainly in the eastern (Fig. 6.25) yielded an assemblage of trilobites, bra- outcrops of the Lashkarak Formation in the Dam- chiopods, machaeridians and graptolites, tentatively ghan region. This unit is composed by more than assigned to the Llandeilo by Sudbury (1957) and lat- 200 m of silty shales and sandstones with intercala- ter correlated with the Bedinan Formation of the tions of nodular argillaceous to silty limestones and Border Folds of SE Turkey (Dean 1975, p. 369). The marls. The formation ranges from lowermost record of Oepikograptus bekkeri [=‘Diplograptus Tremadocian to upper Darriwilian and still has not spinulosus’] allows the correlation of part of the been subdivided in formal members, although the fauna to lower Berounian strata. Benthic faunas of Simeh-Kuh and Gerd-Kuh sections (eastern Alborz) South Gondwanan type are probably represented comprises numerous fossiliferous beds, many of here by Colpocoryphe grandis?[=‘C. arago’], Nobil- them reasonably well studied. Tremadocian strata iasaphus nobilis,?Aegiromena [=Sowerbyella? sp.], ? yielded diverse assemblages of trilobites, bra- Paldiskites [=Lingula cf. rugosa] and Plumulites cf. chiopods, conodonts and rare graptolites, described fraternus. From a different drill core close to the in the papers of Bruton et al. (2004), Ghobadi Pour Turkish border, Flugel€ (1963) adds the record of (2006), Ghobadi Pour et al. (2007a, 2011a,b, 2015a), some biserial graptolites, among them the Bohe- Popov et al. (2008, 2009, 2015), Evans et al. (2013), mian-related taxa (Berounian to early Kralodvorian) Kebria-ee Zadeh et al. (2015) and Jahangir et al. Archiclimacograptus cf. vulgatus (Perner) and Anti- (2016), among others. Of special interest is the early costia cf. teres (Perner), but the assemblage needs record of the orthoidean brachiopods Paralenorthis, re-examination. Ranorthis, Gondwanorthis and Tarfaya (a ‘Moroc- can–Argentinian’ genus; see Benedetto & Munoz~ Iraq 2016), which were widespread during the Arenigian in South Gondwana. Also of interest is the late Tre- Ordovician rocks are poorly known from northern madocian occurrence of Taihungshania miqueli and Iraq, being restricted to the upper part of the Khabour Asaphellus in the Paroistodus proteus conodont bio- Formation and the Sinat Shale. The former is a thick zone (Tremadocian 3 stage slice), of clear affinities sequence of sandstones and shales, broadly equivalent with localities in SE France, Sardinia, Turkey and SE to the Seydisehir Formation of Turkey, which in their Arabia. Graptolites from the same strata probably uppermost part contained some Arenigian bra- are ‘Tetragraptus’ bulmani and ?Kiaerograptus, chiopods (Palaeoglossa cf. attenuata,?Lingulepis), rare instead of Tetragraptus ex. gr. quadribrachiatus and endoceratid cephalopods and trace fossils (Cruziana Didymograptus, respectively. Higher beds in the furcifera, C. goldfussi, C. rugosa, Rusophycus carleyi, ? same formation recorded the opportunistic Thysan- Didymaulichnus). These occurrences come from the otos–Leptembolon association of lingulate bra- core of the Ora anticline (north Amadia; Fig. 6.27), chiopods in the Floian, prior to the onset of listed – and partly figured – by Dunnington et al. carbonate sedimentation. After a relatively con- (1959) and Seilacher (1963, 1970, 1993). densed sedimentation in the Dapingian, Darriwilian In the Sinat area of NW Iraq (Fig. 6.26), deep- strata yielded Neseuretus sp. – ex ‘N. aff. tristani’ –, water shales with trace fossils of the Nereites ichnofa- ostracods, brachiopods, conodonts and rhombiferan cies (Seilacher 1963) yielded a thorax of the trilobite and diploporite cystoids (Echinosphaerites, Heliocri- Selenopeltis buchi, probably representing Berounian nites, Sinocystis, Tholocystis, Glyptosphaerites) that strata. show similarities to Baltic, South Gondwanan and Asiatic faunas (Lefebvre et al. 2005; Ghobadi Pour Iran et al. 2007b, c, 2011c; Popov et al. 2008; Bassett et al. 2013; Evans et al. 2013; Bogolepova et al. 2014). Knowledge of the Ordovician faunas from Iran has Upper Ordovician strata in the Alborz Mountains experienced a significant boost in the last two are represented by a well-developed ‘Lashkarak LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 279

Formation’ sensu lato in the western areas (Karim Lower Ordovician trilobite assemblages are domi- 2009), and by the upper member of the Ghelli For- nated by asaphids and nileids, with some warm-water mation in eastern Alborz areas (Ghavidel-Syooki trilobites (Carolinites, Peltabellia, Brachyhipposiderus) et al. 2011a). A rich trilobite fauna recorded together with raphiophorids and calymenaceans from limestones in Tatavrud area (Fig. 6.28) bears (Pseudocalymene superba, Neseuretus ghavideli). Darri- clear affinities with the Kralodvorian faunas of wilian assemblages yielded early records of Asiatic South Gondwana, Avalonia, Baltica, and several immigrants and the appearance of new taxa among peri-Gondwanan Asiatic terranes. It is an outer- trilobites (Neseuretinus birmanicus, Ovalocephalus), shelf assemblage that includes Ovalocephalus, ostracods (?Aechmina cf. ventadorni, Cerninella, Dicranopeltis, Symphysops, Amphoriops, Birmanites Vogdesella, Ogmoopsis), brachiopods (Nicolella), gas- and ?Eccoptochile, among other widespread genera tropods (Lesueurilla)andechinoderms(Heliocrinites), (Karim 2009). In eastern Alborz, the shales of the among other taxa which are widely distributed in Ghelli Formation provided assemblages of acri- South Gondwana. Based on this, Ghobadi Pour & tarchs, cryptospores and chitinozoans which, mostly, Popov (2009) proposed a modified palaeogeographi- overlap the range of the Tatavrud trilobite fauna, cal peri-Gondwanan reconstruction, suggesting that but represent more inshore environments as already central Iran was part of an ‘overlap’ zone where tropi- indicated by the sedimentological data and trace cal and high-latitude benthic taxa coexisted. fossil record from its middle member (Bayet-Goll & The Kerman Region of east-central Iran Neto de Carvalho 2016). There is an important (Fig. 6.34) is another key area for reconstructing the thickness of strata (150 m) assigned to the Armorico- palaeobiogeographical links with the western chitina nigerica plus Ancyrochitina merga chitino- Mediterranean region. The Ordovician succession is zoan biozones, which are directly followed by the represented by the Katkoyeh Formation. Its lower Spinachitina oulebsiri biozone in the upper 95 m of shale member produced various benthic and plank- the Ghelli Formation (Ghavidel-Syooki et al. 2011a). tonic graptolites characteristic for upper Arenigian In central Iran, Tremadocian strata from the area strata (Rickards et al. 1994, 2001). Higher beds between Bojnurd and Esfaraen (Fig. 6.31) have been include calcareous sandstones rich in bryozoans and compared with the Lashkarak Formation sensu lato. the brachiopod genus Rostricellula, followed by Jahangir et al. (2015) recorded early Tremadocian highly fossiliferous younger Ordovician beds. graptolites (Rhabdinopora flabelliformis) towards the Among these are silty argillites containing bivalves middle part of the Cordylodus lindstromi conodont (Modiolopsis), cephalopods (Sactorthoceras) and gas- biozone, which spans the Cambrian–Ordovician tropods (Tritonophon peeli, Nonorios, Slehoferia, boundary. The single benthic taxon mentioned in Tropidodiscus), which are overlain by a brachiopod– that paper is a trilobite (Asaphellus inflatus) occur- bryozoan assemblage with Drabovia aff. crassior and ring in higher beds. In a different region of central Rhynchotrema (Bassett et al. 1999; Dastanpour et al. Iran (Anarak; Fig. 6.32), Popov et al. (2015) and 2006; Ebbestad et al. 2008). The Kerman assemblage Hairapetian et al. (2015) mentioned, within the is directly comparable with that of middle Berounian Chahgonbad Formation, a lower assemblage with strata of Bohemia, SW Europe and Morocco. brachiopods, cephalopods and ostracods (Schallreuter The Zagros Mountain Range includes other sig- et al. 2006) of Middle Ordovician age. In the upper nificant Ordovician outcrops in southern Iran. One third of the same formation, trilobites (Vietnamia cf. site, poorly known and pending revision, is located teichmuelleri), brachiopods (Rostricellula cf. ambi- in the Kuh-e Surmeh area of the Fars Province gena) and ostracods show affinities with the Upper (Fig. 6.35). In it there is an unnamed unit consisting Ordovician assemblages of Sardinia and United Arab in more than 60 m of micaceous shales with Emirates. interbedded sandstones and ending in a ferruginous Ordovician sections from the Derenjal Mountains dolomite. Its Berounian trilobites (Dalmanitina, north of Tabas (central Iran; Fig. 6.33) include the ‘Eohomalonotus’ bohemicus – ?Platycoryphe) and rare very fossiliferous Shirgest Formation, which ranges graptolites indicate affinities with faunas from the from Tremadocian to Darriwilian. Stratigraphy and Bedinan Formation of Turkey and the Abba Group faunas from this unit (trilobites, conodonts, ostra- of northern Iraq (Dean 1980, p. 17). A better-known cods, cephalopods, gastropods, echinoderms) have area lies in the Faraghan Mountains of the High been studied by Pillet (1973), Bruton et al. (2004), Zagros (Fig. 6.36), where the Seyahou Formation Lefebvre et al. (2005), Ghobadi Pour et al. (2006), starts with thin Floian strata followed by a strati- Ghobadi Pour & Turvey (2009), Ghobadi Pour & graphical gap (Ghavidel-Syooki et al. 2014). Above Popov (2009), Ghaderi et al. (2009), Evans et al. it, the formation yielded important Berounian (2013) and Ebbestad et al. (2016), among others. assemblages of brachiopods (Svobodaina havliceki) 280 Gutierrez-Marco et al. LETHAIA 50 (2017) and trilobites (Dalmanitina cf. acuta, Iberocoryphe, (Cruziana rugosa and others) as well as some yet-to- Deanaspis aff. vysocanensis, Sardoites aff. pillolai, be-described trilobites (El Fiky, unpublished obser- Neseuretinus aff. malestanus) that indicate close sim- vation cited in Kora 1991). The lower member of the ilarities with SW European and Turkish Berounian formation consists of purple and olive-grey siltstones faunas (Ghobadi Pour et al. 2015b). The age of these and shales, probably coeval with Middle Ordovician assemblages is well constrained by concurrent chiti- strata in south Jordan and Saudi Arabia. nozoans, belonging to the Acanthochitina barbata and Armoricochitina nigerica biozones of the South Jordan Gondwana regional scale (Ghobadi Pour et al. 2015b). Additional occurrences of cephalopods, Fossiliferous Ordovician rocks from Jordan are echinoderms, tentaculitids and machaeridians were restricted to the Southern Desert region (Fig. 6.38), reported in the same beds by Ghavidel-Syooki et al. where the succession displays some similarities with (2015). The Seyahou Formation is capped by an ero- that described from the NW Saudi Arabia. Because sive surface, being followed by the glaciomarine of the predominance of massive sandstones over deposits of the Dargaz Formation, coeval with the extensive epicratonic areas of NE Africa and the Ara- Hirnantian glaciation (Ghavidel-Syooki et al. bian Peninsula, the Ordovician formations of Jordan 2011b). According to the latter authors, these sedi- were originally adscribed to the so-called Nubian ments may indicate the presence of satellite ice caps Sandstone Facies, and were later subdivided with the adjacent the Zagros margin of Arabia. The post-gla- help of trace fossils (Seilacher 1993) and sedimento- cial sedimentation is represented by the Sarchahan logical studies. Formation, mainly Silurian in age, although its basal The oldest Ordovician units from Jordan are the beds include Hirnantian graptolites from the Cruziana sandstones presented in the Disi and Umm Metabolograptus persculptus biozone (Ghavidel- Sahm formations in the upper part of the Ram Syooki et al. 2011b; Ghobadi Pour et al. 2015b). Group, where sparse occurrences of Cruziana furcifera, C. goldfussi and other trace fossils have Afghanistan been identified (Bender & Huckriede 1963; Seilacher 1970; Selley 1970). The overlying Hiswah Formation Owing to its Ordovician sedimentary and palaeonto- (originally referred to as ‘graptolite sandstone’), logical record, the Tezak-Nawar domain of central which constitutes the base of the Khreim Group, Afghanistan was considered by some authors as comprises a lower shaley and an upper sandy mem- directly correlatable either with the Central Iberian ber. Both are fossiliferous, having yielded graptolites, Zone of Spain (Blaise & Bouyx 1980) or with SE lingulacean brachiopods, rare crinoid remains and Turkey (Dean 1980). The succession started with abundant trace fossils, the latter restricted to the Cruziana-sandstone facies, with Tremadocian beds sandstone beds. The graptolite ‘Didymograptus cf. at the base bearing Cruziana semiplicata, followed by bifidus’ has been repeatedly cited in the regional lit- Arenigian sandstones with bivalves (Afghanodesma erature since Bender & Huckriede (1963: pl. 3, fig. desparmeti) and trace fossils (Cruziana furcifera, 2), but their figured specimens, as well as new C. goldfussi, Didymaulichnus lyelli). Overlying shales unpublished material, is here reassigned by yielded Darriwilian graptolites (Didymograptus Gutierrez-Marco to D.(D.) murchisoni, a widespread murchisoni) and higher up an Upper Ordovician upper Oretanian species. The elkaniid brachiopod ? trilobite assemblage very similar to the one recorded Broeggeria and the ichnogenera Skolithos, Cruziana from the Bedinan Formation of Turkey. It comprises and Arthrophycus (=‘Harlania’) have also been iden- Neseuretinus birmanicus and N. malestanus tified (Amireh et al. 2001; Turner et al. 2012). (although both could be preservational variants of Above the shelf deposits of the Hiswah Formation, N. turcicus, see Hammann & Leone 1997), Deanaspis the Dubaydib Formation represents much shallower cf. bedinanensis, Nobiliasaphus, lichids and cyclopy- environments, and corresponds to the ‘Sabellarifex gids (Gutierrez-Marco 1997, and references therein), sandstone’ of earlier authors. Among the ichnofossils but these data remains largely overlooked in more Skolithos predominates in the lower part, whereas recent literature. the upper member yielded a diverse assemblage of Cruziana, Arthrophycus, Rusophycus, Diplichnites,? Egypt Rossellia, Skolithos and rare finds of the bivalve Modiolopsis (Bender & Huckriede 1963; Wolfart Ordovician successions from Egypt lie essentially in et al. 1968; Selley 1970; Seilacher 2000). Seilacher east-central Sinai (Fig. 6.37), where the thin Abu (1970) reviewed in detail the ichnoassemblage from Hamata Formation provided some trace fossils the uppermost beds of the formation at the Sahl el LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 281

Karim locality, west of Wadi Rum, and defined the which contains reworked fossils exhumed from older new ichnotaxa ‘Cruziana petraea’(=Rusophycus beds, as well as some trace fossils. Its age is indicated petraeus), C. lineata, C. flammosa and C. perucca, by the record of Metabolograptus persculptus in the together with the ‘Iberian’ form ‘C.’ almadenensis. overlying post-glacial Batra Formation, which is con- Owing to the stratigraphical position and the particu- tinuous with the Silurian black shales (Loydell 2007). lar morphology deduced for the trace markers (unu- sual trilobites and other arthropod groups), Seilacher Saudi Arabia (1970) suggested an early ‘Caradoc’ age for these beds. This was partly confirmed by later records of Ordovician fossils from Saudi Arabia have been ‘C.’ almadenensis and ‘C. petraea’inSpain,which recovered from surface outcrops and boreholes in come from uppermost Dobrotivian to lowermost the Tabuk region (Fig. 6.39) and the Jabal Sammar– Berounian quartzites (Lorenzo & Gutierrez-Marco Jabal Al Hanadir areas (Fig. 6.40), as well as from 2008; Gamez Vintaned & Carls 2011). the subsurface of the central Arabian basin The Turbeiliyat Formation overlies concordantly (Fig. 6.41). Palaeontological research on Ordovician the previous beds and represents more fossiliferous rocks from SW Saudi Arabia (Dibsiyah and Anamah marine facies. It is equated here, not only to the formations of the Wajid Group) has not yet pro- ‘Conularia sandstone’, but also to the ‘Nautiloid sand- duced significant results. stone’ of earlier authors. The latter includes thin, cal- Besides some Tremadocian palynomorphs usually careous sandstone horizons that were erroneously reworked in younger strata, the older palaeontologi- placed in the Silurian – above the Batra Shale – at a cal occurrences in NW and central Saudi Arabia time when the unawareness of the presence of thick come from the Cruziana-bearing sandstones that glacial-related sediments led to the misunderstanding constitute the upper part (Sajir Member) of the Saq of the stratigraphical relationships with underlying – Formation. These strata where renamed as the Anz or apparently coeval – beds. The first fossils from the Formation by Khalifa (1993, 2015), who restricted Turbeiliyat Formation were discovered by Bender & the Saq Formation to the Cambrian. Ordovician ich- Huckriede (1963) and studied by Wolfart et al. noassemblages from the upper Saq (or Anz) Forma- (1968), but the localities could benefit from a review. tion are composed of Cruziana rugosa, C. furcifera, The original material comes from several beds and C. goldfussi, C.? imbricata, Didymaulichnus lyelli, fossiliferous sites, comprising rare rhynchonelliform Lockeia and Diplichnites (El-Khayal & Romano brachiopods (Rostricellula arabica, Dalmanella aff. 1988). In the Al Hanadir area, the top 1.5 metres of horderleyensis, Platystrophia), linguliforms (Orbicu- the formation is intercalated by a conglomeratic bed lothyris costellata, Trematis tenuiornata, Lingulella), (20–50 cm thick) with lingulacean debris and fish trilobites (Brongniartella benderi, B.aff.platynota), microremains, which in some boreholes provided bivalves (Modiolopsis sp, Ctenodonta, Goniophorina, abundant microfossils of probable terrestrial origin. pterioids), bellerophontoidean gastropods (‘Bucanella There are cryptospores (monads, obligate dyads and aff. bohemica’: probably Tritonophon peeli), cephalo- tetrads), cuticlelike sheets, clusters of sporelike cells, pods, crinoids and rare conularids (‘Exoconularia sp.’: and hilate sporomorphs (Strother et al. 1996; Le probably Archaeoconularia), plus what could be either Herisse et al. 2007) attributable to primitive land tabulate corals or massive bryozoans. Bender & Huck- plants. The sandstones overlying this bed contain riede (1963) and Wolfart et al. (1968) also mentioned trilobites (Neseuretus), some graptolites (Didymo- trace fossils, one of them identified as Cruziana graptus s.str. and unidentified biserials) and the trace pudica by Seilacher (1970). Interestingly, Orbicu- fossils Phycodes fusiforme [=‘P. aff. palmatum’], ? lothyris costellata was later found in pre-glacial and Gordia and Lockeia (El-Khayal & Romano 1988; Sei- glacial-related sediments in the Murzuq Basin of lacher 2000). The graptolites illustrated by El-Khayal Libya, and also from central Saudi Arabia. Although & Romano (1988, fig. 6f) indicate an Oretanian age the entire assemblage was originally regarded as ‘Car- for the topmost beds of the formation in the Al- adoc to Ashgill’ in Jordan, the absence of index taxa Qasim region. However, the same strata in a drill prevented a detailed correlation. The only significant core to the west of the Tabuk –Sßafajah area yielded information is that the Jordanian species O. costellata an older microfossil assemblage that indicates a ranges in Saudi Arabia below the Quwarah Member probable diachronic development for the end of (Kralodvorian) of the Quasim Formation and that a sand sedimentation. This was suggested after the probable late Berounian–?early Kralodvorian age is recognition of the Velatachitina veligera chitinozoan also suggested for their Libyan occurrence. biozone (Al-Hajri 1995), a local equivalent to the The latest Ordovician strata in Jordan are repre- late Arenigian (ca. Dapingian) Desmochitina ornensis sented by the glacial-related Ammar Formation, biozone of Ibero-Armorica. 282 Gutierrez-Marco et al. LETHAIA 50 (2017)

The Cambro-Ordovician Saq Formation – or the Lingulella); some bivalves such as Hemiprionodonta restricted Ordovician Anz Formation – is overlain lusitanica? =‘Glyptarca cf. naranjoana’; and palaeotax- by the Tabuk Group (Middle Ordovician to Lower odonts, cephalopods, bellerophontiform gastropods, Devonian), which starts with the Qasim Formation crinoids, ostracods (a.o. ‘Bollia’), conodonts (Balto- and includes the remaining Ordovician strata. The niodus variabilis, Drepanodus homocurvatus, D. sub- latter begins with the Hanadir Shale, a very fossilifer- erectus, Oistodus abundans) and fish debris, cited in ous lower member which seems broadly correlatable Thomas (1977), Khashogi (1979), El-Khayal (1985), with the lower part of the Hiswah Formation of Jor- El-Khayal & Romano (1985, 1988), Williams et al. dan, the Am5 (Upper Siltstone) Member of the (1986), Vaslet et al. (1987), Henry & Destombes Amdeh Formation of Oman, and the Ayim Member (1991) and Rickards & Khashogji (2000). Probably of the Rann Formation of the United Arab Emirates. the actinocerid and orthocerid cephalopods ?Mesakto- The fossil record of the lower part of the Hanadir ceras and ?Sactoceras, identified by Evans (2000) in a Member consists of abundant graptolites such as sample coming from the Takuk area, were also Didymograptus (D.) murchisoni and rare biserial derived from the sandy storm layers at the upper part fragments, the problematic form Hanadirella bran- of the Hanadir Shale. The age of this unit has been kampi, indeterminate ostracods, bivalves and lingu- debated, but the occurrence of Pterograptus 0.7 m laceans, plus the trilobites Neseuretus cf. tristani, from the base and of Didymograptus only 1 m below Ningkianolithus hanadirensis and an indeterminable its top, clearly indicates that the Hanadir Member can asaphid (Khashogi 1979; Fortey & Morris 1982; El- be correlated with upper Oretanian strata. The rich Khayal 1985; El-Khayal & Romano 1985, 1988). This palynological record of the Hanadir Shale (acritarchs, assemblage occurs 1.5–2.0 m above the base of the transported cryptospores, chlorophycean algae, Hanadir member and is indicative of the lower- chitinozoans) ranges from the upper part of the upper Oretanian (Fig. 4). The outcropping basal- S. formosa to the lower part of the L. pissotensis chiti- most beds only contain N. cf. tristani and Pterograp- nozoan biozones, being the latter represented in the tus cf. elegans (= ?Pterograptus in El-Khayal & middle and upper third of this member (a.o. Al-Hajri Romano 1988 p. 167, judging by a picture by El- 1995; Paris et al. 2000; Le Herisse et al. 2007). Khayal, Gutierrez-Marco pers. Observation), but The overlying Kahfah Member of the Qasim For- core samples from an equivalent position on the mation is formed in their lower part by fine-grained Saqqar structure recorded a mixed assemblage of sandstones with interspersed sparsely fossiliferous chitinozoans that was adscribed by Al-Hajri (1995) beds, containing a few bivalves, trilobites (Kerfor- to the upper Arenigian Desmochitina bulla biozone, nella), ?fish debris (Williams et al. 1986) and biserial despite containing reworked specimens of the index graptolites in need of a thorough taxonomic revi- taxa from the younger Siphonochitina formosa and sion. A puzzling assemblage of chitinozoans, proba- Linochitina pissotensis biozones (lower to basalmost bly belonging to the Lagenonochitina ponceti upper Oretanian strata). As indicated by this author, biozone, was reported by Al-Hajri (1995) and proba- it is possible that part of this mixed assemblage was bly represents uppermost Dobrotivian strata, reworked from the Saq Formation, as seen in the roughly equivalent to the basal Sandbian. Higher subsurface of the Sßafajah area. The oldest non- beds in the succession only provided long-ranging reworked chitinozoan assemblage recorded from the chitinozoan species, maybe equivalent to most of the Hanadir Shale is from the Cyathochitina calix?to lower and middle Berounian, ranging from the Siphonochitina formosa biozones (Al-Hajri 1995), the L. deunffi to the J. tanvillensis biozones (Al-Hajri latter co-occurring with D.(D.) murchisoni. 1995). The middle part of the Hanadir Member contains The next shaley unit is the Ra’an Member of the a depauperated assemblage with abundant D.(D.) Qasim Formation that at its base yielded chitino- murchisoni and Hanadirella, lingulaceans and cepha- zoans of the Tanuchitina fistulosa biozone (Paris lopod remains. However, in the upper 5–17 m the et al. 2000), near another poorly diversified unit becomes richly fossiliferous, coinciding with epi- assemblage bearing Mediterranean-type trilobites sodes of distal storm events. These strata yielded, at (Kerfornella and ?Deanaspis =‘Onnia’), lingulacean two localities, a shelly fauna of Mediterranean affini- braquiopods, biserial graptolites (Normalograptus cf. ties, most significant among the fossils the trilobites brevis,‘Diplograptus’), cephalopod remains and con- Neseuretus tristani, Iberocoryphe verneuili, Kerfornella odonts (Williams et al. 1986). In the Tabuk region, sp., Plaesiacomia vacouvertis, P. aff. rara, Crozonaspis Janjou et al. (1997) reported from the Ra’an Mem- aff. morenensis and Kloucekia cf. drevermanni.There ber some brachiopods (Drabovinella, Fezzanoglossa are also brachiopods – both rhynchonelliform (?Tiss- fezzanica, Orbiculothyris costellata, Lingulella), as well intia)andlinguliform(Monobolina, Schizocrania,? as unidentifiable trilobites, bryozoans, conularids LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 283 and ‘annelids’ (sic: either machaeridians or cornuli- also of mid- and late-Arenigian age (Floian 2– tids?). This assemblage includes lingulacean bra- Dapingian 1 stage slices). From the trace fossil- chiopod taxa previously known only from Libya and rich sandstone intercalations Cruziana furcifera, Jordania. Near the top of the Ra’an Shale, McClure C. rugosa, Rusophycus, Planolites, Skolithos and (1988) cited the occurrence of a biserial graptolite, Arenicolites were identified. erroneously identified as Glyptograptus persculptus The distinctive Ayim Member of the Rann Forma- (sic). tion includes in their upper part thin condensed The youngest member of the Quasim Formation nautiloid limestone beds with large burrow systems is the Quwarah Member, consisting of alternations of Thalassinoides type at some of their bases. Apart of sandstones and shales. Near the base, a distinct of different brachiopod taxa and crinoid ossicles, bed with fossiliferous nodules has been reported by dasycladacean algae, gastropods, bivalves, fragments Janjou et al. (1997), bearing linguliform bra- of illaenid and odontopleurid trilobites, ostracods chiopods (lingulaceans and discinaceans such as O. and foraminifera (Fortey et al. 2011; and references costellata), bivalves, cephalopods, bryozoans, colo- therein), the bioclastic limestones provided con- nial cornulitids and rare gastropods. A trilobite odont assemblages that were reviewed as indicative assemblage, also from the basal part, included Caly- of a probable upper Pygodus serra biozone (Darriwil- menella boisseli, Calymenella sp. and Iberocoryphe ‘al- ian 3 stage slice). The lower beds of the Ayim Mem- mualleni’ – a nomen nudum (Janjou et al. 1997, p. ber contained rare trilobites (Nileus emiratus of 13). Widespread occurrences of chitinozoans in the Hamdan & Mustafa 2009). Quwarah Member (Al-Hajri 1995; Paris et al. 2000) Finally, the upper member of the Raan Formation are broadly indicative of the Armoricochitina nigerica includes some shell beds and fossil remains washed- and Ancyrochitina merga biozones and of the in as storm deposits. Transported assemblages of Kralodvorian stage. Higher up, this member is trun- trilobites with a definite Gondwana signature com- cated everywhere by the glacial erosive surface at the prise characteristic middle Berounian trinucleids base of the Sarah Formation (Hirnantian), which (Deanaspis goldfussi seftenbergi, originally described mainly contains inherited fossils from earlier forma- from Bohemia and related with the Sardic form tions as well as some pre-Silurian chitinozoans D. goldfussi fluminensis), calymenaceans (Vietnamia recorded in Central Arabia (Paris et al. 2000, teichmuelleri, described in Sardinia) and dalmani- ‘Assemblage 1’). taceans (Dreyfussina cf. taouzensis, a widespread form known from Morocco and SW Europe). In United Arab Emirates addition to the trilobites, the index-species of the Tanuchitina fistulosa chitinozoan biozone was prob- Fossiliferous Ordovician rocks crop out in the Dibba ably recognized from this member (Rickards et al. Zone of the northern Oman Mountains, east of 2010). Dubai (Fig. 6.42). They constitute ‘exotic’ rafts of Rann Formation within the Kub Melange, thrusted Oman and deformed during the emplacement of the Semail Ophiolite of the Haybi Complex. The Rann Forma- Outcropping exposures south of Muscat (Fig. 6.43) tion is a relatively thin siliciclastic unit (ca. 100 m) are represented by the Amdeh Formation, which which is subdivided into three members by an inter- corresponds to shallow-water shelf deposits. The val of grey-red shales and griotte-like nodular bio- Upper Quartzite Member (Am4 + lower Am5 carto- clastic limestones (Ayim Member). Palaeontological graphical units) has yielded abundant upper Areni- data comes essentially from the complementary gian trace fossils (Cruziana rugosa, C. furcifera, work of Rickards et al. (2010) and Fortey et al. Rusophycus, Planolites, Teichichnus, Phycodes, Daeda- (2011). The lower member of the Rann Formation lus, Skolithos linearis) as well as fish microremains comprises green shales with alternation of quartzitic (Sacabambaspis), preserved in fine-grained crinoidal sandstones very rich in trace fossils (Cruziana ichno- calcarenites (Lowelock et al. 1981; Sansom et al. facies). The shales bear rare graptolites (Baltograptus 2009). It is possible that the vertical traces illustrated deflexus) and acritarchs, as well as the Mediter- by Seilacher (2000) as Daedalus labechei and ranean-type trilobites Neseuretus cf. arenosus and D. desglandi, and recorded from an unprecise Cruzi- Taihungshania cf. miqueli, closely resembling a mid- ana-rich ‘Arenig quartzite’ located ‘north [sic] of dle Arenigian assemblage from the Montagne Noire. Muscat’, come from the same area and outcrops. Besides these, weathered limestone nodules repre- Above these sandstones, the Upper Siltstone senting a former hyolithid shell bed provided some (=Am5) Member of the Amdeh Formation pro- conodonts belonging to an outer-shelf assemblage, duced South Gondwana trilobites such as Neseuretus 284 Gutierrez-Marco et al. LETHAIA 50 (2017) tristani and Ogyginus, bivalves (Redonia) and crinoid Mali remains (Iocrinus cf. subcrassus and diverse colum- nals). They were partly illustrated by Sansom et al. The only known Ordovician fossils from Mali come (2009) and Donovan et al. (2011), and the assem- from the NW Iullemeden Basin (NE Mali; Fig. 6.48), blage probably represents upper Oretanian strata, where Gatinskiy et al. (1966) reported a single sand- based on the lowest occurrence of N. tristani in stone succession, 12–17 m thick, bearing abundant Ibero-Armorica. brachiopods, graptolites and trilobites. The grapto- Other Ordovician fossils from Oman are known lites are all benthic forms collected from the lower from the subsurface of the Ghaba Salt Basin beds, followed by sandstones with abundant bra- (Fig. 6.44). Two different boreholes (Ghaba-1 and chiopods, and scarce trilobites restricted to the Baqlah-1) have yielded graptolites and paly- upper part of the succession. nomorphs from two closely set horizons located in The brachiopod assemblage is composed by typi- the lower half of the Saih Nihayda Formation of the cal Berounian taxa dominated by ‘Svobodaina ex gr. Haima Supergroup. The former contained Didymo- inclyta’ sp. (=Svobodaina sp.; the Bohemian species graptus (D.) artus, scolecodonts and lingulacean S. inclyta was later synonymized with S. ellipsoides; remains (lower Oretanian), while the latter had see Colmenar et al. 2014). Gatinskiy et al. (1966) Didymograptus (D.) cf. murchisoni (?upper Oreta- also cited Mesodalmanella ex. gr. flava, Heterorthina nian beds). In the absence of index-species of the notata, Rostricellula ex gr. ambigena and ‘Dalmanella Gondwanan chitinozoan zonation, Rickards et al. aff. bouceki’. The naming form of the latter is a Hir- (2010) tentatively attributed the associated chitino- nantian species presently reassigned to the genus zoans to different levels of the Siphonochitina for- Horderleyella, which makes its occurrence in the mosa biozone which, however, is still restricted to association very improbable. Finally, the trilobite lower Oretanian strata in Ibero-Armorica (Fig. 4). material belongs to the genera Dalmanitina and Palynological assemblages from the Ordovician of Calymenella, also common in Berounian strata Oman are quite interesting because two of the chiti- throughout the Mediterranean Province. nozoan species (Desmochitina mortoni and Belone- chitina ghabaensis) also show remote occurrences in Niger low-latitude areas of North Gondwana, indicating an open marine connexion with SW Arabia and that To the west of the Murzuq–Djado Basin, the Palaeo- the distribution of some chitinozoans is not exclu- zoic outcrops that constitute the Tihemboka High of sively latitudinally controlled (Molyneux et al. 2004, Libya and SE Algeria they extend in a SSE direction 2006; Al-Ghammari et al. 2010). reaching the Chirfa region of north Niger (Fig. 6.49). There, the homonymous Chirfa Forma- Mauritania tion contained, near its base, the Late Ordovician graptolite Metabolograptus ojsuensis (Legrand 1993). In the Zemmour area of northern Mauritania Despite this species usually being restricted to lower (Fig. 6.46), Tremadocian shales unconformably over- Hirnantian strata, Legrand (2011) argued in favour lie Archaean rocks of the Reguibat Shield, being the of an older record based in its association with a SW equivalent of the Fezouata Shale of the Moroccan piece of trinucleid trilobite, a group rare in the Hir- Anti-Atlas. From these strata, Destombes et al. (1969) nantian (Owen 2007). Paris (1990) characterized described a similar assemblage of trilobites and grap- these shales as probably falling in the A. merga and tolites of the Adelograptus and Araneograptus murrayi T. elongata chitinozoan biozones. biozones, the former group being represented by the genera Nileus, Platypeltoides, Megistaspis (Ekeraspis), ? Benin and Eritrea Anacheirurus, Bavarilla and Prionocheilus. Further south, the latest Ordovician rocks from the Towards the southern margin of the very wide area Hodh area (Fig. 6.47) provided a post-glacial record occupied by the Hirnantian ice cap, Upper Ordovi- of the Hirnantian graptolite Metabolograptus persculp- cian strata were also represented in the glacial- tus and some elements of the ‘pre-Akidograptus related Kandi Basin which extended from SW Niger ascensus’ fauna (Underwood et al. 1998). From the to Benin, as well as in south-central Eritrea and same sections, Paris et al. (1998) also identified Late northernmost Ethiopia, in the Horn of Africa. Ordovician chitinozoans (Tanuchitina elongata) Upper Ordovician trace fossils from northern followed by assemblages which showed an atypical Benin occur in the Ka Member of the Kandi Forma- mixture of the last Ordovician and first Silurian forms tion, interpreted as post-glacial marine deposits (Spinachitina fragilis). (Konate et al. 2003). From it ‘Cruziana petraea’, LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 285

C. cf. furcifera and Arthrophycus brongniartii mounds developed above these beds, preserve a [=A. ‘linearis protrusiva’] were identified (Seilacher diverse assemblage of bryozoans (Buttler et al. 2007) & Alidou 1988; Alidou et al. 1991; Seilacher 2000). dominated by trepostomes (Jifarahpora libyensis, Ordovician trace fossils from Eritrea include Orbipora, Dekaya, Hallopora and Monotrypa), with Arthrophycus alleghaniensis, A.?brongniartii, minor representation of Cryptostomata, Fenestrata, Didymaulichnus lyelli, Palaeophycus tubularis and Cystosporata and Cyclostomata. Conodonts from Taenidium isp. The assemblage was recorded by the Amorphognathus ordovicicus biozone were also Kumpulainen et al. (2006) from the Adigrat Forma- found (Bergstrom€ & Massa 1992), as well as some tion, which post-dates the glacial beds of the Edaga ostracods (Satiellina jamairensis), brachiopods Arbi Formation of southern Eritrea and northern (Orbiculothyris costellata, strophomenoids), stro- Ethiopia and also yielded some terrestrial matoporoids (Labechia), porifera (Protospongia) and sporomorphs. fragments of trilobites and echinoderms (Massa 1988). A Kralodvorian age is fully justified for most Libya and Tunisia of the Jifarah Formation, but according to Paris (1990) it also comprises some Hirnantian strata Ordovician rocks occur in the subsurface of the (Tanuchitina elongata chitinozoan biozone). northern Ghadamis Basin, which extends from east- Another important area with Ordovician rocks in ern Algeria to Tunisia and NW Libya, including the Libya and adjacent countries is the Murzuq Basin, Djeffara, Southern Chotts, Talemzane–Bou Nemcha which includes superb exposures of Ordovician and Berkine small basins. Numerous drill cores strata in its northern (Gargaf High) and western yielded palaeontological data from southern Tunisia (Tihemboka High) boundaries (Fig. 6.57,58), as well (Fig. 6.55) and Tripolitania (Fig. 6.56), the oldest of as in the extension of the latter in eastern Algeria them coming from Tremadocian sandstones and and northern Niger. The fossiliferous succession shales of the Sanrhar Formation. Massa et al. (1977) starts with the Hawaz (=Achebyat) Formation, a recorded phyllocarid remains and lingulacean bra- sandstone-dominated sequence very rich in trace chiopods such as Tunisiglossa tripolitanea, Palaeo- fossils, with abundant Skolithos linearis and scattered glossa crumena and Siphonotreta. After a sedimentary occurrences of Daedalus multiplex, Bergaueria, gap corresponding to the Late Tremadocian through Cochlichnus, Cruziana, Rusophycus, Arthrophycus early-middle Arenigian (Vecoli et al. 2004), the brongniartii (ex A. ‘linearis’), Teichichnus rectus and Kasbah-Leguine and the Bin Ben Tartar formations Aulichnites (Seilacher 2000; Ramos et al. 2006). A contain a rather complete record of chitinozoans ‘Llandeilo’ age is currently suggested for the forma- ranging from the Cyathochitina calix to the Linochi- tion, which only provided at its base some acritarchs tina pissotensis biozones (Paris 1990). close to the Middle and Late Ordovician boundary Diverse upper Oretanian assemblages of typical (Vecoli & Le Herisse 2004). The overlying Melaz South Gondwana platform type also comprise trilo- Shuqran (=Melez Chograne) Formation yielded bites (Neseuretus tristani, Crozonaspis morenensis cf. upper Berounian faunas, mostly derived from older mayensis, Plaesiacomia cf. vacouvertis, Ogyginus corn- beds and occurring as reworked elements embedded densis) and linguliform brachiopods (Schmidtites, in glaciomarine diamictites. They comprise trilobites Trematis), which were recovered from core samples (Calymenella cf. boisseli, Flexicalymene, Salteroco- in Tunisia and Tripolitania (Massa et al. 1977; ryphe,?Prionocheilus,?Kloucekia, Mucronaspis, Cocks & Fortey 1988; Massa 1988). Dreyfussina cf. struvei and D. libyca), brachiopods A new sedimentary gap occurred in the Dobroti- (Drabovinella, Drabodiscina cf. grandis, Aegiromena vian and the entire Berounian on the Ghadamis descendens, Rafinesquina cf. pseudoloricata,‘Rhyn- Basin, ending in the deposition of the fossiliferous chotrema’ cf. clariondi, Fezzanoglossa fezzanica, Jifarah (Djeffara) Formation of the subsurface of Libyaeglossa colombi, Orbiculothyris costellata, and Tripolitania. This unit comprises a lower siliciclastic Acrosaccus – ex ‘Orbiculoidea’ – massai) and rare member rich in lingulacean brachiopods (Palaeo- echinoderms (Codiacystis bohemica), bryozoans glossa cf. attenuatta, Orbiculothyris costellata, (Polyteichus and ceramoporoids), hyolithids and ‘Torynelasma’ hyolithiformis, Lingulella) and rare conularids. Part of these fossils were cited or rhynchonelliforms (Onnizetina,‘Moridorhynchus’), described, among others, by Havlıcek & Massa together with some bivalves (Synek), gastropods (1973), Cocks & Fortey (1988), Massa (1988) and (‘Bucanella’) and ostracods (Antiaechmina libyensis), Becq-Giraudon & Massa (1996). According to mentioned or described by Massa et al. (1977), Havlıcek & Massa (1973) and Havlıcek (1990), the Massa (1988) and Schallreuter & Hinz-Schallreuter uppermost Ordovician Mamuniyat (=Memouniat) (1998). Dolomitic limestones and bryozoan mud- Formation provided some brachiopods of the 286 Gutierrez-Marco et al. LETHAIA 50 (2017)

Hirnantia fauna preserved in sandstones (Hirnantia & Servais 2005; Kraft et al. 2015). However, this is cf. sagittifera, Plectotyrella lybica). only applicable to the global standard scales under Finally, the Kufra Basin of SW Libya includes the rules of the ICS-IUGS, not to regional or local some Ordovician outcrops of the Hawaz Formation scales. bearing trace fossils (Cruziana gr. rugosa, Researchers who disagree with the convenience of C. rouaulti, Arthrophycus brongniartii, A. simplex, a Bohemo-Iberian regional scale for South Gond- Skolithos, Cylindrichnus) in the areas of Jabal Ehei wana often substantiate their direct comparison with and Jabal Asba (Fig. 6.59–60), cited by Seilacher the global scale through an interpretative dating of (1970), Turner & Benton (1983), Seilacher (2000) regional biozonations based on graptolites and paly- and Seilacher et al. (2002). The occurrences of nomorphs. This is the case of the ‘North’ Gondwana ‘Cruziana petraea’ and C. rugosa respectively chitinozoan zonation (Paris 1990), which is very reported from northern Chad (Seilacher 1970) and useful for correlating Ordovician strata from high eastern Sudan (Seilacher, 2000), both within the palaeolatitudes, but highly speculative regarding the Kufra Basin, can be adscribed to the same unit. precise dating and correlation with the global scale. Paris (1990) compiled data from the east of the As indicated by Gutierrez-Marco et al. (2014), Sirt Basin on the record of the Upper Ordovician Gutierrez-Marco et al. (2016) the use of these bio- E. tanvillensis, A. nigerica, A. merga and T. elongata zones as the main biochronological criteria for South chitinozoan biozones from the subsurface of Gondwana created problems due to its changing Cyrenaica Platform (NE Libya). correlation with either the global or the regional Avalonian scale (Gutierrez-Marco et al. 2014, fig. 1). The FAD of some index chitinozoan taxa may be Conclusions also diachronous around Gondwana. This is the case of the key ‘lower-middle Tremadocian’ Lageno- The Global Ordovician chronostratigraphy, estab- chitina destombesi and ‘Floian’ Euconochitina lished worldwide in deep-water successions and symmetrica biozones, which are mostly restricted to defined by the earliest records of a few graptolite and upper Tremadocian strata in South China (Wang conodont species, is difficult to apply in the South et al. 2013), as well as of the ‘upper Floian’ Eremo- Gondwana palaeobiogeographical region. The gener- chitina brevis, which is massively recorded in the alized absence of the global index taxa and their suit- lower Floian beds in another Chinese section (Liang able facies motivates the replacement of the global et al. 2014). Correlation problems continue with the scale by a regional scheme based on the abundant Linochitina pissotensis Biozone, then and now almost palaeontological record of shallow-water taxa found automatically correlated with ‘Llandeilian’ strata, in in the high-palaeolatitudinal settings around the spite of L. pissotensis having its FAD in upper Oreta- Ordovician South Pole. Sporadic occurrences of nian beds over a wide area including Iberia, the graptolites and shelly faunas of ‘Baltic’ or ‘Avalo- Algerian Sahara and Saudi Arabia. And most nian’ affinities allow for indirect correlation with the recently, a new suggestion for the top of this biozone global scheme through the regional scales for Bal- extends up into lower Sandbian strata, based on a toscandia and British Avalonia, respectively. sequential analysis of the Morgat Member (Pos- However, the Bohemo-Iberian scheme is still not tolonnec Formation) of western France (Dabard widely accepted by followers of the historical British et al. 2015). However, the associated macrofauna regional scale wether or not they incorporated its from these tentative upper beds of the L. pissotensis conceptual evolution in their work. Some even biozone lies at the famous ‘villa Les Atlantes’ fossil considered the South Gondwana regional scale locality that still falls within the uppermost Lower unnecessary ‘for not incorporating substantial Dobrotivian (Morgatia hupei trilobite subzone – see improvements neither in dates nor in correlation Fig. 4). Problems with the chitinozoan scale con- precision’ (Paris & Servais 2005, p. 136). The same tinue with the younger Ancyrochitina merga biozone, group of researchers promoted instead the direct which is so far recorded from Kralodvorian (upper- adoption of the terminology from the global scale, most Katian) strata, but it seems to extend into the disregarding the absence of the great majority of the Avalonian Hirnantian (Challands et al. 2014; fig. graptolite and conodont species that define the base 11). At a Moroccan scale, it is worth mentioning that of the global series, stages and ‘time-slices’. Their the Armoricochitina nigerica biozone was recognized critical view of the use of alternative chronostrati- in the stratotype of the Agadir-Tissint (=Amouguer) graphical scales is motivated by the fact that regional member of the Lower Ktaoua Formation (Elaouad- stages are mainly defined based on regional bio- Debbaj 1986; locality 1580) and in equivalent strata zones, instead of the FADs of certain species (Paris in the complete Upper Ordovician section of Bou LETHAIA 50 (2017) Bohemo-Iberian regional chronostratigraphical scale 287

Ingarf (Paris 1990, p. 201). Regardless, Loi et al. widespread chitinozoan biozones (e.g. the bulla (2010, fig. 3) placed the base of this biozone, defined transgressive event and the formosa flooding event of within the range of A. nigerica, at a higher strati- Paris et al. 2007a). However, uncertainties concern- graphical position within the Upper Tiouririne For- ing the precise dating of some of these cycles still mation at Bou Ingarf. persist, and the biostratigraphical calibration and The use of the regional Bohemo-Iberian scale palaeontological research on other fossil groups resolves these discrepancies and promotes a multi- must continue before this rather theoretical sequen- disciplinary and intergraded view of South Gond- tial stratigraphy can be considered as a standard of wanan palaeontology and palaeogeography, thus correlation for South Gondwana. providing for a higher resolution to interregional The Bohemo-Iberian regional scale must be correlations. Authors that use the global scale in a encouraged as it is a more stable and precise frame tentative manner, in the absence of valuable correla- on which to base sequential stratigraphy and inter- tion criteria, subconsciously introduce inadequate national correlations of the extensive south polar correlations inherited from the use of the British area of Gondwana. This use does not exclude in any scale. One of the most common mistakes observed way the convergence with the evermore precise glo- is, for instance, the frequent confusion of the Darri- bal standard scale, and new studies will help better wilian (often written as ‘Darriwillian’) as a virtual define the direct correlation with other regional synonym of the former Llanvirn+Llandeilo series, schemes. when this global stage also includes the upper Areni- Acknowledgements. – This work has been supported by the gian. There is also the incorrect assumption of the IBEROR project (‘Iberian Ordovician chronostratigraphy and its Sandbian and Katian as equivalent to the traditional correlation with the global scale’, ref. CGL2012-39471/BTE) of Caradoc, which resulted in an imprecise boundary the Spanish Ministry of Economy and Competitiveness. The pre- sent study is dedicated to the memory of Vladimir Havlıcek, with the former Ashgill and in a random assignment Ladislav Marek, Nils Spjeldnaes and Wolfgang Hammann, pio- of the Ka1 to Ka3 stage slices. Similarly, the direct neers in the modern chronostratigraphical development of the dating of well-known fossil localities and horizons in regional scale and for their detailed research of the ‘Mediter- ranean’ brachiopod and trilobite faunas. We are grateful to Prof. terms of global ages generate unfortunate correla- Stan C. Finney (California State University, Long Beach) and tions, such as confusing the ‘upper Darriwilian’ with Prof. Stig M. Bergstrom€ (Ohio State University, Columbus) for the former British Llandeilo or Llandeilian, instead their detailed review and constructive criticism. of using more precise regional terms (e.g. lower Dobrotivian, upper-lower Dobrotivian, lower-upper Dobrotivian – compare biostratigraphical equiva- References lences in Fig. 4). Moreover, ‘upper Darriwilian’ also Al-Ghammari, M., Booth, G.A. & Paris, F. 2010: New chitino- includes age equivalents of the Didymograptus zoan species from the Saih Nihayda Formation, Middle murchisoni graptolite Zone, because the earliest Ordovician of the Sultanate of Oman. Review of Palaeobotany Pygodus serra and Palynology 158, 250–261. occurrence of the conodont (base of Al-Hajri, S. 1995: Biostratigraphy of the Ordovician chitinozoa Dw 3 stage slice) is currently placed within the of northwestern Saudi Arabia. Review of Palaeobotany and D. murchisoni Zone (Webby et al. 2004). Examples Palynology 89,27–48. 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