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Geological Society, London, Memoirs

Chapter 10 Biogeography of Ordovician linguliform and craniiform brachiopods

Leonid E. Popov, Lars E. Holmer, Michael G. Bassett, Mansoureh Ghobadi Pour and Ian G. Percival

Geological Society, London, Memoirs 2013, v.38; p117-126. doi: 10.1144/M38.10

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Chapter 10

Biogeography of Ordovician linguliform and craniiform brachiopods

LEONID E. POPOV1, LARS E. HOLMER2*, MICHAEL G. BASSETT1, MANSOUREH GHOBADI POUR3 & IAN G. PERCIVAL4 1Department of Geology, National Museum of Wales, Cardiff CF10 3NP, UK 2Department of Earth Sciences, Palaeobiology, Uppsala University, 752 36 Uppsala, Sweden 3Department of Geology, Faculty of Sciences, Golestan University, Gorgan, Iran 4Geological Survey of New South Wales, Londonderry, 2753 NSW, Australia *Corresponding author (e-mail: [email protected])

Abstract: The biogeographical patterns shown by Ordovician linguliform and craniiform brachiopods are greatly influenced by their dominance in low-diversity associations in marginal environments. This is particularly evident in the Early Ordovician, when linguliform-dominated dysaerobic assemblages are widely distributed along the deep shelves of Gondwana, the Kazakhstanian terranes and in Baltica. By the Darriwilian, micromorphic linguliforms are characteristic components of the pantropical climatic-controlled faunas of Laurentia, Cuyania and Kazakhstanian terranes, which – in spite of separation by extensive oceans – retain a distinct similarity. Analysis of craniiform biogeographical distribution is impeded significantly by the poor state of craniide taxonomy and lack of reliable data from most regions. However, in general their biogeographical dispersion is similar to other groups of the Palaeozoic Evolutionary Fauna. Unlike the linguliforms, which are important members of the Cambrian Evolutionary Fauna, there is no convincing Cambrian craniiform record; they may have evolved and dispersed from Gondwana and associated microcontinents and island arcs. The earliest well-established record is from the late Tremadocian of temperate to high-latitude peri-Gondwana. During most of the Ordovician, they have a peri-Iapetus distribution. They are very rare or absent in tropical Gondwana, South China and Kazakhstanian terranes and are not yet documented from Siberia. The trimerellides probably evolved in tropical peri-Gondwanan island arc settings. Their dis- persion and major features of biogeography mirror those of atrypides.

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Linguliform and craniiform brachiopods comprise a relatively Ordovician, as discussed below. Recent linguliforms have a plank- minor, but distinctive component of Ordovician benthic faunas. totrophic larva with a prolonged free-swimming stage in their Linguliforms are recognized as one of the major components of ontogeny, which was retained from ancestral Cambrian stocks the Cambrian Evolutionary Fauna, together with trilobites and (Freeman & Lundelius 1999). By the beginning of the Ordovician hexactinellide sponges (Sepkoski 1981), whereas the existing they became adapted to a wide spectrum of marine environments Cambrian record of craniiforms is sparse (Popov et al. 1999a), from near-shore to abyssal depths (Bassett et al. 1999; Tolmacheva and in terms of three evolutionary faunas recognized by Sepkoski et al. 2004). In the Ordovician, linguliform brachiopods played a (1981) they can be considered as a minor component of the significant role in faunal assemblages that were characteristic of Palaeozoic Evolutionary Fauna (Harper et al. 2004). The biogeo- marginal environments, for example in near-shore mobile sands graphy of the Ordovician linguliforms and craniiforms has not and dysaerobic conditions, which explains the special biogeogra- previously been the subject of a separate comprehensive analysis, phical patterns shown by the group. The Ordovician also is charac- but has generally been considered together with biogeographical terized by distinctive benthic associations of micromorphic studies of rhynchonelliform brachiopods. Ordovician linguli- linguliform brachiopods and there is growing evidence (Mergl form and craniiform brachiopods proliferated in marginal 2002; Holmer et al. 2005) that they commonly had symbiotic marine environments where they formed low-diversity associ- relationships with sponges, a life habit which clearly affected the ations often dominated by a single taxon. Some opportunistic pattern of their biogeography. In addition, a wide range of different linguliform-dominated faunal associations, which are associated linguliform biogeographical patterns are evident in cases where with intervals with significant environmental changes, in tandem they form minor components of various benthic communities with trimerellide associations, reveal characteristic distributional that are dominated by other filter feeders, for example rhynchonel- patterns that have proved helpful in palaeogeographical recon- liform brachiopods and bryozoans. structions (Popov & Holmer 1994, 1995). Craniiforms, in particu- The quality of published global data on the biodiversity of Ordo- lar the trimerellides, have also proven biogeographically important vician linguliforms is variable. It is relatively good for Baltica and (Popov et al. 1997). However, because of the low diversity of high- to temperate-latitude Gondwana, and there are also good these associations, the conventional methods (such as cluster and data compiled for Laurentia, the Australasian sector of low lati- principal component analyses) have a limited application here. tude Gondwana, Avalonia and the Kazakhstanian terranes. By con- trast, there is only a single publication on the Mid Ordovician micromorphic linguliforms from South China, and no available Linguliform brachiopods data from Siberia and North China. Moreover, a considerable amount of data on the Ordovician linguliforms unfortunately The life strategies and evolutionary history of linguliforms differ remains unpublished, including descriptions of Late Ordovician significantly from those of other brachiopod subphyla (Bassett microbrachiopods from Ireland (Avalonia), presented in an unpub- et al. 1999). These differences are clearly reflected in their dis- lished PhD thesis by McClean (Wright & McClean 1991), and of tinctive patterns of biogeographical distribution during the the Mid Ordovician faunas of the Argentinean pre-Cordillera.

From:Harper,D.A.T.&Servais, T. (eds) 2013. Early Palaeozoic Biogeography and Palaeogeography. Geological Society, London, Memoirs, 38, 117–126. http://dx.doi.org/10.1144/M38.10 # The Authors 2013. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

118 L. E. POPOV ET AL.

Early Ordovician (Tremadocian–Floian) changes. In Baltoscandia and the Alborz peri-Gondwanan terrane, this expansion occurred just prior to the onset of temperate-latitude The end of the Furongian (Late Cambrian) Epoch saw a major carbonate sedimentation, which coincided in both regions with decline in the generic diversity of linguliform brachiopods. The proliferation of benthic associations with clear characteristics of decline is best seen in outer shelf and basinal faunal associations the Palaeozoic Evolutionary Fauna (Bassett et al. 2002; Popov (Bassett et al. 1999), which in the early Tremadocian Age were et al. 2008). In Bohemia, the Thysanotus–Leptembolon fauna occupied mainly by the low- to medium-diversity Broeggeria appeared because of immigration after a period of non-deposition Association. This association was distributed widely along the corresponding to the entire Furongian Epoch (Mergl 1986). Bed- deep shelves of Gondwana, the Kazakhstanian terranes and in narczyk (1999) suggested that the Thysanotus Leptembolon fauna Baltica (Popov & Holmer 1994, 1995). The Broeggeria Associ- may have been of a peri-Gondwanan origin, and this is now sup- ation often tracks the distribution of dysaerobic environments, ported by the earliest known occurrence of Thysanotus from the which are characterized, in particular, by the olenid trilobite bio- Shirgesht Formation of Derenjal Mountains in Central Iran, facies. The Tremadocian micromorphic brachiopod associations where it is associated with the billingsellide brachiopod Protambo- mostly have a low diversity and they are dominated by Eurytreta nites and the conodont Cordylodus angulatus (Bassett et al. 1999). and Ottenbyella, which have almost cosmopolitan distribution. The Furongian to early Tremadocian interval in Baltica and Similar Tremadocian micromorphic brachiopod associations temperate-latitude Gondwana was a time of proliferation of have been documented from Baltica (Popov & Holmer 1994, spinose siphonotretides (Family Siphonotretidae), which show a 1995), Laurentia (Popov et al. 2002; Holmer et al. 2005) and distinctive geographical distributional pattern (Popov et al. Kazakhstanian terranes (Popov & Holmer 1994; Holmer et al. 2009a; Fig. 10.1). The earliest spinose siphonotretides are confined 2001), and from various peri-Gondwanan regions (Popov et al. to Gondwana (Bohemia, Alborz, Central Iran). In Baltoscandia, 2008; Streng et al. 2011). the siphonotretides are represented by the enigmatic genus Hel- In the transitional period from the Furongian to the early Trema- mersenia, which is characterized by having a coarsely punctuate docian, a number of low-diversity obolid associations can be found shell without spines (Williams et al. 2004) and forms almost on the shallow clastic shelves of Baltica and high- to temperate- monotaxic associations. The equally enigmatic siphonotretide latitude Gondwana, including the Obolus Association in Baltica Schizambon lacks both spines and perforations; it is found on (Popov & Holmer 1994, 1995), and the Tunisiglossa Association in both sides of Iapetus, but is more abundant in Laurentia (Holmer the North African sector of Gondwana (Havlı´cˇek 1989; Fig. 10.1). et al. 2005; Popov et al. 2009a). By the end of the Tremadocian, The late Tremadocian to early Floian transition (Paltodus delti- spinose siphonotretides replaced Helmersenia in Baltica and also fer to Prioniodus elegans biozones) is associated with an increased spread to Laurentia (Holmer et al. 2005). turnover and a significant proliferation of lingulate brachiopod In Laurentia, Tremadocian micromorphic brachiopod faunas faunas (Bassett et al. 1999). In the shallow temperate-latitude were of low diversity, and dominated by genera of the Family clastic shelves of Gondwana and Baltica, the early Tremadocian Acrotretidae (e.g. Conotreta, Eurytreta and Ottenbyella; Popov obolid associations were replaced mainly by the Hyperobolus et al. 2002; Holmer et al. 2005). Diverse micromorphic brachiopod Association (Bohemia, Australasian sector of Gondwana and the associations with early representatives of the acrotretide fami- South Urals; Havlı´cˇek 1982, 1989; Popov & Holmer 1994, 1995; lies Eoconulidae (Otariconulus), Ephippelasmatidae (Akmolina, Percival & Engelbretsen 2007) and later by the Thysanotus– Mamatia, Pomeraniotreta), Scaphelasmatidae (Eoscaphelasma) Leptembolon Association (Fig. 10.1), which then spread widely and Torynelasmatidae (Sasyksoria) have been described from in Bohemia, the East Baltic, South Urals, Alborz and Central late Tremadocian–early Floian deposits of Kazakhstanian ter- Iran (Mergl 1986, 1996, 1997; Popov & Holmer 1994, 1995; ranes, which were located in a subequatorial peri-Gondwana Ghobadi Pour 2008; Popov et al. 2008; Fig. 10.1). In addition to setting (Popov & Holmer 1994). By contrast, contemporaneous epibenthic obolids, which form the core of the association, there micromorphic linguliform brachiopod associations of the Austra- are also some infaunal taxa (e.g. Leptembolon and Lingulella) lasian sector of Gondwana documented by Brock & Holmer together with epibenthic spinose siphonotretides (e.g. Eosiphono- (2004) and Percival & Engelbretsen (2007) are of relatively treta and Siphonobolus). low biodiversity. In addition to widespread genera, for example The expansion of the Thysanotus–Leptembolon Association Otariconulus, Ottenbyella, Semitreta and Wahwahlingula, they was diachronous and associated with significant environmental contain Libecoviella otherwise known only from Bohemia

Fig. 10.1. Geographical distribution of selected linguliform brachiopod taxa and associations in the Tremadocian. Relative position of the main palaeocontinents is mainly after Cocks & Torsvik (2002). Sibumasu and Central Iran are considered parts of mainland Gondwana (Torsvik & Cocks 2009); Alai and Alborz are isolated microplates located in temperate latitudes between Gondwana and South China (Popov et al. 2009a). Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

ORDOVICIAN LINGULIFORM AND CRANIIFORM BRACHIOPODS 119

(Mergl 2002), whereas an unidentified ephippelasmatid described The major features of temperate-latitude lingulate brachiopod and illustrated by Percival & Engelbretsen (2007) is most probably faunas can be typified by those described from the Mid Ordovician congeneric with Ghavidelia otherwise known from the Alborz of Baltica (Gorjansky 1969; Holmer 1989), Iran (Popov et al. peri-Gondwanan terrane (Popov et al. 2008). 2008) and South China (Zhang 1995). They are characterized by In Baltoscandia, the late Tremadocian to early Floian micro- a low taxonomic diversity and abundance of ephippelasmatids morphic brachiopod associations also have diverse Acrotretidae, belonging to the genera Aipyotreta, Myotreta and Numericoma. including Eoconulidae (Otariconulus), Ephippelasmatidae (Akmo- Other characteristic genera are Scaphelasma (Scaphelasmatidae), lina, Mamatia, Pomeraniotreta) and the earliest Biernatiidae Torynelasma (Torynelasmatidae), Biernatia and Eoconulus (Biernatia), but lack representatives of the Scaphelasmatidae (Eoconulidae). Most of these genera, with the exception of Aipyo- and the Torynelasmatidae (Popov & Holmer 1994; Holmer & treta, Myotreta and Sasyksoria, are cosmopolitan. Torynelasma- Biernat 2002). During the late Tremadocian–Floian, micromor- tids are present in the most of known faunas of that age, but phic brachiopod associations of Baltica show strong links to absent in Avalonia, Bohemia and Alborz, which were located at those of adjacent parts of Gondwana, as indicated by the geo- temperate latitudes, and they are also absent from the Cuyania graphical distribution of such genera as the dysoristid Ferrobolus, Terrane. The torynelasmatids are the most diverse in the Kazakh- the zhanatellid Rowellella, the obolid Elliptoglossa, the paterulid stanian terranes, where they are represented by the endemic Diencobolus and the acrotretides Acrotreta, Akmolina, Eoconulus, genera, such as Christicoma, Issedonia and Polylasma. The ephip- Otariconulus and Ombergia (Holmer & Popov 1994; Popov & pelasmatid Lurgiticoma is known from Kazakhstan and New Holmer 1994, 1995; Holmer et al. 2000; Mergl 2002; Ghobadi Zealand (Nazarov & Popov 1980; Percival et al. 2009) and was Pour et al. 2011). probably tropical peri-Gondwanan. A remarkable feature of the Chinese fauna is the lack of the lingulids Elliptoglossa and Pater- ula, an almost complete absence of the family Acrotretidae Mid Ordovician (Dapingian–Darriwilian) (e.g. Acrotreta, Conotreta, Cyrtonotreta and Spondylotreta), which are rather characteristic of the contemporaneous faunas in There is little available information on Dapingian linguliform Baltica, Laurentia and Kazakhstanian terranes, and the abundance brachiopods, which mostly have been described from Baltica of Myotreta, which is known otherwise only from Baltica (Gorjansky 1969; Biernat 1973) and temperate-latitude peri- (Fig. 10.2). The co-occurrence of Myotreta indicates that the Gondwana (Bohemia, Alborz, Avalonia; Mergl 1996, 2002; Chinese fauna has distinct links to the contemporaneous Baltic Popov et al. 2008). By contrast, Dapingian linguliform faunas fauna, which is also shown by the joint occurrence of common from low latitudes, including Laurentia, Siberia, North China species like Numericoma perplexa, N. spinosa, Acanthambonia and equatorial Gondwana, remain almost unknown. By the Dapin- cf. delicata and Scaphelasma mica. Another characteristic Baltic gian, micromorphic brachiopod associations of Baltoscandia genus is Biernatia, and it is remarkable that biernatids are incorporated the Scaphelasmatidae and the Torynelasmatidae. restricted in geographical distribution to Baltica during the Early Lingulides, including Paterula and Rowellella and the acrotretides Ordovician, whereas in the Dapingian and most of the Darriwilian Acrotreta, Biernatia and Numericoma, are common in Baltica and they occur only in Baltica and Alborz (Popov et al. 2008; temperate-latitude Gondwana (Gorjansky 1969; Holmer 1989; Fig. 10.2). Only from the late Darriwilian–early Sandbian interval Mergl 2002; Popov et al. 2008). (Pygodus serra – Pygodus anserinus biozones) did the Biernatidae The Middle Ordovician interval from the Lenodus variabilis become almost cosmopolitan in distribution, and by that time the to Pygodus anserinus biozones is the best known interval in family had spread to Laurentia, Cuyania, New Zealand (Australa- terms of the biogeographical distribution of lingulate microbra- sian sector of Gondwana) and Kazakhstanian terranes, as well as chiopod assemblages. They are represented on the eastern to South China. The Biernatidae is also represented by two (Alabama) and western (Nevada) margins of Laurentia (Cooper endemic genera in Avalonia (Sutton et al. 2000). 1956; Krause & Rowell 1975), in several Kazakhstanian terranes The late Darriwilian was a time of significant changes in taxo- (Popov 2000a), Iran (Popov et al. 2008), New Zealand (Percival nomic composition of microbrachiopod assemblages, with the et al. 2009) and South China (Zhang 1995), and they are widely number of cosmopolitan taxa increased, whereas the biogeogra- spread in Baltoscandia (Gorjansky 1969; Holmer 1989). In phical differences between individual faunas decreased. In South addition, significant unpublished data are available for the China the generic diversity of micromorphic lingulates nearly Cuyania Terrane in the Argentinean Precordillera. doubled in the Pygodus anserinus Biozone and such genera as

30° 30° North China

0°

0° North Annamia Laurentia Tien Shan Tarim 30° Siberia Chu-Ili South China Central Sibumasu Iran

Avalonia Turan 60° KazakhstanianAlborz 60° Baltica Terranes Turkish 30° Gondwana Urals Tauri des Armorica Hellenic Urals Perunica s urkish T Turkish Pontides PerunicaPontide Baltica a Gondwana Hellenic Turkish Taurides Turan Armoric

Central Avalonia Iran Fig. 10.2. Geographical distribution of

Undiferina Ephippelasma Biernatia (Dapingian) selected genera of micromorphic Spinilingula Myotreta Biernatia (late Darriwilian) linguliform brachiopods in the Darriwilian. Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

120 L. E. POPOV ET AL. ) . a

South China m n m o ) a Darriwilian i i t . sh Alborz lab ue Fm. ue m m rma s nus o i am u a (A c h e Valley L a i Z ilian lik F lik z o F ci i ser i a a r a e n lop enti c one one -Il d on Treh on i ri r chi akan L akan u lt n e d-Darriwilian asu- i z un a . su ioz ar valon inde At Laurentia (Utah Laurentia Baltica A Ante Kur E Cuyania Terrane Cuyania L Fm Bestamak L. variabilis L. Fm Uzunbulak Ching Dapingian Dar riw P. as Chu-Ili Terrane Cuyania B Biozone Biozone Biozone Zealand New K Ch Lau K Baltic B m Po Uhaku Pratt Ferry Fm. A Llanvirn Sharka Fm. Sharka Boh emia

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Fig. 10.3. Results of the pair-group cluster analysis for presence-absence data (Euclidean similarity index) for the Middle Ordovician linguliform Fig. 10.4. Palaeogeographical reconstruction for the Mid Ordovician (after microbrachiopod genera from 19 localities (for details see the Appendix). Ghobadi Pour & Popov 2009, modified) showing geographical distribution of the Early Ordovician craniides and Pseudocrania. Position of polar and subtropical convergences and major patterns of the oceanic circulation for Ephippelasma, Biernatia, Undiferina, Acanthambonia and Nush- the winter in the Southern Hemisphere mainly after Wilde (1991). biella make their first appearance, but representatives of the Family Acrotretidae remain very rare. The results of the Cluster Analysis (Euclidean distance, using Undiferina (the latter genus makes its first appearance in South the PAST statistical package distributed by Hammer et al. 2001) China only in the Pygodus anserinus Biozone, and in Baltoscandia of 17 Darriwilian, one early Sandbian and one Dapingian lingulate it is known only in the Sandbian). microbrachiopod assemblages from peri-Gondwana (Cuyania, In general, the observed patterns of biogeographical distri- Alborz and Bohemia), Baltica, Laurentia, Kazakhstanian terranes, bution of micromorphic lingulate brachiopods through the Mid New Zealand and South China show the significance of climatic Ordovician Epoch suggest that this was controlled mostly by control of the distribution of taxa incorporated into micromorphic climatic factors. Thus, in spite of a considerable oceanic space linguliform brachiopod assemblages. Two major clusters have separating Laurentia, Cuyania, the Kazakhstanian terranes and been recognized (Fig. 10.3). One includes faunas from South the Australasian sector of Gondwana, they retain a distinct simi- China, Alborz, Bohemia and Avalonia. In the Darriwilian, all of larity across the low latitudes. these regions were situated in high or temperate latitudes in rela- tive proximity, or within a Gondwanan margin affected by the Late Ordovician (Sandbian to Hirnantian) influence of the cool Southern Subpolar Current (Fig. 10.4). The second cluster is confined mainly to Laurentia, the tropical Linguliforms from the Late Ordovician Epoch are the least well Kazakhstanian terranes and to Baltica, which was situated in studied faunas, and there are only a few published studies from temperate latitudes. Unlike the Early Ordovician Epoch, by the Baltica (Holmer 1986, 1989, 1991a; Popov et al. 1994), Kazakh- Darriwilian Baltica had moved on a considerable distance from stanian terranes (Nazarov & Popov 1980; Popov 2000a, b), Avalo- Gondwana (Torsvik & Rehnstro¨m 2003) and supported essentially nia (Wright 1963) and the Australian sector of low-latitude endemic faunas of rhynchonelliform brachiopods and trilobites Gondwana (Percival 1978; Percival et al. 1999). There is there- (Fortey & Cocks 2003). By that time it had drifted away from fore no good basis for a detailed analysis of the biogeographical the Southern Subpolar Current, which controlled faunal migra- distribution of linguliforms within the Late Ordovician Epoch. tions along the western Gondwanan coast (Wilde 1991; Ghobadi In Baltica, micromorphic linguliforms are comparatively well Pour et al. 2010). This location may also suggest that there was known, and by the Sandbian–early Katian the associations a relatively free exchange of micromorphic linguliform brachio- include cosmopolitan lingulides such as Paterula and Rowellella, pod faunas across the Iapetus between Baltica and Laurentia and the equally cosmopolitan acrotretides Hisingerella, Biernatia, during the Darriwilian. Opsiconidion and Acrotretella, together with the siphonotretide Remarkably, Baltica forms a subcluster with the Darriwilian Acanthambonia (Holmer 1986; Popov et al. 1994). There is also faunas from the Cuyania Terrane (yet to be formally described), a moderate abundance of poorly known large lingulides, including which was located in proximity to the South American sector of several species of the infaunal Pseudolingula in Baltica plus Gondwana on the opposite side of the ocean in approximately several poorly known macroscopic linguliforms and siphonotretids the same southern latitudes (Fig. 10.3). (Gorjansky 1969; Holmer 1991a, b). The low-latitude micromorphic linguliform brachiopod Linguliforms described from the Australian sector of low lati- faunas are characterized in general by higher taxonomic diversity tude Gondwana include both endemic taxa, such as Casquella and the occurrence of pantropical genera like Rhysotreta and and Anomaloglossa, and the cosmopolitan Paterula, Elliptoglossa Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

ORDOVICIAN LINGULIFORM AND CRANIIFORM BRACHIOPODS 121 and Hisingerella (Percival 1978; Percival et al. 1999). Mid Katian of the Chu–Ili peri-Gondwanan terrane (Nazarov & Popov 1980; microbrachiopod assemblages from the lower Malongulli For- Popov et al. 1999a), and from the upper Darriwilian deposits of mation of New South Wales (Percival et al. unpublished data) the Alborz terrane (Ghobadi Pour 2008; Bassett et al. 2013) contain characteristic pantropical acrotretide genera, for exam- (Fig. 10.5). ple, Torynelasma, Undiferina, Rhysotreta and Nushbiella, in addi- The oldest known Baltoscandian craniide is Pseudocrania, first tion to the numerous cosmopolitan genera, which constitute a documented from the late Dapingian (mid Volkhovian) and background assemblage. ranging up to the top of the early Darriwilian (Kundian) (Gor- Kazakhstanian terranes, which mainly occupied subequatorial jansky 1969). From the Aserian (late Darriwilian), Pseudocrania positions west of the Australian sector of Gondwana (Popov was replaced in the Baltic Ordovician Fauna by Orthisocrania. et al. 2009b), also supported diverse micromorphic linguliform There is no record of craniides in the Early to Mid Ordovician of faunas (including Acrotreta, Biernatia, Dictyonites, Ephippe- Laurentia and Siberia (Popov et al. 1999a). lasma, Rowellella and Scaphelasma). There are a few endemic Kazakhstanian peri-Gondwanan island arcs preserved the ear- taxa, for example, acrotretoids Naimania and Tasbulakia, liest known record of trimerellides. In the Chingiz terrane they whereas Veliseptum is also known from the Katian of Baltica occur in the lower Sandbian Bestamak Formation (Nikitin & (Popov 2000a, b; Holmer 1986). Popov 1984). This assemblage includes Ussunia (which appears There is little data from late Katian and Hirnantian linguliform to retain some craniopsid-like plesiomorphic characters), together brachiopods; however, there is little doubt that micromorphic with such genera as Palaeotrimerella and Ovidiella. lingulate brachiopod assemblages were affected severely by the first wave of the terminal Ordovician extinction events at the end Late Ordovician of the supernus Biozone (Harper et al. 2004). The craniides went through a significant diversification during the Late Ordovician Epoch, and they became a distinctive minor Craniiform brachiopods component of benthic assemblages of shallow and mid-shelf set- tings in temperate zones and low latitudes. By that time they had A lecithotrophic craniiform larva with its short planktonic stage spread widely to Laurentia, Baltica, Avalonia and high-latitude had evolved in craniiforms by the Mid Ordovician Epoch (Popov peri-Gondwana, but they are as yet unknown from Siberia, North et al. 2010, 2012). This suggests that craniforms are a potentially and South China, and the Australian sector of Gondwana (Popov valuable group in palaeogeographical reconstructions. However, et al. 1999a). Remarkably, Late Ordovician craniides are extre- this analysis is impeded significantly by the issue related to the cur- mely rare in Kazakhstanian peri-Gondwanan terranes, where just rent interpretation of craniide taxonomy. The generic affiliation of a single Late Ordovician species (Orthisocrania shidertensis a number of Ordovician species referred currently to Petrocrania Gorjansky 1972) has been described from the Katian (Tauken For- and Philhedra is uncertain, and both genera are a ‘waste basket’ mation) of northeastern Central Kazakhstan. A number of Ordovi- for numerous poorly defined craniide taxa of uncertain affinity. cian craniides (e.g. Acanthocrania and Petrocrania) are known in Laurentia, Avalonia and in Baltica, on both sides of the Iapetus Early to Mid Ordovician Ocean. Orthisocrania was endemic to Baltica in the late Darriwi- lian, but spread into Avalonia by the beginning of the Katian. Phil- The oldest known craniides are Petrocrania? sp. from the upper hedra is probably the only Late Ordovician craniide genus that is Tremadocian deposits of Bavaria (Sdzuy et al. 2001) and endemic to Baltica. Its type species, P. baltica (Koken 1889), Petrocrania? caputium Mergl 2002 from the Milina Formation has a unique morphology, including hollow spines – a character- (Tremadocian) of Bohemia. As yet undescribed craniides are also istic otherwise unknown in other craniides. reported from Lower Ordovician olistoliths in the Silurian Pulgon The earliest known Ordovician craniopsides are from the Formation of southern Kyrgyzstan (Harper et al. 2004). The Sandbian deposits of Baltoscandia (Pseudopholidops) and the Mid Ordovician record of peri-Gondwanan craniides is sparse. Avalon microcontinent (Paracraniops), which occupied a similar It includes Celidocrania from the Dapingian to lower Darriwilian geographical position. During the Katian, craniopsides (mostly [Arenig] deposits of North China (Liu et al. 1985) and Pseudocra- Paracraniops) became widely distributed in low latitudes. In par- nia from the uppermost Darriwilian to lower Sandbian deposits ticular, they are known from Baltoscandia (Popov & Pushkin

Fig. 10.5. Geographical distribution of selected trimerellide taxa in the Katian. Relative position of the main palaeocontinents is mainly after Cocks & Torsvik (2002) with Eodinobolus Eodinobolus Belubula Eastwardly directed trimerellide migrations modifications for peri-Gondwanan Palaeotrimerella Palaeotrimerella Corystops during the Katian Ussunia Monomerella Costitrimerella terranes after Popov et al. (2009b). Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

122 L. E. POPOV ET AL.

1986), Laurentia (including North America (Cooper 1956) and Silurian. Almost all local trimerellide lineages outside of South South Scotland (Williams 1962)), Kazakhstan (Gorjansky 1972) China and Australia became extinct just before the Hirnantian, and Taimyr (Nikiforova 1982). Pseudopholidops was confined and there is no documented record of trimerellides in the mostly to Baltica, but it has been found recently in the mid Hirnantian and Rhuddanian brachiopod faunas. Katian deposits of the Zagros Mountains (M. Ghavidel Syooki, pers. comm. 2011) where it is associated with brachiopod and trilobite faunas characteristic of high-latitude Gondwana (e.g. Tafi- Discussion latia, Svobodaina, Neseuretinus, Dalmanitina). During the Hirnan- tian, Pseudopholidops was a relatively rare, but widely distributed The Ordovician biogeography of linguliform brachiopods is component of the Hirnantia brachiopod assemblage, and is basically controlled by the radiation and dispersion of those reported from Baltoscandia, Wales, Poland and South China lineages, which survived the severe crisis of that the group experi- (Popov et al. 1999a). enced during the late Furongian–early Tremadocian interval. The The distribution of trimerellides during the Sandbian–Katian early Tremadocian biogeography of linguliform brachiopods, with was consistent with their dispersion from equatorial peri- low-diversity, near-shore obolid communities inhabiting shal- Gondwanan island arc settings. By that time they are known from low clastic shelves of high-latitude Gondwana and Baltica and the Australian sector of Gondwana (Percival 1995) and they morphologically archaic taxa such as siphonotretides that are made their first appearance in Laurentia in the early Katian characteristic of Laurentia, Baltica and Gondwana, has a clear (Cooper 1956; Norford & Steele 1969). In South China, an abun- Cambrian origin. On the shallow shelves of temperate- and high- dant trimerellide fauna is known from Katian deposits (Li & latitude Gondwana and Baltica, lingulide communities were domi- Han 1980; Rong & Li 1993; Xu & Li 2002). However, the trimer- nated by epibenthic obolids (e.g. Obolus, Schmidtites, Ungula) ellide fossil record of this area is incomplete. Large numbers that were replaced rapidly by mixed associations of burrowing of endemics and the diachronous appearance of some genera and epibenthic lingulides (e.g. Thysanotus–Leptembolon Associ- (e.g. Eodinobolus, Monomerella and Palaeotrimerella) suggest ation) and finally by low-diversity associations of burrowing a significant geographical isolation of all these faunas (Fig. lingulides (e.g. the Tunisiglossa Association). The distributional 10.5) defined by adaptation to the restricted near-shore environ- pattern of the Thysanotus–Leptembolon Association, which ments. However, all of these faunas also exhibit some simi- emerged rapidly in Baltica and the Alborz terrane just prior to larities, which can allow establishment of the time and possible the shift from clastic to temperate-latitude carbonate sedimen- direction of faunal exchange (Popov et al. 1997). For example, tation, suggests that it was an opportunistic fauna that flourished early species of Eodinobolus were reported from the Sandbian at a time of significant environmental changes (Popov & Holmer to lower Katian deposits of Kazakhstan and Australia, while 1994, 1995; Mergl 1997; Popov et al. 2008). During the late Tre- closely related Palaeotrimerella is known from the late Darri- madocian–early Floian interval, the archaic siphonotretides, such wilian to Katian of Kazakhstan (Popov et al. 1997) and Katian as Helmersenia and Schizambon, were replaced on the shelves of of South China (Li & Han 1980). Monomerella made its first all major continents by spinose siphonotretides, which evolved appearance in the early to middle Katian of Australia (Percival originally in temperate-latitude Gondwana (Popov et al. 2009a). 1995), whereas in Kazakhstan it is recorded from the mid Another remarkable feature of linguliform biogeography is Katian (Abak Regional Stage) and appeared in South China the origin and dispersion of the associations of borrowing lingu- approximately contemporaneously in the mid Katian (Percival lides during the Early to Middle Ordovician. The earliest associ- 1995; Popov et al. 1997). ations of borrowing lingulides (e.g. Tunisiglossa Association) The observed pattern of geographical distribution of Ordovician are documented from high-latitude Gondwana (Havlı´cˇek 1989). trimerellides can be explained as the result of two major mig- This group emerged in subequatorial latitudes only from the rations from Kazakhstanian terranes: (1) during the late Sand- Darriwilian onwards. There is no clear Mid Ordovician record of bian–early Katian interval to Gondwana and North America; (2) the ‘Lingula’-type associations (after Ziegler et al. 1968) in during the mid to late Katian to Baltica and Siberia. The westerly Laurentia and Baltica. However, they are relatively well documen- direction of the migration of the Eodinobolus kazakhstanicus ted from Siberia (Kanygin & Yadrenkina 1977) and the Kazakhsta- group of species in the early Katian (Popov et al. 1997; Fig. nian terranes (Nikitina et al. 2006; Popov et al. 2007). Lingulides 10.5) suggests that there were some significant changes in are able to survive through the stress of strong changes of palaeogeography during the late Late Ordovician, because it dupli- salinity variations resulting from freshwater input (Emig 1986; cates the somewhat later dispersion of atrypoids (Popov et al. Hammen & Lum 1977). It is possible that such adaptation 1999b) and the early virgianids (mostly so-called Holorhynchus evolved originally in reaction to survival pressure in near-shore fauna; Rong et al. 2004). It is also noticeable that the Eodinobolus environments subjected to seasonal ice and snow melting at high kazakhstanicus group of species often occurs as a minor com- latitude. Their expansion to the low latitudes along the western ponent of rhynchonelliform brachiopod-dominated associations Gondwanan coast occurred during short-term cooling events in of the shallow and mid shelf, which is outside the main habitat the early to mid Darriwilian. Remarkably, these lingulide associ- of trimerellides (in restricted near-shore environments). ations settled in peri-Gondwanan Kazakhstanian terranes that The geographical distribution of the trimerellide Gasconsia were affected by tropical monsoons, where they inhabited near- in the late Katian also mirrors the distribution of the Holorhynchus shore tidal flats (Popov et al. 2007). Fauna (Popov et al. 1997; Rong et al. 2004). Ordovician trimerel- Recovery and proliferation of micromorphic linguliform bra- lide biogeography indicates that the microplates and island arcs chiopod associations took place in the late Tremadocian–Dapin- corresponding to some Kazakhstanian terranes together with gian interval. By the Darriwilian, they formed a characteristic South China were most probably situated in relative proximity to element of pantropical faunas. An important component of these the Australian sector of Gondwana, and by the Katian, Baltica faunas includes taxa descendant from Furongian faunas of tropical and South China had entered low latitudes. It is also possible peri-Gondwanan island arcs (acrotretide families Eoconulidae, that some Kazakhstanian terranes drifted westwards towards Ephippelasmatidae, Scaphelasmatidae and Torynelasmatidae; Baltica some time during the Late Ordovician (Popov et al. 1997). Popov & Holmer 1994, 1995). However, initial radiation of the The Ordovician trimerellide fauna from South China is unusual Ordovician acrotretide lineages related to the family Acrotretidae in that it contains taxa with excavated and vaulted visceral plat- occurred in Laurentia (Holmer et al. 2005). The lingulide genera forms; this pattern of shell morphology is otherwise unknown in Elliptoglossa and Paterula and the acrotretide family Biernatii- the majority of the Ordovician trimerellides with the exception dae probably dispersed from temperate-latitude Gondwana and of the Australian Belubula , but became widespread in the Baltica (Mergl 1999, 2002; Popov & Holmer 1994, 1995; Popov Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

ORDOVICIAN LINGULIFORM AND CRANIIFORM BRACHIOPODS 123 et al. 2007; Ghobadi Pour et al. 2011). Temperate-latitude micro- Appendix morphic brachiopod associations differ mainly in their lower diversity and in the absence of such taxa as Ephippelasma, Rhyso- Faunal list used in multivariate analysis treta, Spinilingula, Spondylotreta, Undiferina and torynelasma- tids, which were confined mainly to tropical latitudes. There are (1) Cuyania terrane, Lower Darriwilian, Ponon Trehue Formation (new): also a few taxa (e.g. Aipyotreta and Myotreta) that are confined Acrosaccus, Conotreta, Cyrtonotreta, Eoconulus, Numericoma, Nush- to temperate-latitude faunas. biella, Rowellella, Scaphelasma, Schizotreta. It is probable that the positions of major continents and climatic (2) Cuyania terrane, Upper Darriwilian, Pygodus serra to Pygodus anserinus belts were major controlling factors in the distribution of linguli- biozones (new): Acrosaccus, Biernatia, Conotreta, Cyrtonotreta, Dictyo- forms through the Ordovician, while in the longer term, the nites, Elliptoglossa, Eoconulus, Ephippelasma, Faveola, Acrotretidae oceans did not represent a significant barrier to their dispersal. gen. nov., Paterula, Rowellella, Rhysotreta, Scaphelasma, Schizotreta. Latitudinal differentiation of the micromorphic lingulate (3) Alborz terrane, Dapingian, Baltoniodus navis Biozone (Popov et al. 2008): brachiopod faunas is more evident during the early to mid Acanthambonia, Acrotreta, Aipyotreta, Alichovia, Biernatia, Numericoma, Darriwilian, but mainly disappeared by the late Darriwilian to Paterula, Rowellella, Scaphelasma, Schizotreta, Wahwahlingula. Sandbian, when pantropical taxa (e.g. Ephippelasma, Rhysotreta, (4) Alborz terrane, Darriwilian, Lenodus pseudoplanus Biozone (Popov et al. Spinilingula, Undiferina) spread to Baltica and South China. A 2008): Biernatia, Eoconulus, Paterula, Rowellella. possible explanation is that this is related to the suggested cool- (5) Bohemia (Perunica), Darriwilian, Sharka Formation (Mergl 2002): Cyrto- ing episode during the early to mid Darriwilian (Ghobadi Pour notreta, Eoconulus, Mytoella, Numericoma, Paldiskites, Paterula, Rowel- et al. 2007; Cherns & Wheeley 2009), which is most evident lella, Schizotreta, Spondyglossella, Wosekella. (6) Avalonia, Britain, Llandeilo Series (Sutton et al. 1999, 2000): Aipyotreta, from the regions of temperate-latitude peri-Gondwana, within Apatobolus, Bathmoleta, Eoconulus, Litoperata, Meristopacha, Opsiconi- the so-called ‘overlap zone’ (see also Fortey & Cocks 2003). dion, Paterula, Schizotreta, Tarphyteina, Trematis. The subsequent warming, which probably occurred synchro- (7) Laurentia, Alabama, Darriwilian, Pratt Ferry Formation, Pygodus serra nously, as indicated by the mid-Darriwilian carbon isotope excur- Biozone (Cooper 1956 plus new observations): Acanthambonia, Acrosac- sion (Bergstro¨m et al. 2008), resulted in a southern shift of the cus, Acrotreta, Biernatia, Conotreta, Cyrtonotreta, Dictyonites, Ellipto- climatic belts and the migration of some characteristic taxa from glossa, Eoconulus, Ephippelasma, Eurytreta, Faveola, Glossella, the tropics to higher latitudes (Ghobadi Pour & Popov 2009). Nushbiella, Paterula, Rowellella, Rhysotreta, Scaphelasma, Schizotreta, In the character of their proliferation and dispersion, cranii- Spinilingula, Spondylotreta, Talasotreta, Torynelasma, Trematis, Undifer- forms share similarities with other groups of the Palaeozoic Evol- ina, Westonia? utionary Fauna, including atrypide brachiopods, bryozoans and (8) Laurentia, Nevada, lower Darriwilian, Antelope Valley Limestone (Krause ostracods. There is no convincing Cambrian record of the & Rowell 1975): Conotreta, Cyrtonotreta, Dictyonites, Elliptoglossa, Subphylum Craniiformea, but it is likely that it originally Eoconulus, Eurytreta, Hansotreta, Acrotretidae gen. nov., Numericoma, evolved and dispersed from Gondwana and associated micro- Nushbiella, Paterula, Rowellella, Rhysotreta, Scaphelasma, Schizotreta, continents and island arcs. The earliest craniides are known Spinilingula, Spondylotreta, Torynelasma, Trematis, Undiferina. from the late Tremadocian of temperate- to high-latitude peri- (9) Kazakhstan, Chingiz terrane, lower Sandbian, Bestamak Formation Gondwana (Mergl 2002; Sdzuy et al. 2001), and they had a (Popov 2000a): Acrosaccus, Aktassia, Apatobolus, Biernatia, Conotreta, peri- Iapetus geographical distribution through most of the Ordovi- Cyrtonotreta, Elliptoglossa, Eoconulus, Ephippelasma, Eurytreta, Nush- cian. They are very rare or absent in tropical Gondwana, South biella, Paterula, Rowelella, Scaphelasma, Schizotreta, Torynelasma, China and Kazakhstanian terranes, and are not yet documented Veliseptum. from Siberia. (10) Kazakhstan, Chu–Ili terrane, upper Darriwilian, Uzunbulak Formation (Nikitina et al. 2006): Biernatia, Cyrtonotreta, Elliptoglossa, Eoconulus, The trimerellides probably evolved in a tropical peri- Eurytreta, Numericoma, Nushbiella, Paterula, Pomeraniotreta, Rowel- Gondwanan island arc setting (Chingiz terrane). Their dispersion lella, Rhysotreta, Scaphelasma, Schizotreta, Spinilingula, Spondyglos- and major features of biogeography mirror those of atrypoids, sella, Talasotreta, Torynelasma, Westonia? which were also characterized by lecithotrophic larvae (Freeman (11) Kazakhstan, Atasu–Zhamshi microplate, lower Darriwilian, Kurchilik & Lundelius 2005) and the early virgianids (Popov et al. 1997). Formation (Popov 2000a): Cyrtonotreta, Elliptoglossa, Issedonia, Craniiform biogeography reflects the approach of Avalonia Lurgiticoma, Numericoma, Rhysotreta, Scaphelasma, Spinilingula, towards Baltica some time in the Sandbian (e.g. migration of Westonia? Orthisocrania to Avalonia). The first appearance of the trimerel- (12) Kazakhstan, Chu–Ili terrane, lower Darriwilian, Karakan Limestone lide Eodinobolus in Laurentia during the early Katian suggests (Nazarov & Popov 1980): Cristicoma, Cyrtonotreta, Dictyonites, Eoconu- relative proximity to the Australian sector of Gondwana at that lus, Elliptoglossa, Faveola, Fascicoma, Numericoma, Paterula, Rhyso- time. However, the direct faunal exchange between Laurentia treta, Rowellella, Spinilingula, Scaphelasma, Schizotreta, Talasotreta, and equatorial Gondwana became minimized by the mid Katian. Westonia? The major route of trimerellide migration pathways was directed (13) New Zealand, mid Darriwilian, Thompson Creek, NW Nelson (allochtho- to the west, towards Baltica and Siberia, probably because of nous limestone) (Percival et al. 2009): Hyperobolus?, Spinilingula, Schi- numerous island arcs and microplates (concentrated near the zotreta, Trematis, Cyrtonotreta, Physotreta?, Scaphelasma, Torynelasma, equator west of Gondwana), which may have served as bridges for Lurgiticoma?, Undiferina, Nushbiella. those groups of benthos characterized by specific environmental (14) South China, Lenodus variabilis Biozone (Zhang 1995 and new obser- requirements and low dispersion potential. The pantropical distri- vations): Eoconulus, Scaphelasma, Torynelasma, Myotreta, Numericoma. bution of the trimerellides Eodinobolus and Monomerella during (15) South China, Eoplacognathus suecicus Biozone; Eoconulus, Ephippe- the Katian may be related to the ecological tolerance of these lasma, Myotreta, Numericoma, Scaphelasma, Torynelasma. (16) South China, Pygodus anserinus Biozone (Zhang 1995 and new obser- taxa, because they often occur together with various associations vations): Acanthambonia, Biernatia, Eoconulus, Eurytreta, Myotreta, dominated by rhynchonelliform brachiopods, outside the main Numericoma, Scaphelasma, Torynelasma, Sasyksoria, Undiferina. habitat of trimerellides. (17) Baltica, lower Darriwilian, Kunda Regional Stage (Holmer 1989 and new data): Acrotreta, Biernatia, Conotreta, Elliptoglossa, Eoconulus, Glos- L. Popov, M. Ghobadi and M. Bassett acknowledge logistical and financial sella, Hansotreta, Lingulasma, Myotreta, Numericoma, Paterula, Rowe- support from the National Museum of Wales. The work of Lars Holmer was sup- lella, Scaphelasma, Schizotreta, Siphonotreta, Torynelasma. ported by grants from the Swedish Research Council (VR 2009-4395, 2012- (18) Baltica, middle Darriwilian, Azeri Regional Stage (Holmer 1989): 1658). Research in Iran by M. Ghobadi Pour was supported by Golestan Univer- Acanthambonia, Biernatia, Conotreta, Cyrtonotreta, Elliptoglossa, Eoco- sity. I. Percival publishes with permission of the acting Director of the Geological nulus, Faveola, Glossella, Myotreta, Numericoma, Nushbiella, Paterula, Survey of New South Wales. Rowellella, Scaphelasma, Schizotreta, Siphonotreta, Torynelasma. Downloaded from http://mem.lyellcollection.org/ by guest on November 27, 2013

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