Journal of Earth Science, Vol. 25, No. 1, p. 1 44, February 2014 ISSN X Printed in China DOI: /s

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1 Journal of Earth Science, Vol. 25, No. 1, p. 1 44, February 2014 ISSN X Printed in China DOI: /s Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis: Additional Evidence from the Lower Triassic of the Russian Far East and Kazakhstan Yuri D Zakharov*, Alexander M Popov Far Eastern Geological Institute, Far Eastern Branch, Russian Academy of Sciences, Stoletiya Prospect 159, Vladivostok , Russia ABSTRACT: After the End-Permian mass extinction, ammonoids reached levels of taxonomic diversity higher than in the Changhsingian by the Dienerian Substage of the Induan. However, brachiopods exhibit a prolonged delay in recovery, and their taxonomic diversity had not recovered to Late Permian levels even by the Olenekian. The differential patterns of recovery between these two clades may reflect fundamental differences in physiology and behavior. Brachiopods were benthic organisms that were dependent on specific trophic sources, and their general reduction in size during the Early Triassic may have been a response to a relative paucity of food resources. In contrast, ammonoids were sluggishnektic organisms that utilized a wider range of trophic sources and that suffered no comparable size decrease during the Early Triassic. Brachiopods may have been at a disadvantage also due to vulnerabilities associated with their larval stage, during which they had to locate a suitable substrate for settlement. In contrast, ammonoids had no larval stage and juveniles may have been dispersed widely into favorable habitats. These factors may account for differences in the relative success of ammonoids and brachiopods at high-latitude regions following the End-Permian mass extinction: ammonoids successfully recolonized the Boreal region during the Early Triassic whereas brachiopods did not. KEY WORDS: Lower Triassic, South Primorye, Kazakhstan, brachiopod, ammonoid, biotic recovery. 1 INTRODUCTION The Permian-Triassic boundary (PTB) is associated with the largest extinction event of Phanerozoic time (e.g., Kozur, 2007; Yin and Zhang, 1996; Erwin, 1994). Early Triassic successions of both benthonic and nektonic marine organisms contain important information about the postextinction biotic recovery interval. However, in spite of significant progress in the past decades in investigation of Early Triassic brachiopods (e.g., Ruban, 2009; Chen et al., 2006, 2005a, b; Shen and Shi, 1996; Shen and He, 1994; Xu and Grant, 1994; Jordan, 1993; MacFarlan, 1992; Chen, 1983; Xu and Liu, 1983) and ammonoids (e.g., Brühwiler et al., 2012a, b, 2010a, b, c, 2008; Smyshlyaeva and Zakharov, 2012; Ware et al., 2011; Guex et al., 2010; Brayard et al., 2009a, b, c, 2007a, b, 2006a, b; Shigeta et al., 2009; Brayard and Bucher, 2008; Mu et al., 2007; Shevyrev, 2006, 1986; Krystyn et al., 2003; Ermakova, 2002; Zakharov, 1997, 1996, 1978; Dagys and Ermakova, 1996, 1988; Krystyn and Orchard, 1996; Waterhouse, 1996; Tozer, 1994; Guex, 1978), our knowledge of these important invertebrate groups, especially Mesozoic-type Brachiopodaremains incomplete. *Corresponding author: yurizakh@mail.ru China University of Geosciences and Springer-Verlag Berlin Heidelberg 2014 Manuscript received June 11, Manuscript accepted September 28, Early Triassic (mainly Olenekian) ammonoids are abundant and diverse in many regions of the world, but the most diverse Olenekian brachiopod faunas have been discovered in the former USSR area (South Primorye and Mangyshlak). The goal of this work is to analyze taxonomic diversity patterns of Early Triassic articulated brachiopods and ammonoids from the former USSR (Fig. 1) in order to reconstruct biotic recovery patterns after the End-Permian mass extinction. 2 MATERIALS Original materials on Early Triassic invertebrates used for our investigation were obtained by the authors from Induan and Olenekian sections of South Primorye during the last five years. Investigation of Olenekian brachiopods from Mangyshlak, Kazakhstan was made on the basis of Zakharov Y D s collection from his Kara-Tau expedition in LOWER TRIASSIC AMMONOID AND BRACHIO- POD BIOSTRATIGRAPHY 3.1 South Primorye (Russian Far East) The Early Triassic epiplatformal terrigenous assemblage of South Primorye (Fig. 1) lies unconformably on the Upper Palaeozoic and more ancient marine and continental sedimentary and volcano-sedimentary deposits, volcanics and granitoids (Zakharov et al., 2010; Zakharov, 1978, 1968). In many areas of South Primorye (e.g., Russian Island, Ussuri and Amur gulfs, Abrek Bay and Artyom area), the bulk of the Induan or Zakharov, Y. D., Popov, A. M., Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis: Additional Evidence from the Lower Triassic of the Russian Far East and Kazakhstan. Journal of Earth Science, 25(1): 1 44, doi: /s

2 2 Yuri D Zakharov and Alexander M Popov Induan to earliest Olenekian Lazurnaya Bay Formation consists of shallow-marine shelf sediments. These are predominantly conglomerate and sandstone with lenses of sandy limestone-coquina dominated by bivalves. The lower part of the Induan is represented by the Tompophiceras ussuriense Zone, its upper part corresponds to the Gyronites subdharmus Zone (Figs. S1 and S2). The Tobizin Cape Formation, of Smithian age, is present on Russian Island and on the western coast of Ussuri Gulf, where it conformably overlies the Lazurnaya Bay Formation. It is represented by sandstone with lenses of coquinoid calcareous sandstone with mainly cephalopod and bivalve remains. On Russian Island, the following zones can be recognized: Hedenstroemia bosphorensis below and Anasibirites nevolini above (Figs. S3 and S4). The Schmidt Cape Formation of early Spathian age is exposed only on Russian Island and consists of sandstone with thick lenses of sandy limestone. It is represented by a single zone, the Tirolites-Amphistephanites Zone (Figs. S3 and S6). The Zhitkov Cape Formation of Smithian to Spathian age represents deeper shelf facies characterised by the dominance of mudstone and siltstone with numerous calcareous-marly nodules and lenses and thin, irregular beds of sandstone. The lower part of the formation, which is found in the West Ussuri Gulf-Strelok Strait area (Figs. S1 and S7), is represented by the Hedenstroemia bosphorensis and Anasibirites nevolini zones and is Smithian in age. The upper part of the formation was formed in this area during the Early (Tirolites-Amphistephanites Zone) and Late (Neocolumbites insignis and Subfengshanites multiformis zones) Spathian. On Russian Island (Figs. S5 S7) and the western coast of Amur Gulf (Zakharov et al., 2005), only the upper Olenekian Zhitkov Cape Formation is present. However, the 44 N Razdol naia ( a) N 80 N 60 USSURIYSK Rakovka Rakovka Borisovka Zanadvorovka 40 N Razdol'naya Razdol'noye 70 Komarovka Vol no-nadezhdinskoe Tavrichanka 2 Perevoznaia Bol. Kiparisovka Ivniachka Uglovoe 90 Kaimanovka Zavodskoy Artyom Barsukovka Study area A 110 E vka o rtyom km Steklyanukha 40 N Bogataya Smolyaninovo 43 Popov Isl. Amur Gulf 3 4 Vladivostok Russian Island Ussuri Gulf Bol shoi Kamen Fokino 2 Sukhodol (b) 43 Reyneke Isl. 7 Dunai 0 15 km Sea of Japan Askol d Isl. Putyatin Isl. 132 E Figure 1. (a) Location of study areas in the former USSR. 1. South Primorye and 2. Mangyshlak. (b) Study sections in South Primorye area. 1. Seryj-Tri Kamnya; 2. Abrek; 3. Konechnyj; 4. Tobizin; 5. Ayax-Balka; 6. Zhitkov; 7. Golyj; 8. Paris.

3 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 3 Olenekoceras-bearing part of the Spathian on the western coast of Amur Gulf is characterised mainly by fine-grained, striped sandstone (Atlasov Cape Formation). The Karazin Cape Formation represents the Anisian everywhere in South Primorye. No benthic forms of allochthonous origin have been found in this formation and therefore it seems to have accumulated under offshore environmental conditions. It composed mainly of fucoid sandstone with large calcareous septarian nodules. The Lower Anisian in South Primorye is represented by the Ussuriphyllites amurensis and Leiophyllites pradyumna zones (Figs. S5 S7). The detailed lithological succession and distribution of brachiopod and ammonoid species in the Lower Triassic of South Primorye are shown in some figures (Figs. S1 and S8). Figure 2 illustrates the stratigraphic distribution of brachiopods and ammonoids at the family and order levels (within the Upper Changhsingian-Spathian interval) Tompophiceras ussuriense Zone (Griesbachian) The ammonoid assemblage of the Griesbachian Tompophiceras ussuriense Zone (at least 18 m thick) is characterised by the presence of two ceratitid ammonoid genera: Tompophiceras (T.), occurred together with inarticulated brachiopods (Lingula sp.) and some bivalves, and Lythophiceras (Lythophiceras sp.), the latter occurring in the upper part of the zone. For detailed ammonoid distribution data, see Fig. S1. However, no Griesbachian articulated brachiopods were found in the Far East apparently because of their extreme rarity there. A remarkable feature of the Griesbachian ceratitid ammonoid assemblage from South Primorye is that Tompophiceras is a representative of the Palaeozoic-type family (Dzhulfitidae) (Fig. 2). However, Lythophiceras belongs to a Mesozoic-type family (Ophiceratidae) Gyronites subdharmus Zone (Dienerian) The Dienerian Gyronites subdharmus Zone ( m thick) is marked by the first appearance datum of the ceratitid ammonoid genus Gyronites (G. subdharmus Kiparisova), used here for recognition of the base of the Dienerian. Other ceratitid ammonoid genera in the lower part of the zone are Lythophiceras (L. eusakuntala), Proptychites (P. hiemalis and possibly Tompophiceras (Tompophiceras sp.), in the upper part Ussuridiscus (U. varaha), Wordieoceras (W. cf. wordiei), Dunedinites (D. magnumbilicatus), Melagathiceratidae (Melagathiceratidae gen. et sp. indet.), and Pachiproptychites (P. otoceratoides). All these ceratitid ammonoids of the Gyronites subdharmus Zone, with the exception of the Dzhulfitidae (Tompophiceras) and Proptychitidae (e.g., Proptychites and Pachiproptychites), are representatives of typical Mesozoictype families (Ophiceratidae, Meekoceratidae, Melagathiceratidae, and possibly Paranannitidae). The Paleozoictype Dzhulfitidae was discussed earlier, but the ancestral group of the Proptychitidae is unknown at present (Fig. 2). The articulated brachiopod assemblage of the Dienerian Gyronites subdharmus Zone is characterised by the presence of four families of the order Rhynchonellida (Figs. 2 and 3): Pontisiidae (Lissorhynchia sp., Pontisiinae gen. and sp. nov.), Wellerellidae (Wellerellidae gen. and sp. nov. A, Wellerellidae gen. and sp. nov. B), Norellidae (Piarorhynchella? sp. nov.) and Rhynchonellidae (Abrekia sulcata, Rhynchonellidae gen. and sp. nov.). For detailed biostratigraphic distribution data, see Figs. S1 and S2. Among these brachiopod families, the Pontisiidae and the Wellerellidae are Paleozoic-type ones, but the Norellidae and the Rhynchonellidae seem to have originated during the Early Triassic (Induan). Newly found Dienerian brachiopod shells in South Primorye, including specimens of Abrekia, first described by A. S. Dagys (see Supplement), are generally very small as compared with Late Permian ones (Tables S2 S4) their maximal shell length is only 7.8 mm Hedenstroemia bosphorensis Zone (Smithian) The early Smithian Hedenstroemia bosphorensis Zone has been subdivided into a lower unit ( Gyronites separatus (=Ussuriflemingites abrekensis) beds) and an upper unit (Euflemingites prynadai beds) (Zakharov, 1997). The 16 to 50-mthick Gyronites separatus beds are characterised by the occurrence of the prolecanitid ammonoids Aspenitidae (Parahedensroemia sp., P. kiparisovae) and Sageceratidae (Pseudosageceras cf. multilobatum). Numerous ceratitid ammonoids also occur: Meekoceratidae (Ambitoides orientalis, Gurleyites sp.), Flemingitidae (Ussuriflemingites abrekensis), Xenoceltitidae (Preflorianites cf. radiatus), Proptychitidae (Paranorites varians, Kummelia? sp.), Arctoceratidae (Arctoceras septentrionale), Clypeoceratidae (Clypeoceras spitiense), and Columbitidae (Proharpoceras carinatitabulatum). The Euflemingites prynadai beds, which are at least 106 m thick, are also characterised by the presence of prolecanitid ammonoids Hedenstroemiidae ( Hedenstroemia bosphorensis), Aspenitidae (Parahedenstroemia conspicienda), Sageceratidae (e.g., Pseudosageceras multilobatum), Ussuriidae (e.g., Ussuria schamarae), and numerous ceratitid ammonoids: Meekoceratidae (e.g., Meekoceras subcristatum, Abrekites edites, A. planus), Dieneroceratidae (e.g., Dieneroceras chaoi), Inyoitidae (Inyoites spicini), Prionitidae (Radioprionites abrekensis), Flemingitidae (Euflemingites prynadai, E. artyomensis, Flemingites radiatus, F. aff. glaber, Balhaeceras balhaense, Rohillites laevis, Palaeokazakhstanites ussuriensis), Palaeophyllitidae (Anaxenaspis orientalis, Eophyllites sp., E. ascoldiensis), Xenoceltitidae (Preflorianites cf. radiatus), Arctoceratidae (e.g., Arctoceras septentrionale), Clypeoceratidae (Clypeoceras timorense), Melagathiceratidae (Juvenites simplex, J. dieneri, J. cf. septentrionalis, J. aff. sinuosus, Prosphingitoides ovalis), and Owenitidae (Owenites koeneni). The Early Smithian ammonoid succession is dominated by ceratitid ammonoids, all of which seem to be Mesozoic-type ones, as well as by four other prolecanitid ammonoid families. No articulated brachiopods were found in the lower part of the Hedenstroemia bosphorensis Zone. The articulated brachiopod assemblage of the Euflemingites prynadai beds is characterised by species of the families Wellerellidae (Wellerellidae gen. and sp. nov.), Norellidae (Piarorhynchella? sp. nov.), and Rhynchonellidae (Abrekia sulcata). For detailed biostratigraphic distribution data, see Figs. S1 and S4. Among these three early Smithian articulated brachiopod families, only the Wellerellidae is a Palaeozoic-type one (Fig. 2). The

4 4 Yuri D Zakharov and Alexander M Popov Ammonoids Brachiopods Order Ceratitida Prolecanitida Terebr. Sp. At. Rhynchonellida Permian Triassic Changh. Induan Olenekian - Griesbachian Dienerian Smithian Spathian Pleuronodoceratidae Wellerellidae Rhynchonellidae Huananoceratidae Xenodiscidae Ophiceratidae Tompophiceras Dzhulfitidae Lissorhynchia Dieneroceratidae Arctoceratidae Clypeoceratidae Pontisiidae Piarorhynchella? Norellidae Abrekia Hustedtiella Neoretziidae Lepismatina Lepismatinidae Fletcherithyris Dielasmatidae Proptychitidae Zeilleriidae? Meekoceratidae Sageceratidae Tirolitidae Khvalynitidae Ussuriidae Aspenitidae Hedenstroemiidae Inyonitidae Prionitidae Xenoceltitidae Melagathiceratidae Flemingitidae Palaeophyllitidae Keyserlingitidae Olenikitidae Stephanitidae Paranannitidae Procarnitidae Owenitidae Columbitidae Figure 2. Stratigraphic ranges of latest Permian and Early Triasic articulated brachiopod and ammonoid families in South Primorye. Dark and light gray fields indicate known and inferred ranges of families; dots and horizontal lines indicate occurrences of some important (named) genera. Also shown are inferred evolutionary relationships among ammonoid families. Abbreviations: Changh.. Changhsingian; Terebr.. Terebratulida; Sp.. Spiriferinida; At.. Athyridida.

5 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 5 Figure 3. Articulated brachiopods from the Lower Triassic of South Primorye and Mangyshlak. 1 and 2. Lissorhynchia sp., two views of same specimen No , x 2, Seryj Cape, Dienerian; 3 6. Prelissorhynchia (?) sp. nov., four views of same specimen Nos (4), x 2, Dolnapa, upper Spathian; Rhynchonellaceae gen. and sp. indet., four views of same specimen No (1), x 2, Tri Kamnya Cape, Smithian; Rhynchonellidae gen. and sp. nov. A, four views of same specimen No (2), x 2, Tri Kamnya Cape, Smithian; Rhynchonellidae gen and sp. nov. 2, four views of same specimen No , x 2, Tri Kamnya Cape, Smithian; Lissorhynchia sp. nov., four views of same specimen No , x 2, Dolnapa, Upper Spathian; Hustedtiella planicosta Dagys, four views of same specimen No , x 2, Dolnapa, upper Spathian; Spirigerellina sp., four views of same specimen No (2), x 2, Dolnapa, Upper Spathian; Lepismatina mansfieldi (Girty), four views of same specimen No (12), x 2, Dolnapa, Upper Spathian; Spiriferinidae gen. and sp. nov. A, four views of same specimen No. 20P, x 1, Paris Bay, upper Spathian.

6 6 Yuri D Zakharov and Alexander M Popov Lilliput effect continuous to be evident among early Smithian brachiopods of these families, which have maximal shell lengths of only 9.2 mm (observation from 53 specimens) (Table S4) Anasibirites nevolini Zone (lower Olenekian) The late Smithian Anasibirites nevolini Zone (20 68 m thick) is marked by the first appearance datum of the genus Anasibirites. The assemblage is characterised by rare occurrences of the prolecanitid ammonoids Sageceratidae (Pseudosageceras sp. indet.) and Aspenitidae (Parahedenstroemia nevolini), but by numerous ceratitid ammonoids, represented by Meekoceratidae (e.g., Meekoceras subcristatum), Tirolitidae (Bandoites elegans), Prionitidae (e.g., Hemiprionites contortus, Shigetaceras dunajense, Arctoprionites ovalis, Anasibirites nevolini, Wasatchites sikhotealinensis, Gurleyites maichensis), Palaeophyllitidae (Burijites skorochodi, Anaxenaspis sp. nov.), Xenoceltitidae (Preflorianites? sp.), Arctoceratidae (e.g., Arctoceras labogense, Churkites syaskoi), Melagathiceratidae (Juvenites simplex), Paranannitidae (Paranannites minor, Prosphingitoides ovalis), and Owenitidae (Owenites koeneni). For detailed ammonoid distribution data, see Figs. S3 and S4. The late Smithian ammonoid succession is dominated by ceratitid ammonoids, all of which seem to be Mesozoic-type ones, as well as by two other prolecanitid ammonoid families (Fig. 2). Very rare and small rhynchonellid brachiopods (up to 6 mm in length) were found in the zone, but their preservation is often moderate to poor in clayey facies Tirolites-Amphistephanites Zone (lower Spathian) The early Spathian Tirolites-Amphistephanites Zone (40 50 m thick) is marked by the first appearance datum of the genera Tirolites and Amphistephanites, used here for recognition of the base of the Spathian. The assemblage is characterised by the prolecanitid ammonoids Sageceratidae (Pseudosageceras sp.) and Aspenitidae (Parahedenstroemia sp.) and by ceratitid ammonoids, represented by Meekoceratidae (Bajarunia dagysi), Tirolitidae (Tirolites ussuriensis, T. subcassianus, Bandoites elegans), Dinaritidae (Tchernyschewites costatus), Olenikitidae (e.g., Kazakhstanites sonticus), Stephanitidae (Amphistephanites parisensis) and Flemingitidae (Guangxiceras tobisinense). The early Spathian ammonoids assemblages of the Tirolites-Amphistephanites Zone are dominated by Ceratitida, whereas Prolecanitida are very rare. All recognized ammonoid families seem to be Mesozoic-type ones. This time interval studied in South Primorye is characterised by the presence of abundant articulated brachiopods, represented by the next families of the orders Rhynchonellida, Athyritida, Spiriferinida, and Terebratulida: Rhynchonellidae (e.g., Rhynchonellidae gen. and sp. nov. A), Neoretziidae (e.g., Hustedtiella planicosta), Lepismatinidae (Lepismatina aff. mansfieldi), Diplospirellidae (Spirigerellina pygmaea), Dielasmatidae ( Fletcherithyris margaritovi), Zeilleriidae? (Zeilleriidae? gen. and sp. indet.), and Zeilleriidae? (e.g., Zeilleriidae? gen. and sp. nov. A). For detailed biostratigraphic distribution data (see Figs. S3 and S8). Observed ranges across the Tirolites-Amphistephanites Zone are summarized in Fig. 2. However, of the five brachiopod families known from this stratigraphic interval, two (Neoretziidae and Dielasmatidae) are Palaeozoic-type ones. The shell sizes of all these early Spathian brachiopod families are quite reduced (Tables S5 and S6)), with the exception of some dielasmatid brachiopods (e.g., Fletcherithyris margaritovi) that are up to 27 mm long (Table S7) Neocolumbites insignis Zone (middle Spathian) The middle Spathian Neocolumbites insignis Zone (66 95 m thick) is marked by the first appearance datum of the genus Neocolumbites. The ammonoid assemblage is characterised by species of both the Prolecanitida and Ceratitida orders. Ceratitid ammonoids are represented by Meekoceratidae (Svalbardiceras zhitkoviense), Tirolitidae (Tirolites cf. subcassianus Zakharov), Palaeophyllitidae (Burijites skorochodi, Leiophyllites praematurus), Keyserlingitidae (e.g., Olenekoceras meridianus), Columbitidae (e.g., Neocolumbites insignis, Columbites ussuriensis, Procolumbites subquadratus, Hellenites inopinatus), and Procarnitidae (Procarnites sp.). The prolecanitid ammonoids Sageceratidae (Pseudosageceras sp.) and Khvalynitidae (Khvalynites unicus) are present in lower abundance. The ammonoid assemblages of this zone are dominated by Ceratitida. All middle Spathian ammonoid families seem to be Mesozoic-type ones. Small spiriferinid brachiopod shells, up to 12 mm in length, were found in this zone Subfengshanites multiformis Zone (upper Spathian) The late Spathian Subfengshanites multiformis Zone (16 18 m thick) is marked by the first appearance datum of the genus Subfengshanites. This zone is characterised by the occurrence of abundant ceratitid and very rare prolecanitid (Pseudosageceras sp.) ammonoid species. The main ceratitid ammonoid taxa known from this level are the Dieneroceratidae (Dieneroceras karasini), Palaeophyllitidae (Palaeophyllites superior, Leiophyllites sp.), Paranannitidae (e.g., Zhitkovites insularis (Kiparisova), Pseudoprosphingites globosus, Isculitoides? suboviformis), Columbitidae (Subfengshanites multiformis, Arnautoceltites gracilis, and Prenkites aff. timorensis). All recognized ammonoid families seem to be Mesozoic-type ones. The Subfengshanites multiformis Zone is characterised also by the occurrence of Rhynchonellidae (Rhynchonellidae gen. et sp. nov. 1, 2 and 3), Lepismatinidae (Lepismatinidae gen. and sp. nov.), and Neoretziidae (Hustedtiella planicosta). For detailed biostratigraphic distribution data and rhynchonellid brachiopod shell size (see Figs. S5, S8, and Table S4). Observed ranges across the Subfengshanites multiformis Zone are summarized in Fig. 2. Among brachiopod families only the Neoretziidae crossed the PTB 3.2 Mangyshlak (Kazakhstan) The Spathian in Mangyshlak has been investigated in detail in the Dolnapa Well reference section (e.g., Zakharov et al., 2008; Gavrilova, 2007; Balini et al., 2000; Shevyrev, 1968), located in the Kara-Tau area. Two lithostratigraphic units, corresponding to the Olenekian Tjururpiniaya Series, can be recognized in this section: the Tartalinskaya Fm. (Tirolites cassianus-kiparisovites carinatus and Columbites parisianus- Procolumbites zones), which is composed of mudstone, silt-

7 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 7 stone and fine-grained sandstone with calcareous interlayers, and the Karadzhatykskaya Fm. (Arnautoceltites bajarunasi- Stacheites undatus Zone and Eumorphotis beds), which consists of intercalations of mudstone with calcareous boulders, siltstone with plant detritus, and fine-grained sandstone with interlayers of limestone (Fig. S9) Tirolites cassianus-kiparisovites carinatus Zone (Spathian) The ammonoid assemblage of the early Spathian Tirolites cassianus-kiparisovites carinatus Zone (which is at least 113 m thick) is characterised by the presence of two ammonoid orders: the Prolecanitida (Sageceratidae Pseudosageceras longilobatum) and the Ceratitida. The latter consists of Dinaritidae ( Dinarites asiaticus, D. orientalis), Olenikitidae (Tjururpites cf. costatus, Kiparisovites carinatus, Hyrcanites nodosus, Kazakhstanites dolnapaensis), Tirolitidae (e.g., Tirolites cf. cassianus),?prionitidae (Albanites gracilis) and Doricranitidae (Doricranites) (Fig. 4). The early Spathian Order Ammonoids Brachiopods Ceratitida Prolec. Terebr. Sp. At. Rhynch. Smithian Triassic Upper Olenekian Spathian Piarorhynchella Rhynchonellidae Prelissorhynchia (?) Pontisiidae Piarorhynchella Lissorhynchia Norellidae Hustedtiella Neoretziidae Spirigerellina Diplospirellidae Lepismatina Lepismatinidae Fletcherithyris Dielasmatidae Antezeilleria Antezeilleriidae Thyratryaria Family uncertain Proanadyrella? Sageceratidae Khvalynitidae Procarnitidae Dinaritidae Xenoceltitidae Olenikitidae Tirolitidae Kashmiritidae Doricranitidae Prionitidae Palaeophyllitidae Columbitidae Figure 4. Stratigraphic range of Olenekian articulated brachiopod and ammonoid families in Mangyshlak. Abbreviations: Prolec.. Prolecanitida; Rhynch.. Rhynchonellida. See Fig. 2 for further details. ammonoid assemblages of the Tirolites cassianus-kiparisovites carinatus Zone are dominated by Ceratitida, and no Palaeozoic-type ammonoid families were identified. Articulated brachiopods are represented by three orders (Rhynchonellida, Spiriferida, and Terebratulida) and several families: Norellidae (Piarorhynchella mangyshlakensis), Lepismatinidae (Lepismatina sp. nov.), and Dielasmatidae ( Fletcherithyris margaritovi) (Fig. 4). No Palaeozoic-type brachiopod families were discovered in this zone. As in the case with the Lower Triassic of South Primorye, size reduction ( Lilliput effect ) has been observed in many brachiopods from the Tirolites cassianus-kiparisovites carinatus Zone (Table S4), with the exception for very rare Dielasmatidae (up to 20 mm in length) Columbites parisianus-procolumbites Zone (middle Spathian) The ammonoid assemblage of the middle Spathian Columbites parisianus-procolumbites Zone (274 m thick) is characterised by the presence of prolecanitid ammonoids Sageceratidae (Pseudosageceras longilobatum) and Khvalynitidae (Khvalynites mangyshlakensis) and abundant ceratitid ammonoids: Dinaritidae ( Dinarites asiaticus, D. orientalis), Xenoceltitidae (Xenoceltites mangyshlakensis, X. bajarunasi, Preflorianites kiparisovae), Kashmiritidae (Kashmirites subdimorphus, K. popowi Shevyrev, Eukashmirites subdimorphus), Olenikitidae (Tjururpites costatus, Kiparisovites ovalis, Kazakhstanites dolnapaensis), Tirolitidae (Tirolites armatus, T. longilobatus, T. rossicus), Doricranitidae (Doricranites acutus, D. bogdoanus),?prionitidae (Albanites gracilis), Palaeophyllitidae (Leiophyllites exacudus), Procarnitidae (Procarnites kokeni), and Columbitidae (Columbites cf. parisianus, C. ventroangustus, Procolumbites karatauchicus, Hellenites kazakhstanicus, Mangyshlakites mirificus) (Fig. 4). The early Spathian ammonoid assemblages of the Columbites parisianus-procolumbites Zone are dominated by Ceratitida, and no Palaeozoic-type ammonoid families were recognized. Articulated brachiopods are represented by four orders (Rhynchonellida, Athyridida, Spiriferinida, and Terebratulida), among which a number of families were recognized: Pontisiidae (Prelissorynchia (?) sp. nov., Lissorhynchia sp. nov., Piarorhynchella mangyshlakensis), Neoretziidae (Hustedtiella planicosta), Diplospirellidae (Spirigerellina sp., Spirigerellinae gen. and sp. indet.), Lepismatinidae (Lepismatina sp. nov.), Antezeilleriidae (Antezeilleria sp.), and an uncertain family (Proanadyrella (?) sp., Thyratryaria sp. nov. A, Thyarotryaria sp. nov. B) (Fig. 4). The genus Prelissorynchia (Pontisiidae) possibly crossed the PTB; formerly, this genus was considered to be only Late Paleozoic in age. Brachiopod miniaturization is also recorded for this level, with shell lengths for all of the families above measuring between 3.4 and 9.4 mm (Tables S5 and S6) Arnautoceltites bajarunasi-stacheites undatus Zone (upper Spathian) The ammonoid assemblage of the late Spathian Arnautoceltites bajarunasi-stacheites undatus Zone (172 m thick) is characterised by the presence of prolecanitid ammonoids Sage-

8 8 Yuri D Zakharov and Alexander M Popov ceratidae (Pseudosageceras sp.) and some ceratitid ammonoid families: Dinaritidae ( Dinarites orientalis, Stacheites undatus, S. concavus), Xenoceltitidae sp., Olenikitidae (Kazakhstanites dolnapaensis, Tjururpites costatus, Kiparisovites ovalis), Tirolitidae (Tirolites armatus), Prionitidae (Arctoprionites sp., Albanites gracilis), and Columbitidae (Arnautoceltites bajarunasi). The late Spathian ammonoid assemblages of the Arnautoceltites bajarunasi-stacheites undatus Zone are dominated by Ceratitida, and no Palaeozoic-type ammonoid families were recognized. Three articulated brachiopod orders (Rhynchonellida, Spiriferinida, and Terebratulida) were found in the Arnautoceltites bajarunasi-stacheites undatus Zone. They are represented by the families Pontisiidae (Lissorhynchia sp. nov., Piarorhynchella mangyshlakites), Lepismatinidae (Lepismatina sp. nov.), and an uncertain family (Thyratryaria aff. pertumida). Among these brachiopod families, only a single Palaeozoic-type one (Pontisiidae) has been discovered. The Lilliput effect is also prevalent among the brachiopod fauna of the Arnautoceltites bajarunasi-stacheites undatus Zone. Lissorhynchia exhibits shell lengths to a maximum of 5.3 mm, and other latest Spathian brachiopods are only marginally larger (up to 7 10 mm) (Table S4) Eumorphotis beds (upper Spathian) In the uppermost part of the Spathian in the Dolnapa Well Section, the ~151-m-thick Eumorphotis beds yielded very rare prolecanitid ammonoids (Pseudosageceras sp.) together with bivalves, gastropods, and nautiloids. However, ceratitid ammonoids and articulated brachiopods have not been discovered here. 4 DISCUSSION 4.1 Overview of Early Triassic Brachiopod and Ammonoid Faunas Palaeozoic-type taxa About 13 families of articulated brachiopods survived the End-Permian mass extinction. Among them, 13 Palaeozoictype brachiopod genera have been identified in the Lower Triassic: Cathaysia, Paryphella, Spinomarginifera, Retimarginifera (?), Acosarina, Prelissorhynchia (?), Araxathyris, Spirigerella, Orbicoella, Crurithyris, Paracrurithyris, Paraspiriferina, and Tethyochonetes (e.g., Shen et al., 2006; Chen et al., 2005b; this study, see Table S1). They belong to the Paraspiriferinidae, Schizophoriidae, Productellidae, Ambocoellidae, Pontisiidae, and Athyrididae families. All these genera, with the exception of Orbicoelia, Prelissorhynchia (?) and possibly Retimarginifera, were discovered only in the lower Griesbachian. However, Retimarginifera (?) was reported from the Smithian (Chen et al., 2005b), Prelissorhynchia (?) from both the Griesbachian (Chen et al., 2005b) and the middle Spathian (this study), and Orbicoelia only from the upper Olenekian (Chen et al., 2005b). Besides, at least six additional Permiantype brachiopod genera belonging to six other families (Diplospirellidae, Neoretziidae, Laballidae, Pennospiriferinidae, Dielasmatidae, and Angustothyrididae) also may have survived into the Lower Triassic (Fig. 5). However, for certain only four ammonoid lineages (representing different taxonomic levels) survived the mass extinction: (1) Xenodiscidae and Dzhulfitidae (at the family level), (2) Otoceratoidea (at the superfamily level), and (3) Episageceras (at the generic level). Only a single Palaeozoic-type genus (Episageceras) is for certain known from the Lower Triassic (Fig. 5). In Siberia and the Russian Far East, representatives of Episageceras were collected from the lower and upper parts of the Induan (Dagys and Ermakova, 1996; Zakharov, 1978; Popow, 1961). About 18 Mesozoic-type genera have been reported from the Induan (Table S1). A few ammonoid genera (Hypophiceras fauna) have been recorded in the upper Changhsingian of the Meishan Section in South China, identified as Pseudogastrioceras, Otoceras (?), Hypophiceras, Metophiceras, Tompophiceras, Glyptophiceras, and Pseudosageceras (Yang et al., 1996; Yin et al., 1996). McGovan and Smith (2007) have shown in their Fig. 7 that Aldanoceras, as well as Otoceras, Xenodiscus, Hypophiceras, Tompophiceras, Metophiceras and Episageceras, crossed the PTB (established stratigraphic ranges are shown there as solid lines), using Dagys and Ermakova s (1996) data on the Verkhoyansk area and taking into account information on Hypophiceras fauna of the Meishan area. As was recently emphasized by Shevyrev (2006), ammonoids of the so-called Hypophiceras fauna in Meishan represent a mixed assemblage of Permian and Triassic forms. The composition of the assemblage at Meishan is in doubt because of poor preservation, where ammonoid shells are strongly deformed and suture lines are usually unobservable (Shevyrev, 2006). In this situation, some Permian araxoceratid ammonoids, in Shevyrev s (2006) opinion, could have been mistaken for Triassic Otoceras, Permian Paratirolites or Pseudotirolites for Triassic Tompophiceras, and Permian Xenodiscus for Triassic Hypophiceras. However, the age of Otoceras-bearing sequences in the Boreal realm is still under debate now (e.g., Bjerager et al., 2006; Baud and Beuchamp, 2001; Stemmerik et al., 2001; Kozur, 1998; Orchard and Krystyn, 1998), and only further progress in the carbon-isotopic, palaeomagnetic and micropalaeontological investigation could solve this very important problem. According to Brayard et al. (2007a), the Smithian genus Proharpoceras probably was derived from the Wuchiapingian Anderssonoceratidae (Otoceratina), thus implying that an additional lineage among Ceratitida crossed the PTB. However, this inference concerning two taxa from distant stratigraphic levels needs to be tested through further morphological analysis and additional fossil finds. The new evidences and published data show that in contrast to the brachiopod fossils only restricted ammonoid lineages at the generic and family levels servived the End-Permian mass extinction Mesozoic-type taxa About 15 Mesozoic-type articulated brachiopod genera were found in the Induan (Table S1), some of which are known from Russia (e.g., Lissorhynchia, Abrekia, Rhynchonellidae gen. nov. 4, Piarorhynchella (?), Wellerellidae gen. nov. A, Wellerellidae gen. nov. B, and Pontisiinae gen. nov.). However, Chen et al. (2005a) reported only 7 Mesozoic-type genera from the Induan of South China (e.g., Shen et al., 2006; Chen et al.,

9 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis b; Shen and He, 1994), Idaho (Girty, 1927), Himalayas (Bittner, 1899), and North Caucasus (Dagys, 1974). The taxonomic diversity of brachiopods during the Dienerian was only about 11% of that for the Changhsingian (Fig. 6). Among Olenekian articulated brachiopods, only three Palaeozoic-type genera (Retimarginifera (?), Orbicoelia, and Prelissorhynchia (?)) but 40 Mesozoic-type genera are known. They are identified as Maorirhynchia (?), Rhynchonellidae gen. nov. 1, 2 and 3, Nudirostralina, Abrekia, Sinuplicorhynchia (?), Paranorellina, Piarorhynchella, Aparimarhynchia, Lissorhynchia, Spirigerellina, Spirigerellinae gen. nov., Neoretzia, Hustedtiella, Lepismatina, Spiriferinidae gen. nov. A, Figure 5. Post-extiction survival of Paleozoic-type brachiopods and ammonoids (Chen et al., 2005a, b; this study). Abbreviations: Capitan.. Capitanian; An.. Anisian. See Fig. 2 for Early Triassic substages.

10 10 Yuri D Zakharov and Alexander M Popov Number of genera Changh. Induan Olenek. Anisian Changh. Induan Perm. Mass extinction Ammonoids 68 (136) (129) (60) 27 (27?) 20 (23?) (16) (16) 12 0 Gri. Die. Smi. Spa. Gri. Die. Smi. Spa. Triassic Number of genera Perm. Mass extinction Articulated brachiopods Olenek. Triassic Anisian ? Postextinction miniaturization No any trends Postextinction miniaturization Occured Figure 6. Early Triassic generic richness recovery of the brachiopod and ammonoid faunas. Number in parenthesis means number of genera in substages; number above the horizontal arrow indicates quantity of genera crossed the corresponding boundary interval (e.g., Brühwiler et al., 2011a, b, 2010a, b; Ware et al., 2011; Guex et al., 2010; Shigeta et al., 2009; Brayard and Bucher, 2008; Zakharov et al., 2008; Mu et al., 2007; Brayard et al., 2006a, b; Shen et al., 2006; Chen et al., 2005a, b; Ermakova, 2002; Bogoslowskaya et al., 1999; Tozer, 1994; Dagys, 1974; this study). See Fig. 2 for Early Triassic substages. Spiriferinidae gen. nov. B, Obnixia, Fletcherithyris, Rhaetina, Portneufia, Antezeilleria, Zeilleriidae? gen. nov. A, B, C, D and F, Periallus, Protogusarella, Thyratryaria, Proanadyrella, and Vex, of which twelve are new genera (Table S1). Chen et al. (2005b) reported only 18 Mesozoic-type brachiopod genera in the Olenekian on the basis of data from South China, Southwest Japan, Idaho, Himalayas, Tibet, Dobrogea, New Zealand, and Spitsbergen. The taxonomic diversity of brachiopods during the Smithian and Spathian was only about 16% and 19% of that for the Changhsingian, respectively (Fig. 6). In this stage of our knowledge we conditionally eccept for the Induan ammonoids that they are represented by only a single Palaeozoic-type genus (Episageceras) but 67 Mesozoictype genera (Fig. S12). Only Mesozoic-type ammonoid genera have been reported from the Smithian and Spathian which have yielded 136 and 129 genera, respectively (Fig. 6; Table S1). McGovan (2005) inferred that ammonoids did not fully recover until the Spathian or Anisian. However, recent data (Brühwiler et al., 2011a, b, 2010a, b, c; Guex et al., 2010; Brayard et al., 2009a, b; this study) show that the recovery of ammonoid faunas proceeded significantly more rapidly. Brayard et al. (2009a, b) concluded that ammonoid diversity level was higher than the maximum level recorded during all the Permian by the Smithian. According to our data, based on recently published and original evidences, ammonoids reached levels of taxonomic diversity during the Dienerian that were 146% higher than in the Changhsingian, representing a much faster recovery than that for brachiopods (the diversity of brachiopods during the Dienerian was only about 8% of that for the Changhsingian) (Fig. 6). Maximum taxonomic diversity of Permian ammonoids is associated with the Wordian-Roadian (Bogoslovskaya et al., 1999). However, the diversity of ammonoids during the Smithian (Table S1) reached not less than 200% of that for the Wordian.

11 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis Patterns of Geographical Differentiation of Early Triassic Brachiopod and Ammonoid Faunas Early Triassic ammonoid diversification produced about 290 genera, while Early Triassic articulated brachiopod diversification produced only about 68 genera (Table S1). Griesbachian relict articulated brachiopods have been recovered from South China (e.g., Chen et al., 2005a, b), Salt Range (Grant, 1970), Nepal (e.g., Waterhouse, 1978), Kashmir (Shimizu, 1981) and Oman (Chen et al., 2005a, b; Twitchett et al., 2004). During Olenekian time, one group of Permian relict species continued to exist in Nepal, while another one migrated possibly to the Mangyshlak and Romanian areas. Localities of Induan Mesozoic-type articulated brachiopod are known only in lower to middle-latitude regions: South China (e.g., Chen et al., 2005b; Shen and He, 1994); Northwest Caucasus (Dagys, 1974), Himalayan region (Chen et al., 2005a, b; Bittner, 1899), South Primorye (Dagys, 1974; this study), and Idaho (Girty, 1927). However, the most diverse Induan brachiopod faunas were reported from South China (14 genera, including five Mesozoic-type genera) and South Primorye (six Mesozoic-type genera) (Fig. S10). No Induan articulated brachiopods have been discovered in high palaeolatitudes. Data of the present study show that the most diverse Olenekian articulated brachiopod faunas are present in the former USSR area (South Primorye and Mangyshlak). These faunas consist of 21 and 13 genera, respectively (Fig. S11). Other known localities of Olenekian articulated brachiopods include Idaho (Hoover, 1979; Perry and Chatterton, 1979; Girty, 1927), South China (Perry and Chatterton, 1979), Carpathians- Balkanian area (Jordan, 1993), Tibet (Chen et al., 2005b; Chen, 1983), Japan (Dagys, 1993), Spitsbergen (Dagys, 1993, 1974), Alps (Jordan, 1993), and New Zealand (MacFarlan, 1992). Among known Early Triassic articulated brachiopods, only a few Olenekian genera (Obnixia, Hustedtiella? (Dagys, 1974) and Aparimirhynchia (MacFarlan, 1992)) reached highlatitude areas. In contrast to Early Triassic brachiopods, Induan (Figs. S12) and Olenekian ammonoid assemblages, according to our data, are common for both the tropical-subtropical area and the high-latitude region of the Boreal realm, which is in agreement with previous studies on this topic (e.g., Brayard et al., 2009a, b, 2006b; Tozer, 1994). 4.3 Palaeophysiological and Behavioural Contexts of End- Permian Environmental Stresses The End-Permian Early Triassic interval is dominated by unusual oceanic and climatic conditions (e.g., oceanic anoxia, expansion of desert belts, declining atmospheric levels of O 2 but increasing of levels of CO 2, a biocalcification crisis; e.g., Galfetti et al., 2007; Payne and Kump, 2007; Kidder and Worsley, 2004; Isozaki, 1997; Wignall and Hallam, 1993; Baud et al., 1989). Oceanic anoxia has been invoked as the main kill mechanism of the Permian-Triassic extinction event (Wignall and Hallam, 1993), although this hypothesis has been disputed by Hermann et al. (2011) on the basis of the largely terrigenous origin of particulate organic matter in the Lower Triassic of the Salt Range and Surghar Range. Nevertheless, chemically and/or physiologically harsh environmental conditions certainly existed at the end of Permian, possibly caused by unusual oceanic and climatic conditions that affected the marine biota for several million years (Algeo and Twitchett, 2010). The largest effects were during the Griesbachian, which represents a survival phase for brachiopods and ammonoids following the End-Permian ecological crisis. The patterns of recovery of brachiopods and ammonoids following the End-Permian mass extinction differed owing to fundamental differences in their physiology and behavior. Brachiopods and ammonoids are representatives of physiologically dissimilar groups of marine organisms with possible metabolic and skeletal differences (Knoll et al., 2007). Brachiopods belong to a group of organisms with a calcium carbonate skeleton that was often massive with respect to supporting organic tissue and formed from fluids minimally buffered by physiology. Ammonoids, in contrast, belonged to a group of organisms with a calcium carbonate skeleton of moderate mass and formed from fluids that were relatively well buffered with respect to the factors that govern carbonate precipitation. The latter condition is considered to be more sensitive to high watermass P CO2 (hypercapnic stress). Differences in feeding behaviour may have mattered also. Data on brachiopod sizes changes in the Permian-Triassic transition in South China (e.g., Chen et al., 2005a, b) and our additional evidence on the Late Permian and Early Triassic brachiopods mainly of the former USSR (Tables S2 S7 and Fig. 7) is in an agreement with the hypothesis of Keller and Abramovich (2009) that high-stress conditions led to both biodiversity loss and size reduction ( Lilliput effect in Urbanek s (1993) sence). According to Harries and Knorr (2009), this effect represents a pronounced reduction in size of the biota associated with the aftermath of mass extinctions. However, by contrast with Early Triassic brachiopods, survived the End- Permian mass extinction, ammonoids from the same habitats/ facies in both the Primorye region and Kazakhstan do not show any trends toward smaller size in the Early Triassic. These different outcomes may have been related to differences in trophic resource availability (just after the End-Permian crisis, notably brachiopods may have suffered from a relative paucity of food resources). Epifaunal benthos (suspension-feeding brachiopods as well as bivalves) dominated Lower Middle Triassic marine environments, but the more modern life habits (e.g., infaunalization associated with burrowing suspension and deposit feeders) increased during the Late Triassic (Bonuso and Bottjer, 2008). Judging by the abundance and wide distribution of nektic ammonites during the Early Triassic, one can infer that they generally were not specialized feeders, although they were apparently presumably bottom feeders during the Late Palaeozoic and Mesozoic, because optimal temperatures of their growth are mainly comparable to those obtained from their co-occurring benthos on the shelf (e.g., Moriya et al., 2003; Smyshlyaeva et al., 2002). Thus, Early Triassic ammonoids had greater access to trophic resources and therefore stood a better chance of success than brachiopods. Another factor in the differential recovery of these two clades may have been related to differences in larval/juvenile survival and development. After hatching, brachiopod larvae and juvenile ammonoids may be dispersed with the aid of currents. All planktic brachiopod larvae are capable of delaying

12 12 Yuri D Zakharov and Alexander M Popov settlement until a suitable substratum has been located, although the motile larval stage usually lasts only days or hours (Williams et al., 1997). High sedimentation rates during the Early Triassic may have resulted in generally more mobile substrates that were unfavorable for larval brachiopod settlement (cf., Algeo and Twitchett, 2010). Ammonoids, in contrast, L ( mm ) ( N=11) 11.8 ( N=18) 42 ( N=100) Terebratulida 27 ( N=147) 10 L ( mm ) ( N=177) Mass-extinction Spiriferinida ( N=23) 9.4 L ( mm ) Athyritida 11.4 ( N=43) L ( mm ) ( N=114) 13.5 ( N=32) Rhynchonellida 8.7 ( N=105) ( N=54) L ( mm ) L Spiriferida ( N=107) 69 ( N=64) L ( mm ) ( N=16) 20 Productida 10 0 Lower Upper Lower Upper Wuchiapingian Changhsingian Upper Permian 13.7 ( N=15) Griesb. Diener. Smithian Spathian Induan Olenekian Lower Triassic Figure 7. Post-extinction miniaturization of brachiopods: evidence from the Upper Permian of Transcaucasia and North Caucasus (Tables S2 S4), Griesbachian of South China (Table S2; Chen et al., 2006), Dienerian and Smithian of South Primorye (Table S4) and Spathian of Mangyshlak (Tables S4 S6), South Primorye (Tables S4 S7) and the Western USA (Table S7; Hoover, 1979). N. the number of specimens used for our observation.

13 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 13 are characterised by direct ontogenetic development without a larval stage and, thus, were not dependent on substrate characteristics. The survival of planktic brachiopod larvae was significantly lower because of the short duration of their motile phase, the necessity of finding a suitable substrate, and the specificity of their trophic needs factors that worked to their disadvantage during the Early Triassic. The contrast in recovery patterns for brachiopods and ammonoids was most evident at high palaeolatitudes, most difficult apparently place for brachiopod larvae migration. 5 CONCLUSIONS 1. Ammonoids exceeded their pre-crisis taxonomic diversity and abundance by the late Induan (Dienerian Substage). In contrast, brachiopods never recovered their pre-crisis diversity and abundance anytime during the Triassic or later, in spite of the fact that about 13 brachiopod genera survived the End- Permian mass extinction (recent palaeontological and carbonisotopic records allow to assume that only restricted ammonoid lineages at both the generic and the family levels survived the mentioned mass extinction). 2. The differential recovery patterns of brachiopods and ammonoids seem to be a result of physiological and behavioural differences, reflected in their different life modes and distributions in ecospace. Specifically, brachiopods were at a disadvantage during the Early Triassic because of the short duration of their motile phases, the need to find a suitable substrate for larval settlement, and the specificity of their trophic requirements. 3. Brachiopod s lilliput effect developed during the Early Triassic, judging from published data and partly confirmed by some original evidences from South Primorye, Mangyshlak, North Caucasus and Transcaucasia, may have been a response to a relative paucity of food resources for this specific benthic organisms of that time. 4. Diversification of Early Triassic brachiopods at high latitudes was quite restricted: no articulated brachiopods of Induan age and only three genera of Olenekian age (Obnixia and Hustedtiella? (Dagys, 1974) and Aparimirhynchia (MacFarlan, 1992)) have been discovered in the Boreal realm. However, immediately after the End-Permian extinction event (and possibly just during it), ammonoids colonized and stably inhabited many regions in the Boreal realm throughout the Triassic. The contrast in recovery patterns for brachiopods and ammonoids was most evident in the high palaeolatitudes, apparently most difficult place for brachiopod larvae migrations. ACKNOWLEDGMENTS For help in finding some important Early Triassic brachiopod and ammonoid fossils in South Primorye and regular assistance, we are indebted to Y Shigeta (National Museum of Nature and Science, Tsukuba), H Maeda, and T Kumagae (Kyoto University) and also to V T S edin (Institute of Pacific Oceanology, Vladivostok). Our cordial thanks are due to Prof. T A Algeo (University of Cincinnati) for providing valuable editorial comments, Prof. Shuzhong Shen (Nanjing Institute of Geology and Palaeontology) and anonymous reviewer for stimulating remarks, as well as for their information on latest published papers on Early Triassic ammonoids and data on this topic in press, that substantially improved this paper. This work is a contribution to UNESCO-IUGS IGCP Project 572 and financially supported by the Russian grant RFBR (No a). REFERENCES CITED Algeo, T., Twitchett, R., Anomalous Early Triassic Sediment Fluxes due to Elevated Weathering Rates and Their Biological Consequences. Geology, 38(11): Balini, M., Gavrilova, V. A., Nicora, A., Biostratigraphical Revision of the Classic Lower Triassic Dolnapa Section (Mangyshlak, West Kazakhstan). Zentralblatt für Geologie und Paläontologie I, 1998: Baud, A., Beauchamp, B., Proposals for Redefinition of the Griesbachian Substage and for the Base of the Triassic in the Arctic Regions. Proceedings of the International Symposium The Global Stratotype of the Permian-Triassic Boundary and the Paleozoic-Mesozoic Events. Changxing Baud, A., Magaritz, M., Holser, W. T., Permian-Triassic of the Tethys: Carbon Isotope Studies. Geologische Rundschau, 78(2): Bittner, A., Himalayan Fossils. Trias Brachiopoda and Lamellibranchiata. Palaeontologica Indica, 15(2): 1 76 Bjerager, M., Seidler, L., Stemmerik, L., et al., Ammonoid Stratigraphy and Sedimentary Evolution across the Permian-Triassic Boundary in East Greenland. Geological Magasine, 143(5): Bogoslovskaya, M. F., Kuzina, L. F., Leonova, T. B., Classification and Distribution of Late Paleozoic Ammonoids. In: Rozanov, A. Y., Shevyrev, A. A., eds., Fossil Cephalopods: Recent Advances in Their Study. Izdatelstvo Akademii Nauk SSSR, Moscow (in Russian) Bonuso, N., Bottjer, D. J., A Test of Biogeographical, Environmental, and Ecological Effect on Middle and Late Triassic Brachiopod and Bivalve Abundance Patterns. Palaios, 23(1): Brayard, A., Bucher, H., Smithian (Early Triassic) Ammonoid Faunas from Northwestern Guangxi (South China): Taxonomy and Biochronology. Fossils and Strata, 56: Brayard, A., Bucher, H., Brühwiler, T., et al., 2007a. Proharpoceras Chao: A New Ammonoid Lineage Surviving the End-Permian Mass Extinction. Lethaia, 40(2): Brayard, A., Escarguel, G., Bucher, H., 2007b. The Biogeography of Early Triassic Ammonoid Faunas: Clusters, Gradients, and Networks. Geobios, 40(6): Brayard, A., Bucher, H., Escarguel, G., et al., 2006a. The Early Triassic Ammonoid Recovery: Palaeoclimatic Significance of Diversity Gradients. Palaeogeography, Palaeoclimatology, Palaeoecology, 239(3 4): Brayard, A., Escarguel, G., Bucher, H., 2006b. The Biogeography of Early Triassic Ammonoid Faunas: Clusters, Gradients, and Networks. Geobios, 40: Brayard, A., Escarguel, G., Bucher, H., et al., 2009a. Smithian and Spathian (Early Triassic) Ammonoid Assemblages from Terranes: Paleoceanographic and Paleogeographic Implica-

14 14 Yuri D Zakharov and Alexander M Popov tions. Journal of Asian Earth Sciences, 36(6): Brayard, A., Escarguel, G., Bucher, H., et al., 2009b. Good Genes and Good Luck: Ammonoid Diversity and the End-Permian Mass-Extinction. Science, 325(5944): Brayard, A., Brühwiler, T., Bucher, H., et al., 2009c. Guodunites, a Low-Palaeolatitude and Trans-Panthalassic Smithian (Early Triassic) Ammonoid Genus. Palaeontology, 52(2): Brühwiler, T., Brayard, A., Bucher, H., et al., Griesbachian and Dienerian (Early Triassic) Ammonoid Faunas from Northwestern Guangxi and Southern Guizhou (South China). Palaeontology, 51(5): Brühwiler, T., Bucher, H., Goudemand, N., 2010a. Smithian (Early Triassic) Ammonoids from Tulong, South Tibet. Geobios, 43(4): Brühwiler, T., Ware, D., Bucher, H., et al., 2010b. New Early Triassic Ammonoid Faunas from the Dienerian/Smithian Boundary Beds at the Induan/Olenekian GSSP Candidate at Mud (Spiti, Northern India). Journal of Asian Earth Sciences, 39(6): Brühwiler, T., Bucher, H., Brayard, A., et al., 2010c. High- Resolution Biochronology and Diversity Dynamics of the Early Triassic Ammonoid Recovery: The Smithian Faunas of the Northern Indian Margin. Palaeogeography, Palaeoclimatology, Palaeoecology, 297(2): Brühwiler, T., Bucher, H., Krystin, L., 2012a. Middle and Late Smithian (Early Triassic) Ammonoids from Spiti (India). Special Papers in Palaeontology, 88: Brühwiler, T., Bucher, H., Ware, D., et al., 2012b. Smithian (Early Triassic) Ammonoids from the Salt Range, Pakistan. Special Papers in Palaeontology, 88: Chen, Y. M., New Advance in the Study of Triassic Brachiopods in Tulong District of Nyalam Country, Tibet. Contribution to the Geology of the Qinghai-Xizang Plateau 11. Geological Publishing House, Beijing Chen, Z. Q., Kaiho, K., George, A. D., 2005a. Survival Strategies of Brachiopod Faunas from the End-Permian Mass Extinction. Palaeogeography, Palaeoclimatology, Palaeoecology, 224(1 3): Chen, Z. Q., Kaiho, K., George, A. D., 2005b. Early Triassic Recovery of the Brachiopod Faunas from the End-Permian Mass Extinction: A Global Review. Palaeogeography, Palaeoclimatology, Palaeoecology, 224(1 3): Chen, Z. Q., Kaiho, K., George, A. D., et al., Survival Brachiopod Faunas of the End-Permian Mass Extinction from the Southern Alps (Italy) and South China. Geological Magazine, 143(3): Dagys, A. S., Triassic Brachiopods (Morphology, Classification, Phylogeny, Stratigraphical Significance and Biogeography). Trudy Instituta Geologii i Geophiziki Sibirskogo Otdeleniya Akademii Nauk SSSR, 214: (in Russian) Dagys, A. S., Geographical Differentiation of Triassic Brachiopods. Palaeogeography, Palaeoclimatology, Palaeoecology, 100(1 2): Dagys, A. S., Ermakova, S. P., Boreal nye Pozdneolenekskie Ammonoidei (Boreal Late Olenekian Ammonoids). Trudy Instituta Geologii i Geofiziki Sibirskogo Otdelenija Akademii nauk SSSR, 714: (in Russian) Dagys, A., Ermakova, S., Induan (Triassic) Ammonoids from North-Eastern Asia. Revue de Paléobiologie, 15: Ermakova, S. P., Zonal Standard of the Boreal Lower Triassic. Nauka, Moscow, 110 (in Russian) Erwin, D. H., The Permian-Triassic Extinction. Nature, 367: Galfetti, T., Bucher, H., Brayard, A., et al., Late Early Triassic Climate Change: Insights from Carbonate Carbon Isotopes, Sedimentary Evolution and Ammonoid Paleobiogeography. Palaeogeography, Palaeoclimatology, Palaeoecology, 243(3 4): Gavrilova, V. A., The Upper Olenekian of Gornyj Mangyshlak (Stratigraphy, Correlation, Ammonoids). Vestnik Sankt-Peterburgskogo Universiteta, 7th Seriya (in Russian) Girty, G. H., New Species of Lower Triassic Fossils from the Woodside and Thaynes Formations. In: Mansfield, G. R., ed., Geography, Geology, and Mineral Resources of Part of Southeastern Idaho, with Descriptions of Carboniferous and Triassic Fossils by G. H. Girty. United States Geological Survey Professional Paper, 152: Grant, R. E., Brachiopods from Permian-Triassic Boundary Beds and Age of Chhidru Formation, West Pakistan. In: Teichert, C., Kummel, B., eds., Stratigraphic Boundary Problems: Permian and Triassic of West Pakistan 4. Kansas University Press Special Publication, Kansas Guex, J., Le Trias Inférieur des Salt Range (Pakistan): Problemes Biochronologiques (Lower Triassic of the Salt Range (Pakistan): Biochronological Problems). Eclogae geologicae Helvetia, 71: Guex, J., Jenks, J. K., O Dogherty, L., et al., Spathian (Lower Triassic) Ammonoids from Western USA (Idaho, California, Utah, and Nevada). Mémoire de Géologie (Lausanne), 49: 1 82 Harries, P. J., Knorr, P. O., What does the Lilliput Effect Means? Palaeogeography, Palaeoclimatology, Palaeoecology, 284(1 2): 4 10 Hermann, E., Hochuli, P. A., Méhay, S., et al., Organic Matter and Palaeoenvironmental Signals during the Early Triassic Biotic Recovery: The Salt Range and Surghar Range Records. Sedimentary Geology, 234(1 4): Hoover, P. R., Early Triassic Terebratulid Brachiopods from the Western Interior of the United States. United States Geological Survey Professional Paper, 1057: 1 21 Isozaki, Y., Permo-Triassic Boundary Superanoxia and Stratified Superocean: Records from Lost Deep Sea. Science, 276(5310): , doi: /science Jordan, M., Triassic Brachiopods of Romania. In: Palfy, J., Jordan, M., eds., Mesozoic Brachiopods of Alpine Europe. Hungarian Geological Society, Budapest Keller, G., Abramovich, S., Lilliput Effect in Late Maastrichtian Planktic Foraminifera: Response to Environmental Stress. Palaeogeography, Palaeoclimatology, Palaeoecology, 284(1 2): Kidder, D. L., Worsley, T. R., Causes and Consequences of Extreme Permo-Triassic Warming to Globally Equable

15 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 15 Climate and Relation to the Permo-Triassic Extinction and Recovery. Palaeogeography, Palaeoclimatology, Palaeoecology, 203(3 4): Knoll, A., Bambach, R. K., Payne, L., et al., Paleophysiology and End-Permian Mass Extinction. Earth and Planetary Science Letters, 256(3 4): Kozur, H. W., Some Aspects of the Permian-Triassic Boundary (PTB) and the Possible Causes for the Biotic Crisis around This Boundary. Palaeogeography, Palaeoclimatology, Palaeoecology, 143(4): Kozur, H. W., Biostratigraphy and Event Stratigraphy in Iran around the Permian-Triassic Boundary (PTB): Implications for the Causes of the PTB Biotic Crisis. Global and Planetary Change, 55(1 3): Krystyn, L., Orchard, M. J., Lowermost Triassic Ammonoid and Conodont Biostratigraphy of Spiti, India. Albertiana, 17: Krystyn, L., Richoz, S., Baud, A., et al., A Unique Permian-Triassic Boundary Section from the Neotethyan Hawasina Basin, Central Oman Mountain. Palaeogeography, Palaeoclimatology, Palaeoecology, 191(3 4): MacFarlan, D. A. B., Triassic & Jurassic Rhynchonellacea (Brachiopoda) from New Zealand & New Caledonia. The Royal Society of New Zealand, Bulletin, 31: McGovan, A. J., Ammonoid Recovery from the Late Permian Mass Extinction Event. Comptes Rendus Palevol, 4(6 7): McGovan, A. J., Smith, A. B., Ammonoids across the Permian/Triassic Boundary: A Cladistic Perspective. Palaeontology, 50(3): Moriya, K., Nishi, H., Kawahata, H., et al., Demersal Habitat of Late Cretaceous Ammonoids: Evidence from Oxygen Isotopes for the Campanian (Late Cretaceous) Northwestern Pacific Thermal Structure. Geology, 31(2): Mu, L., Zakharov, Y. D., Shen, S. Z., Early Induan (Early Triassic) Cephalopods from the Daye Formation at Guiding, Guizhou Province, South China. Journal of Paleontology, 81(5): Orchard, M., Krystyn, L., Conodonts of the Lowermost Triassic of Spiti, a New Zonation Based on Neogondolellasuccessions. Rivista Italiana di Paleontologia e Stratigrafia, 104(3): Payne, J. L., Kump, L. R., Evidence for Recurrent Early Triassic Massive Volcanism from Quantitative Interpretation of Carbon Isotope Fluctuations. Earth and Planetary Science Letters, 256(1 2): Perry, D. G., Chatterton, B. D. E., Late Early Triassic Brachiopod and Conodont Fauna. Thaynes Formation, Southeastern Idaho. Journal of Paleontology, 53(2): Popow, Y. N., Triassic Ammonoids of the North-East USSR. Trudy Nauchno-Issledovatelskogo Instituta Geologii Arktiki Ministerstva Geologii i Okhrany Nedr SSSR, 79: (in Russian) Ruban, D. A., Phanerozoic Changes in the High-Rank Suprageneric Diversity Structure of Brachiopods: Linear and Non-Linear Effects. Palaeoworld, 18(4): Shen, S. Z., He, X. L., The Changhsingian Brachiopod Fauna from Guiding, Guizhou. Acta Palaeontologica Sinica, 33(4): (in Chinese with English Abstract) Shen, S. Z., Shi, G. R., Diversity and Extinction Patterns of Permian Brachiopoda of South China. Historical Biology, 12(2): Shen, S. Z., Zhang, H., Li, W. Z., et al., Brachiopod Diversity Pattern from Carboniferous to Triassic in South China. Geological Journal, 41(3): Shevyrev, A. A., Triassic Ammonoids of the South USSR. Trudy Paleontologicheskogo Instituta Akademii nauk SSSR, 110: (in Russian) Shevyrev, A. A., Triassic Ammonoids. Trudy Paleontologicheskogo Instituta Akademii nauk SSSR, 217: (in Russian) Shevyrev, A. A., Triassic Biochronology: State of the Art and Main Problems. Stratigraphy and Geological Correlation, 14(6): Shigeta, Y., Zakharov, Y. D., Maeda, H., et al., The Lower Triassic System in the Abrek Bay Area, South Primorye, Russia. National Museum of Nature and Science Monographs No. 38. National Museum of Nature and Science, Tokyo. 218 Shimizu, D., Upper Permian Brachiopod Fossils from Guryul Ravine and the Spur Three Kilometers North of Barus. In: Nakazawa, K., Kapoor, H. M., eds., The Upper Permian and Lower Triassic faunas of Kashmir. Palaeontologica Indica, 46: Smyshlyaeva, O. P., Zakharov, Y. D., Shigeta, Y., et al., Optimal Temperatures of Growth in Campanian Ammonoids of Sakhalin (Krilyon) and Hokkaido: Oxygen and Carbon Isotopic Data. The IV International Symposium of International Geological Correlation Program Project 434. Cretaceous Continental Margin of East Asia: Stratigraphy, Sedimentation, and Tectonics. Program and Abstracts, Khabarovsk Smyshlyaeva, O. P., Zakharov, Y. D., New Representatives of the Family Melagathiceratidae (Ammonoidea) from the Lower Triassic of South Primorye. Paleontologicheskii Zhurnal, 2: (in Russian) Stemmerik, L., Bendix-Almgreen, S. E., Plasecki, S., The Permian-Triassic Boundary in Central East Greenland: Past and Present Views. Bulletin of the Geological Society of Denmark, 48(2): Tozer, E. T., Canadian Triassic Ammonoid Fauna. Geological Survey of Canada, Bulletin, 467: Twitchett, R. J., Krystyn, L., Baud, A., et al., Rapid Marine Recovery after the End-Permian Mass-Extinction Event in the Absence of Marine Anoxia. Geology, 32(9): Ware, D., Jenks, J. F., Hautmann, M., Dienerian (Early Triassic) Ammonoids from the Candelaria Hills (Nevada, USA) and Their Significance for Palaeobiogeography and Palaeocenography. Swiss Journal of Geoscience, 104(1): , doi: /s Waterhouse, J. B., Permian Brachiopoda and Mollusca from Nepal. Palaeontographica, Abteilung A, 160(1 6): 1 175

16 16 Yuri D Zakharov and Alexander M Popov Waterhouse, J. B., The Early and Middle Triassic Ammonoid Succession of the Himalayas in Western and Central Nepal. Part 2. Systematic Studies of the Early Middle Scythian. Palaeontographica, Abteilung A, 241(1 3): Wignall, P. B., Hallam, A., Griesbachian (Earliest Triassic) Paleoenvironmental Changes in the Salt Range, Pakistan and Southeast China and Their Bearing on the Permo- Triassic Mass Extinction. Palaeogeography, Palaeoclimatology, Palaeoecology, 101(3 4): Williams, A., James, M. A., Emig, C. C., et al., Anatomy. In: Kaesler, R. L., ed., Treatise on Invertebrate Paleontology. Part H, Brachiopoda, Revised, Vol. 1. Introduction. Geological Society of America & University of Kansas Press, Boulder, Colorado & Lawrence, Kansas Xu, G. R., Liu, G. C., Brachiopods. In: Yang, Z. Y., Yin, H. F., Xu, G., eds., Triassic of the South Qilian Mountains. Geological Publishing House, Beijing (in Chinese with English Abstract) Xu, G. R., Grant, R. E., Brachiopods near the Permian- Triassic Boundary in South China. Smithsonian Contributions to Paleobiology, 76: 1 68 Yang, Z. I., Yang, F., Wu, S., The Ammonoid Hypophiceras Fauna near the Permian-Triassic Boundary at Meishan Section and in South China: Stratigraphical Significance. In: Yin, H. F., ed., The Palaeozoic-Mesozoic Boundary. Candidates of the Global Stratotype Section and Point of the Permian-Triassic Boundary. China University of Geosciences Press, Wuhan Yin, H. F., Wu, S. B., Ding, M. H., et al., The Meishan Section, Candidate of the Global Stratotype Section and Point of Permian-Triassic Boundary. In: Yin, H. F., ed., The Palaeozoic-Mesozoic Boundary. Candidates of the Global Stratotype Section and Point of the Permian-Triassic Boundary. China University of Geosciences Press, Wuhan Yin, H. F., Zhang, K. X., Eventostratigraphy of the Permian-Triassic Boundary at Meishan Section, South China. In: Yin, H. F., ed., The Palaeozoic-Mesozoic Boundary. Candidates of the Global Stratotype Section and Point of the Permian-Triassic Boundary. China University of Geosciences Press, Wuhan Zakharov, Y. D., Biostratigraphy and Ammonoids of the Lower Triassic of South Primorye. Nauka, Moscow. 175 (in Russian) Zakharov, Y. D., Lower Triassic Ammonoids of the USSR. Nauka, Moscow. 224 (in Russian) Zakharov, Y. D., Ammonoid Evolution and the Problem of the Stage and Substage Division of the Lower Triassic. In: Baud, A., Popova, I., Dickins, J. M., eds., Late Paleozoic and Early Mesozoic Circum-Pacific Events: Biostratigraphy, Tectonic and Ore Deposits of Primorye (Far East Russia). Mémoires de Géologie (Lausanne) 30: Zakharov, Y. D., The Induan-Olenekian Boundary in the Tethys and Boreal Realm. Annali dei Musei Civici di Rovereto, Sezione: Archeologia, Storia e Scienze Naturali, 11: Zakharov, Y. D., Popov, A. M., Biakov, A. S., Late Permian to Middle Triassic Palaeogeographic Differentiation of Key Ammonoid Groups: Evidence from the Former USSR. Polar Research, 27(3): Zakharov, Y. D., Popov, A. M., Buryi, G. I., Unique Marine Olenekian-Anisian Boundary Section from South Primorye, Russian Far East. Journal of China University of Geosciences, 16(3): Zakharov, Y. D., Smyshlyaeva, O. P., Safronov, P. P., et al., Triassic Ammonoid Succession in South Primorye: 5. Stratigraphical Position of the Early Olenekian Meekoceras fauna. Albertiana, 38: 23 33

17 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 17 APPENDIX: Original Data on Brachiopod and Ammonoid Remains from the Lower Triassic of South Primorye and Mangyshlak 1 SOUTH PRIMORYE The first geological studies in Russian southern Far East (Ussuri region, later named as Primorye) were made by V. P. Margaritov, a graduate from the St.-Peterburg University, arriving in Vladivostok in 1880 as a teacher of math. On the western coast of the Ussuri Gulf, near Shamara Bay (now Lazurnaya) he discovered some fossils, including ammonoids and inarticulated brachiopods. His collection fell into the hands of the President of the Russian Academy of Sciences, A. P. Karpinsky, a recognized expert in Late Palaeozoic ammonoids, who distinguished Early Triassic ceratitid ammonoids among collected fossils. Later, Early and Middle Triassic marine deposits in the environs of Vladivostok were studied by D. L. Ivanov, the chief of a geological team making reconnaissance work for the construction of the Trans-Siberian railroad. He collected mollusc remains in the area of the Shamara Bay and articulated brachiopod and mollusc shells on Russian Island. On the initiative of A. P. Karpinsky, collections V. P. Margaritov and D. L. Ivanov were forwarded to Austrian palaeontologists K. Diener and A. Bittner, who identified and described them (Bittner, 1899b; Diener, 1895). Later, Triassic cephalopods were investigated by Kiparisova (1961), Burij and Zharnikova (1981, 1962), Zakharov (1997a, 1978, 1968), Zharnikova (1985, 1972), Shigeta et al. (2009), Smyshlyaeva and Zakharov (2011), and Smyshlyaeva (2010). However, after Bittner s investigation, only a few Early Triassic brachiopod taxa, based on material from South Primorye, was described by Dagys (1974, 1965) and Shigeta et al. (2009). Observed ranges of ammonoids and brachiopods across the Induan-Lower Anisian interval in South Primorye (Seryj-Tri Kamnya, Abrek, Konechnyj, Tobizin, Ayax-Balka, Zhitkov, Golyj, and Paris), summarized in Figs. S1 S9 and Table S1, help to understand the main features of their recovery on specific level from the End-Permian extinction. 1.1 Seryj-Tri Kamnya Capes The Seryj Cape-Tri Kamnya Cape Section, exposed at the western Ussuri Gulf, is represented by the Lazurnaya and Tobizin formations (Zakharov et al., 2010; Markevich and Zakharov, 2004; Zakharov, 1997b, 1996, 1968; Kiparisova, 1972, 1961; Burij, 1959; Korzh, 1959; Diener, 1895) Brachiopod occurrences Lissorhynchia sp. (Pontiidae), the first record of this genus from South Primorye, seems to be the oldest Induan articulated brachiopod species found in the region. A single representative of this species was collected by T. Kumagae in the lower part of the Lazurnaya Bay Formation (Gyronites subdharmus Zone, about 20 m above the top of the underlying early Induan Tompophiceras ussuriense Zone). Other Induan brachiopod fossils from this section are represented by Lingula borealis Bittner. A few articulated brachiopods, mainly rhynchonellids, were collected in the lower part of the Smithian Tobizin Formation (Euflemingites prynadai beds) (Fig. S1) Ammonoid occurrences Early Triassic ammonoids are significantly more diverse in the mentioned section, as well as in other South Primorye sections, than associated brachiopods. Induan ammonoids from the Lazurnaya Bay Formation, exposed on the western coast of Ussuri Gulf, are represented by a single genus (Tompophiceras), belonging to the Palaeozoic-type family Dzhulfitidae, and five genera within the Mesozoic-type families Ophiceratidae, Proptychitidae, Meekoceratidae, and Xenoceltitidae?. Early Olenekian (Smithian) ammonoids from the uppermost portion of the Lazurnaya Bay Formation (Ussuriflemingites abrekensis (= Gyronites separatus ) beds of the Hedenstroemia bosphorensis Zone) and the lower portion of the Tobizin Cape Formation (Euflemingites prynadai beds of the mentioned zone) in this section are more diverse, representing by 22 genera within Mesozoic-type families Sageceratidae, Hedenstroemiidae, Aspenitidae, Ussuriidae, Paranannitidae, Clypeoceratidae, Arctoceratidae, Meekoceratidae, Dieneroceratidae, and Flemingitidae (Fig. S1). 1.2 Abrek Bay The Abrek Bay Section, located at Strelok Strait and represented by the Lazurnaya Bay and Zhitkov Cape formations, is worthy of consideration as one of the reference sections for Lower Triassic stratigraphy in South Primorye (Shigeta et al., 2009; Markevich and Zakharov, 2004; Zakharov et al., 2002; Zakharov and Popov, 1999; Kiparisova, 1972, 1961; Burij, 1959) Brachiopod occurrences In the lower part of the Induan Lazurnaya Bay Formation of the Abrek Bay Section only inarticulated brachiopods (Lingula borealis Bittner and Orbiculoidea sp.) have been discovered (Shigeta et al., 2009). Among late Induan brachiopods from the Lytophiceras-bearing beds of the Gyronites subdharmus Zone about five genera of articulated brachiopods, including Abrekia, have been recognized (Shigeta et al., 2009; Dagys, 1974). All of them belong to Mesozoic-type families (Wellerellidae, Pontisiidae, Rhynchonellidae, and Norellidae). Representatives of the Abrekia (A. sulcata Dagys) have been discovered also in the Lower Olenekian (Smithian) portion of the Zhitkov Cape Formation ( Hedenstroemia bosphorensis Zone, presumably Euflemingites prynadai beds) (Fig. S2) Ammonoid occurrences The Induan ammonoid succession in the Abrek Bay Section consists of a single genus (Tompophiceras), belonging to a Permian-type family Dhulfitidae, and 12 genera within five Mesozoic-type families (Proptychitidae, Ophiceratidae, Meekoceratidae, Paranoritidae, and?paranannitidae). Among Smithian ammonoids of the mentioned section 27 genera belonging to 15 Mesozoic-type families have been recognized (Fig. S2). They are following: Sageceratidae, Paranannitidae, Aspenitidae, Xenoceltitidae, Proptychitidae?, Clypeoceratidae, Arctoceratidae, Owenitidae, Inyoitidae?, Prionitidae, Kashmiritidae, Meekoceratidae, Dieneroceratidae, Flemingitidae, and Palaeophyllitidae.

18 18 Yuri D Zakharov and Alexander M Popov 1.3 Konechnyj Cape The Konechnyj Cape, located at Novik Bay on Russian Island, is represented by the Schmidt Formation (Markevich and Zakharov, 2004). At Konechnuj Cape, only a small portion, about 6 m thick, of the middle Olenekian (early Smithian) Schmidt Cape Formation is Figure S1. Observed stratigraphic distribution of Early Triassic brachiopods and ammonoids in the Seryj-Tri Kamnya Capes Section, South Primorye.

19 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 19

20 20 Yuri D Zakharov and Alexander M Popov Figure S3. Observed stratigraphic distribution of Olenekian brachiopods and ammonoids in the Tobizin Cape Section, South Primorye. T.. Tompophiceras ussuriensis; L.. Lazurnaya Bay; Hedenstr.. Hedenstroemia. Other designations as in Fig. S1. exposed. No ammonoids, but abundant of articulated brachiopods were recently collected there. The two brachiopod assemblages have been recognized in the section: (1) Hustedtiella planicosta- Spiriferinida-Rhynchonellida- Fletcherithyris margaritovi- Zeilleriidae? below (occurs mainly in blocks) and (2) monospecies- Fletcherithyris margaritovi assemblage above. Both the assemblages correspond to the Tirolites ussuriensis beds of the early Spathian Tirolites-Amphistephanites Zone in other sections at the Russian Island (Markevich and Zakharov, 2004). Representatives of five genera within five Mesozoic-type families (Rhynchonellidae, Neoretziidae, Spiriferinidae, Dielasmatidae, and Zeilleriidae?) were discovered in this locality. 1.4 Tobizin Cape The Tobizin Cape Formation, exposed on the south part of Russian Island (Novyj Dzhigit Bay area) is represented by the Lazurnaya Bay (basal part), Tobizin Cape and Schmidt Cape formations (Markevich and Zakharov, 2004; Buryi, 1979; Zakharov, 1968; Burij, 1959; Korzh, 1959; Diener, 1895) Brachiopod occurrences From the early Olenekian (Smithian) Tobizin Cape Formation only inarticulated brachiopod Lingula borealis Bittner is known. The articulated brachiopods (Zeilleriidae? gen. and sp. nov. E and Spiriferinidae gen. and sp. indet.) (Fig. S3) have been collected only from the upper part of the middle Olenekian (early Spathian) Schmidt Cape Formation Ammonoid occurrences Only early to middle Olenekian (Smithian-early Spathian) ammonoid succession, represented by 20 genera belonging to 14 Mesozoic-type families is known there. These families are following: Aspenitidae, Ussuriidae, Paranannitidae, Owenitidae, Meekoceratidae, Dieneroceratidae, Prionitidae, Tirolitidae, Melagathiceratidae, Stephanitidae, Dinaritidae, Clypeoceratidae, Arctoceratidae, and Flemingitidae) (Fig. S3). 1.5 Ayax Bay-Balka Cape The Ayax Bay-Balka Cape, exposed on the northern part of Russian Island, consists of Lazurnaya Bay, Tobizin Cape and Schmidt Cape formations (Markevich and Zakharov, 2004; Zakharov, 1997a, 1978, 1968; Buryi, 1979; Kiparisova, 1972, 1961; Korzh, 1959; Diener, 1895) Brachiopod occurrences Only inarticulated brachiopod Lingula borealis Bittner

21 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 21 Figure S4. Observed stratigraphic distribution of Early Triassic brachiopods and ammonoids in the Ayax Bay-Balka Cape area, South Primorye. Reg. substage. regional substage. Other designations as in Figs. S1 and S3. and fragments of rhynchonellid and terebratulid brachiopod shells were collected from the lower part (Induan-Smithian) of the section, which consists of the Lazurnaya Bay and Tobizin Cape formations. Articulated brachiopods Lepismatina aff. mansfieldi (Girty) and spiriferinids belonging to Mesozoictype families Lepismatinidae and Spiriferinidae have been reported from the lower part of the middle Olenekian (early Spathian) Schmidt Cape Formation (Fig. S4) Ammonoid occurrences At the top of the Induan conglomerate member of the Lazurnaya Cape Formation only bad preserved Gyronites? sp. (Meekoceratidae) has been recognized. Among early to middle Olenekian ammonoids from the uppermost portion of the Lazurnaya Bay Formation, Tobizin Cape and Schmidt Cape formations 22 ammonoid genera belonging to 17 Mesozoictype families have been listed (Zakharov, 1997a, b). Among them are following: Sageceratidae, Hedenstroemiidae, Ussuriidae, Paranannitidae, Arctoceratidae, Meekoceratidae, Dieneroceratidae, Owenitidae, Inyotidae, Prionitidae, Xenoceltitidae, Tirolitidae, Melagathiceratidae, Stephanitidae, Doricranitidae?, Dinaritidae, and Palaeophyllitidae (Fig. S4).

22 22 Yuri D Zakharov and Alexander M Popov

23 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 23 Figure S6. Observed stratigraphic distribution of Olenekian and early Anisian brachiopods and ammonoids in the Zhitkov Cape Section, South Primorye. Designations as in Figs. S1 and S Schmidt Cape-Tchernyschev Bay The Schmidt Cape-Tchernyschev Bay, located on the south-eastern part of Russian Island, is represented by the Schmidt Cape and Zhitkov Cape formations (Markevich and Zakharov, 2004; Buryi, 1979; Zakharov, 1978, 1968; Kiparisova, 1972, 1961; Burij, 1959; Korzh, 1959) Brachiopod occurrences Schmidt Cape area is one of the places with abundantearly Spathian Mesozoic-type articulated brachiopods (mainly in the Tirolites ussuriensis beds of the Tirolites-Amphistephanites Zone) (Fig. S5). In this part of the mentioned section, there are five genera belonging to the following Mesozoic-type families: Spiriferinidae, Neoretziidae, Dielasmatidae, and Zeilleriidae (?). Earliest Anisian spiriferinid and terebratulid brachiopods are reported from the Ussuriphyllites amurensis Zone of the Tchernyschev Bay area (Fig. S5) Ammonoid occurrences Spathian ammonoids from the Schmidt and Zhitkov Cape formations of this section are represented by 21 genera belonging to 12 Mesozoic-type families (Sageceratidae, Paranannitidae, Khvalynitidae, Meekoceratidae, Prionitidae, Columbitidae, Kazakhstanitidae, Xenoceltitidae, Tirolitidae, Dinaritidae, and Palaeophyllitidae) (Fig. S5).

24 24 Yuri D Zakharov and Alexander M Popov Figure S7. Observed stratigraphic distribution of Early Triassic and early Anisian invertebrates in the Golyj (Kom-Pikho- Sakho) Cape Section, South Primorye. Heden. bosph.. Hedenstroemia bosphorensis; Neocol.. Neocolumbites;?S.m..?Subfengshanites multiformis; reg. substage. regional substage. Other designations as in Figs. S1 and S5.

25 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 25 Figure S8. Observed stratigraphic distribution of Olenekian brachiopods and ammonoids in the Paris Bay Section, South Primorye. Designations as in Figs. S1 and S7. Sub. mul.. Subfengshanites multiformis; Leioph. prad.. Leiophyllites pradyumna; reg. substage. regional substage. 1.7 Zhitkov Cape The Zhitkov Cape Section, exposed on the northern part of Russian Island, is represented by the Tobizin Cape, Zhitkov Cape and Karazin Cape formations (Markevich and Zakharov, 2004; Zakharov, 1997a, 1978, 1968; Buryi, 1979; Kiparisova, 1972, 1961; Burij, 1959; Korzh, 1959) Brachiopod occurrences Majority of articulated brachiopods from this section was collected in the Tirolites ussuriensis beds of the early Spathian Tirolites-Amphistephanites Zone (Schmidt Formation), others were collected in the late Spathian Subfengshanites multiformis Zone (Fig. S6). Olenekian portion of the Zhitkov Cape Section includes seven genera belonging to six Mesozoic-type families (Rhynchonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae, Dielasmatidae, and Zeilleriidae?) Ammonoid occurrences Smithian and Spathian ammonods of this section were collected in the upper portion of the Tobizin Cape Formation (Anasibirites nevolini Zone) and Schmidt Cape (Tirolites- Amphistephanites Zone), and Zhitkov Cape (Neocolumbites insignis and Subfengshanites multiformis zones) formations (Fig. S6). They are represented by 24 genera belonging to 13 families (Sageceratidae, Paranannitidae, Khvalynitidae, Isculitidae, Meekoceratidae, Dieneroceratidae, Columbitidae, Kazakhstanitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Dinaritidae, and Megaphyllitidae). 1.8 Golyj (Kom-Pikho-Sakho) Cape The Golyj Cape Section, exposed at eastern part of the Ussuri Gulf, consists of Lazurnaya Bay, Zhitkov Cape and Karazin formations (Markevich and Zakharov, 2004; Buryi, 1979; Zakharov, 1968; Burij, 1959; Korzh, 1959) Brachiopod occurrences Articulated brachiopods in this section were discovered only in the middle Olenekian (early Spathian) portion of the Zhitkov Cape Formation (Tirolites-Amphistephanites Zone). They are represented by five genera of four Mesozoic-type families (Rhynchonellidae, Neoretziidae, Spiriferinidae, and Zeilleriidae?) (Fig. S7) Ammonoid occurrences Representatives of a single genus Gyronites (Mesozoictype family Meekoceratidae) were reported from the Induan

26 26 Yuri D Zakharov and Alexander M Popov Lazurnaya Bay Formation (Fig. S7). Early to late Olenekian ammonoids were discovered from the lower ( Hedenstroemia bosphorensis Zone), middle (Tirolites-Amphistephanites Zone) and upper (Neocolumbites insignis Zone) portions of the Zhitkov Cape Formation. Olenekian ammonoid assemblage of the Golyj Cape section consists of 24 genera belonging to 17 Mesozoic-type families: Sageceratidae, Hedenstroemiidae, Aspenitidae, Ussuriidae, Paranannitidae, Khvalynitidae, Arctoceratidae, Owenitidae, Meekoceratidae, Inyoitidae, Prionitidae, Columbitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Melagathiceratidae, and Flemingitidae. 1.9 Paris Bay The Paris Bay Section, located on the northern part of Russian Island, is represented by the Tobizin Cape, Schmidt Cape, Zhitkov Cape and Karazin Cape formations (Markevich and Zakharov, 2004; Buryi, 1979; Zakharov, 1968; Burij, 1959; Korzh, 1959) Brachiopod occurrences Articulated brachiopods from the Paris Bay Section were collected from the early Spathian Schmidt Formation (Tirolites-Amphistephanites Zone) and uppermost beds of the late Spathian Zhitkov Formation (Subfengshanites multiformis Zone). Spathian assemblage of articulated brachiopods consists of six genera belonging to five Mesozoic-type families: Rhynchonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae, and Zeilleriidae? (Fig. S8) Ammonoid occurrences Ammonoids from the Paris Bay Section are originated from the upper portion of the Smithian Tobizin Formation (the upper part of the Anasibirites nevolini Zone), early Spathian Schmidt Formation (Tirolites-Amphistephanites Zone), and late Spathian Zhitkov Formation (Neocolumbites insignis and Subfengshanites multiformis zones). Olenekian (mainly Spathian) ammonoids from this section are represented by 18 genera within 12 Mesozoic-type families: Khvalynitidae,?Proptychitidae Figure S9. Observed stratigraphic distribution of Olenekian brachiopods and ammonoids in the Dolnapa Well Section, Mangyshlak. A. Anisian; Tir. cassianus-kip. carinatus. Tirolites cassianus-kiparisovites carinatus; Col.. Columbites; Ar.b.-S.u.. Arnautoceltites bajarunasi-stacheites undatus; Eum.. Eumorphotis; K. Karadunskaya. Other designations as in Fig. S1.

27 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 27 Figure S10. Geographical differentiation of the recovery brachiopod faunas in the Induan (base map after Ziegler et al., 1998). Realms: I. Boreal; II. Tethyan Palaeoequatorial; III. American Palaeoequatorial; IV. Gondwanan (Zakharov et al., 2008). The arrows represent inferred major ocean currents. Localities: 1. Primorye (Dagys, 1974; this study); 2. Idaho (Girty, 1927); 3. South China (Chen et al., 2005, 2000; Chen and Shi, 1999; Shen and He, 1994; Xu and Grant, 1994; Liao, 1987, 1984, 1980, 1979; Sheng et al., 1984); 4. North Caucasus (Dagys, 1974); 5. Salt Range (Grant, 1970; Kummel and Teichert, 1970); 6. Nepal (Waterhouse, 1994, 1978; Waterhouse and Shi, 1991). Figure S11. Geographical differentiation of the recovery brachiopod faunas in the Olenekian (base map after Ziegler et al., 1998). Designations are as listed in Fig. S10. Localities: 1. Spitzbergen (Dagys, 1993, 1974); 2. Primorye (Dagys, 1974; this study); 3. Japan (Dagys, 1974); 4. Idaho (Hoover, 1979; Perry and Chatterton, 1979; Girty, 1927); 5. Carpatian-Balkanian area (Jordan, 1993); 8. South China (e.g., Chen et al., 2005; Xu and Liu, 1983); 9. Tibet (Chen, 1983; Sun et al., 1981); 10. Himalayas (Bittner, 1899a); 11. New Zealand (MacFarlan, 1992).

28 28 Yuri D Zakharov and Alexander M Popov Figure S12. Geographical differentiation of the recovery ammonoid faunas in the earliest Induan (base map after Ziegler et al., 1998). Designations are as listed in Fig. S10. Localities (Zakharov et al., 2008; Brayard et al., 2006a, b): 1. Verkhoyansk area; 2. Svalbard; 3. Arctic Canada; 4. Alaska; 5. Greenland; 6. Primorye; 7. South China; 8. Transcaucasia; 9. Iran; 10. Kashmir; 11. southern Tibet; 12. Himalayas; 13. Oman. (Proptychitoides), Arctoceratidae, Meekoceratidae, Dieneroceratidae, Columbitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Stephanitidae, Dinaritidae, and Hungaritidae (Fig. S8). 2 MANGYSHLAK, DOLNAPA (KAZAKHSTAN) The first Triassic fossils in Mangyshlak were found in 1914 by M. V. Bayarunas, who identified and described some ammonoids among them 22 years later (Bajarunas, 1936). Later, Early Triassic of Mangyshlak were investigated by Astachova (1960), Shevyrev (1968), Gavrilova (2007, 1989, 1980), Balini et al. (2000) and Zakharov et al. (2008). Before our investigation, Triassic brachiopods from Mangyshlak were described by Dagys (1974) on the bases of A. A Shevyrev s collection. The reference section for the Lower Triassic in Mangyshlak, located at the Dolnapa Well area in Kara-Tau, is represented by the Tartalinskaya, Karadzhatykskaya and Karaduanskaya formations. 2.1 Brachiopod Occurrences Spathian brachiopod assemblage of the mentioned section consists of 8 11 genera belonging to eight Mesozoic-type families (Pontisiidae, Norellidae, Diplospirellidae, Neoretziidae, Lepismatinidae, Dielasmatidae, Antezeilleridae, and Plectoconchidae) (Fig. S9). 2.2 Ammonoid Occurrences Spathian ammonoids are represented by 23 genera belonging to 14 Mesozoic-type families (Sageceratidae, Khvalynitidae, Dinaritidae, Prionitidae, Columbitidae, Kazakhstanitidae, Kashmiritidae, Xenoceltitidae, Olenikitidae, Tirolitidae, Doricranitidae, Dinaritidae, Palaeophyllitidae, and Megaphyllitidae (Fig. S9). 3 DISCUSSION The Induan of South Primorye (SP) is characterised by seven articulated brachiopod species: early Induan Lissorhynchia sp. and six late Induan species, including Abrekia sulcata and new ones (Figs. S1 and S2 and Table S1). Early Olenekian brachiopods from SP show a slight reduction in species number, largely due to a variety of facial conditions. Middle Olenekian brachiopods from SP (about eight species, including new ones) (Figs. S3 S9 and Table S1) are more diverse than those of Mangyshlak. In addition, information on late Olenekian brachiopods from SP (about five species, including new ones) and Mangyshlak (12 species), as well as the published data on Olenekian species from Eurasia (e.g., Chen et al, 2005; Xu and Grant, 1994; Chen, 1983; Xu and Liu, 1983) and North America (e.g., Perry and Chatterton, 1979; Girty, 1927), illustrates a general trend in marked rising of taxonomic diversity of Mesozoic-type brachiopods from the Lower Induan through the Upper Olenekian. The Induan, lower, middle and upper Olenekian in SP are characterised by 20, 11 and 30 species, respectively. Similar changes in ammonoid succession, with highest diversification for the early Olenekian, occur in some other Tethyan regions. According to published data (Chen et al., 2005) 13 articulated brachiopod lineages at familial level survived the End-Permian mass extinction. As a result, at least 12 Palaeozoic type articulated brachiopod genera occur in the Lower Triassic, mainly in the Lower Griesbachian; at the same time the Permian-type Prelissorhynchia at Mangyshlak was found by us in the first time in the upper Olenekian. However, only four ammonoid lineages at familysuperfamily level survived the mentioned mass extinction; apparently only a single Palaeozoic-type ammonoid genus (Episageceras) is known from the Lower Triassic (Zakharov, 1978).