Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis: Additional Evidence from the Lower Triassic of the Russian Far East and Kazakhstan

Home , Albanites, Leiophyllites, Ophiceras, Smolyaninovo, Tirolites, Ussuria

Journal of Earth Science, Vol. 25, No. 1, p. 1–44, February 2014 ISSN 1674-487X Printed in China DOI: 10.1007/s12583-014-0398-6

Yuri D Zakharov*, Alexander M Popov Far Eastern Geological Institute, Far Eastern Branch, Russian Academy of Sciences, Stoletiya Prospect 159, Vladivostok 690022, 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 sluggish- nektic 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 vulner- abilities 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 success- fully 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 Early Triassic (mainly Olenekian) ammonoids are abundant The Permian-Triassic boundary (PTB) is associated with and diverse in many regions of the world, but the most diverse the largest extinction event of Phanerozoic time (e.g., Kozur, Olenekian brachiopod faunas have been discovered in the for- 2007; Yin and Zhang, 1996; Erwin, 1994). Early Triassic suc- mer USSR area (South Primorye and Mangyshlak). The goal of cessions of both benthonic and nektonic marine organisms this work is to analyze taxonomic diversity patterns of Early contain important information about the postextinction biotic Triassic articulated brachiopods and ammonoids from the for- recovery interval. However, in spite of significant progress in mer USSR (Fig. 1) in order to reconstruct biotic recovery pat- the past decades in investigation of Early Triassic brachiopods terns after the End-Permian mass extinction. (e.g., Ruban, 2009; Chen et al., 2006, 2005a, b; Shen and Shi, 1996; Shen and He, 1994; Xu and Grant, 1994; Jordan, 1993; 2 MATERIALS MacFarlan, 1992; Chen, 1983; Xu and Liu, 1983) and ammon- Original materials on Early Triassic invertebrates used for oids (e.g., Brühwiler et al., 2012a, b, 2010a, b, c, 2008; Smy- our investigation were obtained by the authors from Induan shlyaeva and Zakharov, 2012; Ware et al., 2011; Guex et al., and Olenekian sections of South Primorye during the last five 2010; Brayard et al., 2009a, b, c, 2007a, b, 2006a, b; Shigeta et years. Investigation of Olenekian brachiopods from Man- al., 2009; Brayard and Bucher, 2008; Mu et al., 2007; Shevyrev, gyshlak, Kazakhstan was made on the basis of Zakharov Y D’s 2006, 1986; Krystyn et al., 2003; Ermakova, 2002; Zakharov, collection from his Kara-Tau expedition in 1978. 1997, 1996, 1978; Dagys and Ermakova, 1996, 1988; Krystyn and Orchard, 1996; Waterhouse, 1996; Tozer, 1994; Guex, 3 LOWER TRIASSIC AMMONOID AND BRACHIO- 1978), our knowledge of these important invertebrate groups, POD BIOSTRATIGRAPHY especially Mesozoic-type Brachiopodaremains incomplete. 3.1 South Primorye (Russian Far East) The Early Triassic epiplatformal terrigenous assemblage *Corresponding author: [email protected] of South Primorye (Fig. 1) lies unconformably on the Upper © China University of Geosciences and Springer-Verlag Berlin Palaeozoic and more ancient marine and continental sedimen- Heidelberg 2014 tary and volcano-sedimentary deposits, volcanics and grani- toids (Zakharov et al., 2010; Zakharov, 1978, 1968). In many Manuscript received June 11, 2012. areas of South Primorye (e.g., Russian Island, Ussuri and Amur Manuscript accepted September 28, 2012. gulfs, Abrek Bay and Artyom area), the bulk of the Induan or

Zakharov, Y. D., Popov, A. M., 2014. Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis: Addi- tional Evidence from the Lower Triassic of the Russian Far East and Kazakhstan. Journal of Earth Science, 25(1): 1–44, doi:10.1007/s12583-014-0398-6 2 Yuri D Zakharov and Alexander M Popov

Induan to earliest Olenekian Lazurnaya Bay Formation con- posed only on Russian Island and consists of sandstone with sists of shallow-marine shelf sediments. These are predomi- thick lenses of sandy limestone. It is represented by a single nantly conglomerate and sandstone with lenses of sandy zone, the Tirolites-Amphistephanites Zone (Figs. S3 and S6). limestone-coquina dominated by bivalves. The lower part of The Zhitkov Cape Formation of Smithian to Spathian age the Induan is represented by the Tompophiceras ussuriense represents deeper shelf facies characterised by the dominance of Zone, its upper part corresponds to the Gyronites subdharmus mudstone and siltstone with numerous calcareous-marly nodules Zone (Figs. S1 and S2). and lenses and thin, irregular beds of sandstone. The lower part The Tobizin Cape Formation, of Smithian age, is present of the formation, which is found in the West Ussuri Gulf-Strelok on Russian Island and on the western coast of Ussuri Gulf, Strait area (Figs. S1 and S7), is represented by the “Hedenstroe- where it conformably overlies the Lazurnaya Bay Formation. It mia” bosphorensis and Anasibirites nevolini zones and is is represented by sandstone with lenses of coquinoid calcare- Smithian in age. The upper part of the formation was formed in ous sandstone with mainly cephalopod and bivalve remains. this area during the Early (Tirolites-Amphistephanites Zone) and On Russian Island, the following zones can be recognized: Late (Neocolumbites insignis and Subfengshanites multiformis “Hedenstroemia” bosphorensis below and Anasibirites nevolini zones) Spathian. On Russian Island (Figs. S5–S7) and the west- above (Figs. S3 and S4). ern coast of Amur Gulf (Zakharov et al., 2005), only the upper The Schmidt Cape Formation of early Spathian age is ex- Olenekian Zhitkov Cape Formation is present. However, the

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132°E Figure 1. (a) Location of study areas in the former USSR. 1. South Primorye and 2. Mangyshlak. (b) Study sections in South Pri- morye area. 1. Seryj-Tri Kamnya; 2. Abrek; 3. Konechnyj; 4. Tobizin; 5. Ayax-Balka; 6. Zhitkov; 7. Golyj; 8. Paris.

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 3

Olenekoceras-bearing part of the Spathian on the western coast gen. and sp. nov. B), Norellidae (Piarorhynchella? sp. nov.) of Amur Gulf is characterised mainly by fine-grained, striped and Rhynchonellidae (Abrekia sulcata, Rhynchonellidae gen. sandstone (Atlasov Cape Formation). and sp. nov.). For detailed biostratigraphic distribution data, The Karazin Cape Formation represents the Anisian eve- see Figs. S1 and S2. Among these brachiopod families, the rywhere in South Primorye. No benthic forms of allochthonous Pontisiidae and the Wellerellidae are Paleozoic-type ones, but origin have been found in this formation and therefore it seems the Norellidae and the Rhynchonellidae seem to have origi- to have accumulated under offshore environmental conditions. nated during the Early Triassic (Induan). It composed mainly of fucoid sandstone with large calcareous Newly found Dienerian brachiopod shells in South Pri- septarian nodules. The Lower Anisian in South Primorye is morye, including specimens of Abrekia, first described by A. S. represented by the Ussuriphyllites amurensis and Leiophyllites Dagys (see Supplement), are generally very small as compared pradyumna zones (Figs. S5–S7). with Late Permian ones (Tables S2–S4)—their maximal shell The detailed lithological succession and distribution of length is only 7.8 mm. brachiopod and ammonoid species in the Lower Triassic of South Primorye are shown in some figures (Figs. S1 and S8). 3.1.3 “Hedenstroemia” bosphorensis Zone (Smithian) Figure 2 illustrates the stratigraphic distribution of brachiopods The early Smithian “Hedenstroemia” bosphorensis Zone and ammonoids at the family and order levels (within the Up- has been subdivided into a lower unit (“Gyronites separatus” per Changhsingian-Spathian interval). (=Ussuriflemingites abrekensis) beds) and an upper unit (Eu- flemingites prynadai beds) (Zakharov, 1997). The 16 to 50-m- 3.1.1 Tompophiceras ussuriense Zone (Griesbachian) thick “Gyronites separatus” beds are characterised by the oc- The ammonoid assemblage of the Griesbachian Tompo- currence of the prolecanitid ammonoids Aspenitidae (Parahe- phiceras ussuriense Zone (at least 18 m thick) is characterised densroemia sp., P. kiparisovae) and Sageceratidae (Pseudos- by the presence of two ceratitid ammonoid genera: Tompo- ageceras cf. multilobatum). Numerous ceratitid ammonoids phiceras (T.), occurred together with inarticulated brachiopods also occur: Meekoceratidae (Ambitoides orientalis, Gurleyites (Lingula sp.) and some bivalves, and Lythophiceras (Lytho- sp.), Flemingitidae (Ussuriflemingites abrekensis), Xenoceltiti- phiceras sp.), the latter occurring in the upper part of the zone. dae (Preflorianites cf. radiatus), Proptychitidae (Paranorites For detailed ammonoid distribution data, see Fig. S1. However, varians, Kummelia? sp.), Arctoceratidae (Arctoceras septen- no Griesbachian articulated brachiopods were found in the Far trionale), Clypeoceratidae (Clypeoceras spitiense), and Colum- East apparently because of their extreme rarity there. A re- bitidae (Proharpoceras carinatitabulatum). markable feature of the Griesbachian ceratitid ammonoid as- The Euflemingites prynadai beds, which are at least 106 semblage from South Primorye is that Tompophiceras is a rep- m thick, are also characterised by the presence of prolecanitid resentative of the Palaeozoic-type family (Dzhulfitidae) (Fig. ammonoids Hedenstroemiidae (“Hedenstroemia” bosphoren- 2). However, Lythophiceras belongs to a Mesozoic-type family sis), Aspenitidae (Parahedenstroemia conspicienda), Sage- (Ophiceratidae). ceratidae (e.g., Pseudosageceras multilobatum), Ussuriidae (e.g., Ussuria schamarae), and numerous ceratitid ammonoids: 3.1.2 Gyronites subdharmus Zone (Dienerian) Meekoceratidae (e.g., Meekoceras subcristatum, Abrekites The Dienerian Gyronites subdharmus Zone (45–110 m edites, A. planus), Dieneroceratidae (e.g., Dieneroceras chaoi), thick) is marked by the first appearance datum of the ceratitid Inyoitidae (Inyoites spicini), Prionitidae (Radioprionites abre- ammonoid genus Gyronites (G. subdharmus Kiparisova), used kensis), Flemingitidae (Euflemingites prynadai, E. artyomensis, here for recognition of the base of the Dienerian. Other cerati- Flemingites radiatus, F. aff. glaber, Balhaeceras balhaense, tid ammonoid genera in the lower part of the zone are Lytho- Rohillites laevis, Palaeokazakhstanites ussuriensis), Palaeo- phiceras (L. eusakuntala), Proptychites (P. hiemalis and possi- phyllitidae (Anaxenaspis orientalis, Eophyllites sp., E. ascoldi- bly Tompophiceras (Tompophiceras sp.), in the upper part— ensis), Xenoceltitidae (Preflorianites cf. radiatus), Arcto- Ussuridiscus (U. varaha), Wordieoceras (W. cf. wordiei), ceratidae (e.g., Arctoceras septentrionale), Clypeoceratidae Dunedinites (D. magnumbilicatus), Melagathiceratidae (Mela- (Clypeoceras timorense), Melagathiceratidae (Juvenites sim- gathiceratidae gen. et sp. indet.), and Pachiproptychites (P. plex, J. dieneri, J. cf. septentrionalis, J. aff. sinuosus, Pro- otoceratoides). All these ceratitid ammonoids of the Gyronites sphingitoides ovalis), and Owenitidae (Owenites koeneni). The subdharmus Zone, with the exception of the Dzhulfitidae Early Smithian ammonoid succession is dominated by ceratitid (Tompophiceras) and Proptychitidae (e.g., Proptychites and ammonoids, all of which seem to be Mesozoic-type ones, as Pachiproptychites), are representatives of typical Mesozoic- well as by four other prolecanitid ammonoid families. type families (Ophiceratidae, Meekoceratidae, Mela- No articulated brachiopods were found in the lower part gathiceratidae, and possibly Paranannitidae). The Paleozoic- of the “Hedenstroemia” bosphorensis Zone. The articulated type Dzhulfitidae was discussed earlier, but the ancestral group brachiopod assemblage of the Euflemingites prynadai beds is of the Proptychitidae is unknown at present (Fig. 2). characterised by species of the families Wellerellidae The articulated brachiopod assemblage of the Dienerian (Wellerellidae gen. and sp. nov.), Norellidae (Piarorhynchella? Gyronites subdharmus Zone is characterised by the presence of sp. nov.), and Rhynchonellidae (Abrekia sulcata). For detailed four families of the order Rhynchonellida (Figs. 2 and 3): Pon- biostratigraphic distribution data, see Figs. S1 and S4. Among tisiidae (Lissorhynchia sp., Pontisiinae gen. and sp. nov.), these three early Smithian articulated brachiopod families, only Wellerellidae (Wellerellidae gen. and sp. nov. A, Wellerellidae the Wellerellidae is a Palaeozoic-type one (Fig. 2). The

4 Yuri D Zakharov and Alexander M Popov

Permian Triassic Changh. Induan Olenekian

Order - Griesbachian Dienerian Smithian Spathian Lissorhynchia Pontisiidae

Wellerellidae Piarorhynchella? Norellidae

Rhynchonellida Abrekia Rhynchonellidae Hustedtiella Neoretziidae

Brachiopods Lepismatina Lepismatinidae

Sp. At. “”Fletcherithyris Dielasmatidae

Zeilleriidae?

Terebr.

Sageceratidae

Khvalynitidae

Ussuriidae

Prolecanitida Pleuronodoceratidae Aspenitidae

Huananoceratidae Hedenstroemiidae Xenodiscidae Ophiceratidae

Tompophiceras Meekoceratidae Dzhulfitidae Dieneroceratidae Tirolitidae

Inyonitidae

Prionitidae

Ammonoids Flemingitidae

Palaeophyllitidae

Ceratitida Xenoceltitidae Keyserlingitidae Proptychitidae Olenikitidae Arctoceratidae Stephanitidae Clypeoceratidae Melagathiceratidae

Procarnitidae Paranannitidae

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.

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. 90–41, x 2, Seryj Cape, Dienerian; 3–6. Prelissorhynchia (?) sp. nov., four views of same specimen Nos. 405-9(4), x 2, Dolnapa, upper Spathian; 7–10. Rhynchonellaceae gen. and sp. indet., four views of same specimen No. 97-42(1), x 2, Tri Kamnya Cape, Smithian; 11–14. Rhynchonellidae gen. and sp. nov. A, four views of same specimen No. 97-42(2), x 2, Tri Kamnya Cape, Smithian; 15–18. Rhynchonellidae gen and sp. nov. 2, four views of same specimen No. 97-42, x 2, Tri Kamnya Cape, Smithian; 19–22. Lissorhynchia sp. nov., four views of same specimen No. 405-4, x 2, Dolnapa, Upper Spathian; 23–26. Hustedtiella planicosta Dagys, four views of same specimen No. 405-9, x 2, Dolnapa, upper Spathian; 27–30. Spirigerellina sp., four views of same specimen No. 409-4(2), x 2, Dolnapa, Upper Spathian; 31–34. Lepismatina mansfieldi (Girty), four views of same specimen No. 405-4 (12), x 2, Dolnapa, Upper Spathian; 35–38. Spiriferinidae gen. and sp. nov. A, four views of same specimen No. 20P, x 1, Paris Bay, upper Spathian.

6 Yuri D Zakharov and Alexander M Popov

‘Lilliput effect’ continuous to be evident among early Smithian are Palaeozoic-type ones. The shell sizes of all these early brachiopods of these families, which have maximal shell lengths Spathian brachiopod families are quite reduced (Tables S5 and of only 9.2 mm (observation from 53 specimens) (Table S4). S6)), with the exception of some dielasmatid brachiopods (e.g., “Fletcherithyris” margaritovi) that are up to 27 mm long (Ta- 3.1.4 Anasibirites nevolini Zone (lower Olenekian) ble S7). The late Smithian Anasibirites nevolini Zone (20–68 m thick) is marked by the first appearance datum of the genus 3.1.6 Neocolumbites insignis Zone (middle Spathian) Anasibirites. The assemblage is characterised by rare occur- The middle Spathian Neocolumbites insignis Zone (66–95 rences of the prolecanitid ammonoids Sageceratidae (Pseudos- m thick) is marked by the first appearance datum of the genus ageceras sp. indet.) and Aspenitidae (Parahedenstroemia Neocolumbites. The ammonoid assemblage is characterised by nevolini), but by numerous ceratitid ammonoids, represented species of both the Prolecanitida and Ceratitida orders. Cerati- by Meekoceratidae (e.g., Meekoceras subcristatum), Tirolitidae tid ammonoids are represented by Meekoceratidae (Svalbard- (Bandoites elegans), Prionitidae (e.g., Hemiprionites contortus, iceras zhitkoviense), Tirolitidae (Tirolites cf. subcassianus Shigetaceras dunajense, Arctoprionites ovalis, Anasibirites Zakharov), Palaeophyllitidae (Burijites skorochodi, Leiophyl- nevolini, Wasatchites sikhotealinensis, Gurleyites maichensis), lites praematurus), Keyserlingitidae (e.g., Olenekoceras merid- Palaeophyllitidae (Burijites skorochodi, Anaxenaspis sp. nov.), ianus), Columbitidae (e.g., Neocolumbites insignis, Columbites Xenoceltitidae (Preflorianites? sp.), Arctoceratidae (e.g., Arc- ussuriensis, Procolumbites subquadratus, Hellenites inopina- toceras labogense, Churkites syaskoi), Melagathiceratidae tus), and Procarnitidae (Procarnites sp.). The prolecanitid am- (Juvenites simplex), Paranannitidae (Paranannites minor, Pro- monoids Sageceratidae (Pseudosageceras sp.) and Khvalyniti- sphingitoides ovalis), and Owenitidae (Owenites koeneni). For dae (Khvalynites unicus) are present in lower abundance. The detailed ammonoid distribution data, see Figs. S3 and S4. The ammonoid assemblages of this zone are dominated by Cerati- late Smithian ammonoid succession is dominated by ceratitid tida. All middle Spathian ammonoid families seem to be ammonoids, all of which seem to be Mesozoic-type ones, as Mesozoic-type ones. Small spiriferinid brachiopod shells, up to well as by two other prolecanitid ammonoid families (Fig. 2). 12 mm in length, were found in this zone. Very rare and small rhynchonellid brachiopods (up to 6 mm in length) were found in the zone, but their preservation is often 3.1.7 Subfengshanites multiformis Zone (upper Spathian) moderate to poor in clayey facies. The late Spathian Subfengshanites multiformis Zone (16– 18 m thick) is marked by the first appearance datum of the 3.1.5 Tirolites-Amphistephanites Zone (lower Spathian) genus Subfengshanites. This zone is characterised by the oc- The early Spathian Tirolites-Amphistephanites Zone (40– currence of abundant ceratitid and very rare prolecanitid 50 m thick) is marked by the first appearance datum of the (Pseudosageceras sp.) ammonoid species. The main ceratitid genera Tirolites and Amphistephanites, used here for recogni- ammonoid taxa known from this level are the Dieneroceratidae tion of the base of the Spathian. The assemblage is character- (Dieneroceras karasini), Palaeophyllitidae (Palaeophyllites ised by the prolecanitid ammonoids Sageceratidae (Pseudos- superior, Leiophyllites sp.), Paranannitidae (e.g., Zhitkovites ageceras sp.) and Aspenitidae (Parahedenstroemia sp.) and by insularis (Kiparisova), Pseudoprosphingites globosus, Isculi- ceratitid ammonoids, represented by Meekoceratidae (Baja- toides? suboviformis), Columbitidae (Subfengshanites multi- runia dagysi), Tirolitidae (Tirolites ussuriensis, T. subcassianus, formis, Arnautoceltites gracilis, and Prenkites aff. timorensis). Bandoites elegans), Dinaritidae (Tchernyschewites costatus), All recognized ammonoid families seem to be Mesozoic-type Olenikitidae (e.g., Kazakhstanites sonticus), Stephanitidae ones. (Amphistephanites parisensis) and Flemingitidae The Subfengshanites multiformis Zone is characterised (Guangxiceras tobisinense). The early Spathian ammonoids also by the occurrence of Rhynchonellidae (Rhynchonellidae assemblages of the Tirolites-Amphistephanites Zone are domi- gen. et sp. nov. 1, 2 and 3), Lepismatinidae (Lepismatinidae nated by Ceratitida, whereas Prolecanitida are very rare. All gen. and sp. nov.), and Neoretziidae (Hustedtiella planicosta). recognized ammonoid families seem to be Mesozoic-type ones. For detailed biostratigraphic distribution data and rhynchonel- This time interval studied in South Primorye is character- lid brachiopod shell size (see Figs. S5, S8, and Table S4). Ob- ised by the presence of abundant articulated brachiopods, rep- served ranges across the Subfengshanites multiformis Zone are resented by the next families of the orders Rhynchonellida, summarized in Fig. 2. Among brachiopod families only the Athyritida, Spiriferinida, and Terebratulida: Rhynchonellidae Neoretziidae crossed the PTB (e.g., Rhynchonellidae gen. and sp. nov. A), Neoretziidae (e.g., Hustedtiella planicosta), Lepismatinidae (Lepismatina aff. 3.2 Mangyshlak (Kazakhstan) mansfieldi), Diplospirellidae (Spirigerellina pygmaea), Di- The Spathian in Mangyshlak has been investigated in de- elasmatidae (“Fletcherithyris” margaritovi), Zeilleriidae? tail in the Dolnapa Well reference section (e.g., Zakharov et al., (Zeilleriidae? gen. and sp. indet.), and Zeilleriidae? (e.g., 2008; Gavrilova, 2007; Balini et al., 2000; Shevyrev, 1968), Zeilleriidae? gen. and sp. nov. A). For detailed biostratigraphic located in the Kara-Tau area. Two lithostratigraphic units, cor- distribution data (see Figs. S3 and S8). Observed ranges across responding to the Olenekian Tjururpiniaya Series, can be rec- the Tirolites-Amphistephanites Zone are summarized in Fig. 2. ognized in this section: the Tartalinskaya Fm. (Tirolites However, of the five brachiopod families known from this cassianus-Kiparisovites carinatus and Columbites parisianus- stratigraphic interval, two (Neoretziidae and Dielasmatidae) Procolumbites zones), which is composed of mudstone, silt-

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 7 stone and fine-grained sandstone with calcareous interlayers, ammonoid assemblages of the Tirolites cassianus-Kiparisovites and the Karadzhatykskaya Fm. (Arnautoceltites bajarunasi- carinatus Zone are dominated by Ceratitida, and no Stacheites undatus Zone and Eumorphotis beds), which con- Palaeozoic-type ammonoid families were identified. sists of intercalations of mudstone with calcareous boulders, Articulated brachiopods are represented by three orders siltstone with plant detritus, and fine-grained sandstone with (Rhynchonellida, Spiriferida, and Terebratulida) and several interlayers of limestone (Fig. S9). families: Norellidae (Piarorhynchella mangyshlakensis), Le- pismatinidae (Lepismatina sp. nov.), and Dielasmatidae 3.2.1 Tirolites cassianus-Kiparisovites carinatus Zone (“Fletcherithyris” margaritovi) (Fig. 4). No Palaeozoic-type (Spathian) brachiopod families were discovered in this zone. As in the The ammonoid assemblage of the early Spathian Tirolites case with the Lower Triassic of South Primorye, size reduction cassianus-Kiparisovites carinatus Zone (which is at least 113 (‘Lilliput effect’) has been observed in many brachiopods from m thick) is characterised by the presence of two ammonoid the Tirolites cassianus-Kiparisovites carinatus Zone (Table S4), orders: the Prolecanitida (Sageceratidae—Pseudosageceras with the exception for very rare Dielasmatidae (up to 20 mm in longilobatum) and the Ceratitida. The latter consists of Dinari- length). tidae (“Dinarites” asiaticus, “D.” orientalis), Olenikitidae (Tjururpites cf. costatus, Kiparisovites carinatus, Hyrcanites 3.2.2 Columbites parisianus-Procolumbites Zone (middle nodosus, Kazakhstanites dolnapaensis), Tirolitidae (e.g., Tiro- Spathian) lites cf. cassianus), ?Prionitidae (Albanites gracilis) and The ammonoid assemblage of the middle Spathian Colum- Doricranitidae (Doricranites) (Fig. 4). The early Spathian bites parisianus-Procolumbites Zone (274 m thick) is characterised by the presence of prolecanitid ammonoids Sageceratidae (Pseudosageceras longilobatum) and Khvalynitidae (Khvalynites Triassic mangyshlakensis) and abundant ceratitid ammonoids: Dinariti- Upper Olenekian dae (“Dinarites” asiaticus, “D.” orientalis), Xenoceltitidae

Order Smithian Spathian (Xenoceltites mangyshlakensis, X. bajarunasi, Preflorianites Piarorhynchella kiparisovae), Kashmiritidae (Kashmirites subdimorphus, K. Rhynchonellidae Prelissorhynchia (?) popowi Shevyrev, Eukashmirites subdimorphus), Olenikitidae Pontisiidae (Tjururpites costatus, Kiparisovites ovalis, Kazakhstanites Piarorhynchella Lissorhynchia Rhynch. Norellidae dolnapaensis), Tirolitidae (Tirolites armatus, T. longilobatus, T. Hustedtiella rossicus), Doricranitidae (Doricranites acutus, D. bogdoa- Neoretziidae Spirigerellina nus), ?Prionitidae (Albanites gracilis), Palaeophyllitidae (Leio- At. Diplospirellidae phyllites exacudus), Procarnitidae (Procarnites kokeni), and Lepismatina Brachiopods Lepismatinidae Columbitidae (Columbites cf. parisianus, C. ventroangustus, Sp. “”Fletcherithyris Procolumbites karatauchicus, Hellenites kazakhstanicus, Man- Dielasmatidae Antezeilleria gyshlakites mirificus) (Fig. 4). The early Spathian ammonoid Antezeilleriidae assemblages of the Columbites parisianus-Procolumbites Zone Thyratryaria Terebr. Family uncertain are dominated by Ceratitida, and no Palaeozoic-type ammonoid Proanadyrella? families were recognized. Sageceratidae Articulated brachiopods are represented by four orders Khvalynitidae

Prolec. (Rhynchonellida, Athyridida, Spiriferinida, and Terebratulida), Dinaritidae among which a number of families were recognized: Pontisii- dae (Prelissorynchia (?) sp. nov., Lissorhynchia sp. nov., Kashmiritidae Piarorhynchella mangyshlakensis), Neoretziidae (Hustedtiella Xenoceltitidae planicosta), Diplospirellidae (Spirigerellina sp., Spirigerellinae gen. and sp. indet.), Lepismatinidae (Lepismatina sp. nov.), Olenikitidae Antezeilleriidae (Antezeilleria sp.), and an uncertain family Tirolitidae (Proanadyrella (?) sp., Thyratryaria sp. nov. A, Thyarotryaria

Ammonoids

Ceratitida Doricranitidae sp. nov. B) (Fig. 4). The genus Prelissorynchia (Pontisiidae) possibly crossed the PTB; formerly, this genus was considered Prionitidae to be only Late Paleozoic in age. Brachiopod miniaturization is Palaeophyllitidae also recorded for this level, with shell lengths for all of the families above measuring between 3.4 and 9.4 mm (Tables S5 Procarnitidae and S6). Columbitidae

3.2.3 Arnautoceltites bajarunasi-Stacheites undatus Zone Figure 4. Stratigraphic range of Olenekian articulated (upper Spathian) brachiopod and ammonoid families in Mangyshlak. Abbre- The ammonoid assemblage of the late Spathian Arnauto- viations: Prolec.. Prolecanitida; Rhynch.. Rhynchonellida. celtites bajarunasi-Stacheites undatus Zone (172 m thick) is See Fig. 2 for further details. characterised by the presence of prolecanitid ammonoids Sage-

8 Yuri D Zakharov and Alexander M Popov ceratidae (Pseudosageceras sp.) and some ceratitid ammonoid senting different taxonomic levels) survived the mass extinc- families: Dinaritidae (“Dinarites”orientalis, Stacheites undatus, tion: (1) Xenodiscidae and Dzhulfitidae (at the family level), (2) S. concavus), Xenoceltitidae sp., Olenikitidae (Kazakhstanites Otoceratoidea (at the superfamily level), and (3) Episageceras dolnapaensis, Tjururpites costatus, Kiparisovites ovalis), Tiro- (at the generic level). Only a single Palaeozoic-type genus litidae (Tirolites armatus), Prionitidae (Arctoprionites sp., Al- (Episageceras) is for certain known from the Lower Triassic banites gracilis), and Columbitidae (Arnautoceltites bajarun- (Fig. 5). In Siberia and the Russian Far East, representatives of asi). The late Spathian ammonoid assemblages of the Arnauto- Episageceras were collected from the lower and upper parts of celtites bajarunasi-Stacheites undatus Zone are dominated by the Induan (Dagys and Ermakova, 1996; Zakharov, 1978; Ceratitida, and no Palaeozoic-type ammonoid families were Popow, 1961). About 18 Mesozoic-type genera have been re- recognized. ported from the Induan (Table S1). Three articulated brachiopod orders (Rhynchonellida, A few ammonoid genera (Hypophiceras fauna) have been Spiriferinida, and Terebratulida) were found in the Arnauto- recorded in the upper Changhsingian of the Meishan Section in celtites bajarunasi-Stacheites undatus Zone. They are repre- South China, identified as Pseudogastrioceras, Otoceras (?), sented by the families Pontisiidae (Lissorhynchia sp. nov., Hypophiceras, Metophiceras, Tompophiceras, “Glypto- Piarorhynchella mangyshlakites), Lepismatinidae (Lepismat- phiceras”, and Pseudosageceras (Yang et al., 1996; Yin et al., ina sp. nov.), and an uncertain family (Thyratryaria aff. pertu- 1996). McGovan and Smith (2007) have shown in their Fig. 7 mida). Among these brachiopod families, only a single that Aldanoceras, as well as Otoceras, Xenodiscus, Hypo- Palaeozoic-type one (Pontisiidae) has been discovered. The phiceras, Tompophiceras, Metophiceras and Episageceras, ‘Lilliput effect’ is also prevalent among the brachiopod fauna crossed the PTB (established stratigraphic ranges are shown of the Arnautoceltites bajarunasi-Stacheites undatus Zone. there as solid lines), using Dagys and Ermakova’s (1996) data Lissorhynchia exhibits shell lengths to a maximum of 5.3 mm, on the Verkhoyansk area and taking into account information and other latest Spathian brachiopods are only marginally lar- on Hypophiceras fauna of the Meishan area. As was recently ger (up to 7–10 mm) (Table S4). emphasized by Shevyrev (2006), ammonoids of the so-called Hypophiceras fauna in Meishan represent a mixed assemblage 3.2.4 Eumorphotis beds (upper Spathian) of Permian and Triassic forms. The composition of the assem- In the uppermost part of the Spathian in the Dolnapa Well blage at Meishan is in doubt because of poor preservation, Section, the ~151-m-thick Eumorphotis beds yielded very rare where ammonoid shells are strongly deformed and suture lines prolecanitid ammonoids (Pseudosageceras sp.) together with are usually unobservable (Shevyrev, 2006). In this situation, bivalves, gastropods, and nautiloids. However, ceratitid am- some Permian araxoceratid ammonoids, in Shevyrev’s (2006) monoids and articulated brachiopods have not been discovered opinion, could have been mistaken for Triassic Otoceras, Per- here. mian Paratirolites or Pseudotirolites for Triassic Tompo- phiceras, and Permian Xenodiscus for Triassic Hypophiceras. 4 DISCUSSION However, the age of Otoceras-bearing sequences in the Boreal 4.1 Overview of Early Triassic Brachiopod and Ammonoid realm is still under debate now (e.g., Bjerager et al., 2006; Faunas Baud and Beuchamp, 2001; Stemmerik et al., 2001; Kozur, 4.1.1 Palaeozoic-type taxa 1998; Orchard and Krystyn, 1998), and only further progress in About 13 families of articulated brachiopods survived the the carbon-isotopic, palaeomagnetic and micropalaeontological End-Permian mass extinction. Among them, 13 Palaeozoic- investigation could solve this very important problem. type brachiopod genera have been identified in the Lower Tri- According to Brayard et al. (2007a), the Smithian genus assic: Cathaysia, Paryphella, Spinomarginifera, Retimarginif- Proharpoceras probably was derived from the Wuchiapingian era (?), Acosarina, Prelissorhynchia (?), Araxathyris, Spirig- Anderssonoceratidae (Otoceratina), thus implying that an addi- erella, Orbicoella, Crurithyris, Paracrurithyris, Paras- tional lineage among Ceratitida crossed the PTB. However, piriferina, and Tethyochonetes (e.g., Shen et al., 2006; Chen et this inference concerning two taxa from distant stratigraphic al., 2005b; this study, see Table S1). They belong to the Paras- levels needs to be tested through further morphological analy- piriferinidae, Schizophoriidae, Productellidae, Ambocoellidae, sis and additional fossil finds. Pontisiidae, and Athyrididae families. All these genera, with The new evidences and published data show that in con- the exception of Orbicoelia, Prelissorhynchia (?) and possibly trast to the brachiopod fossils only restricted ammonoid line- Retimarginifera, were discovered only in the lower Griesba- ages at the generic and family levels servived the End-Permian chian. However, Retimarginifera (?) was reported from the mass extinction. Smithian (Chen et al., 2005b), Prelissorhynchia (?) from both the Griesbachian (Chen et al., 2005b) and the middle Spathian 4.1.2 Mesozoic-type taxa (this study), and Orbicoelia only from the upper Olenekian About 15 Mesozoic-type articulated brachiopod genera (Chen et al., 2005b). Besides, at least six additional Permian- were found in the Induan (Table S1), some of which are known type brachiopod genera belonging to six other families (Dip- from Russia (e.g., Lissorhynchia, Abrekia, Rhynchonellidae lospirellidae, Neoretziidae, Laballidae, Pennospiriferinidae, gen. nov. 4, Piarorhynchella (?), Wellerellidae gen. nov. A, Dielasmatidae, and Angustothyrididae) also may have survived Wellerellidae gen. nov. B, and Pontisiinae gen. nov.). However, into the Lower Triassic (Fig. 5). Chen et al. (2005a) reported only 7 Mesozoic-type genera from However, for certain only four ammonoid lineages (repre- the Induan of South China (e.g., Shen et al., 2006; Chen et al.,

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 9

2005b; Shen and He, 1994), Idaho (Girty, 1927), Himalayas Prelissorhynchia (?)) but 40 Mesozoic-type genera are known. (Bittner, 1899), and North Caucasus (Dagys, 1974). The taxo- They are identified as Maorirhynchia (?), Rhynchonellidae gen. nomic diversity of brachiopods during the Dienerian was only nov. 1, 2 and 3, Nudirostralina, Abrekia, Sinuplicorhynchia (?), about 11% of that for the Changhsingian (Fig. 6). Paranorellina, Piarorhynchella, Aparimarhynchia, Lis- Among Olenekian articulated brachiopods, only three sorhynchia, Spirigerellina, Spirigerellinae gen. nov., Neoretzia, Palaeozoic-type genera (Retimarginifera (?), Orbicoelia, and Hustedtiella, Lepismatina, Spiriferinidae gen. nov. A,

Figure 5. Post-extiction survival of Paleozoic-type brachiopods and ammonoids (Chen et al., 2005a, b; this study). Abbrevia- tions: Capitan.. Capitanian; An.. Anisian. See Fig. 2 for Early Triassic substages.

10 Yuri D Zakharov and Alexander M Popov

246 240

220

200

180

160 143

140 140 extinction Mass

120 120

Ammonoids

100 extinction Mass 100

Articulated (136) brachiopods 80 68 (129 ) 80

Number of genera of Number 60 60 41 43

40 genera of Number 40 27 (60) (27?) 20 20 (23?) (16) (16) 12 0 0 Gri. Die. Smi. Spa. Gri. Die. Smi. Spa. Changh. InduanOlenek.AnisianChangh. Induan Olenek. Anisian Perm.Triassic Perm. Triassic

1 8 20 19 14 1 7 7?

Postextinction Postextinction No any trends Occured miniaturization miniaturization

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”, Rhaet- until the Spathian or Anisian. However, recent data (Brühwiler ina, Portneufia, Antezeilleria, Zeilleriidae? gen. nov. A, B, C, et al., 2011a, b, 2010a, b, c; Guex et al., 2010; Brayard et al., D and F, Periallus, Protogusarella, Thyratryaria, Pro- 2009a, b; this study) show that the recovery of ammonoid fau- anadyrella, and Vex, of which twelve are new genera (Table nas proceeded significantly more rapidly. Brayard et al. (2009a, S1). Chen et al. (2005b) reported only 18 Mesozoic-type b) concluded that ammonoid diversity level was higher than brachiopod genera in the Olenekian on the basis of data from the maximum level recorded during all the Permian by the South China, Southwest Japan, Idaho, Himalayas, Tibet, Do- Smithian. According to our data, based on recently published brogea, New Zealand, and Spitsbergen. The taxonomic diver- and original evidences, ammonoids reached levels of taxo- sity of brachiopods during the Smithian and Spathian was only nomic diversity during the Dienerian that were 146% higher about 16% and 19% of that for the Changhsingian, respectively than in the Changhsingian, representing a much faster recovery (Fig. 6). than that for brachiopods (the diversity of brachiopods during In this stage of our knowledge we conditionally eccept for the Dienerian was only about 8% of that for the Changhsingian) the Induan ammonoids that they are represented by only a sin- (Fig. 6). Maximum taxonomic diversity of Permian ammon- gle Palaeozoic-type genus (Episageceras) but 67 Mesozoic- oids is associated with the Wordian-Roadian (Bogoslovskaya et type genera (Fig. S12). Only Mesozoic-type ammonoid genera al., 1999). However, the diversity of ammonoids during the have been reported from the Smithian and Spathian which have Smithian (Table S1) reached not less than 200% of that for the yielded 136 and 129 genera, respectively (Fig. 6; Table S1). Wordian. McGovan (2005) inferred that ammonoids did not fully recover

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 11

4.2 Patterns of Geographical Differentiation of Early Tri- usual oceanic and climatic conditions that affected the marine assic Brachiopod and Ammonoid Faunas biota for several million years (Algeo and Twitchett, 2010). Early Triassic ammonoid diversification produced about The largest effects were during the Griesbachian, which repre- 290 genera, while Early Triassic articulated brachiopod diversi- sents a ‘survival phase’ for brachiopods and ammonoids fol- fication produced only about 68 genera (Table S1). Griesba- lowing the End-Permian ecological crisis. chian relict articulated brachiopods have been recovered from The patterns of recovery of brachiopods and ammonoids South China (e.g., Chen et al., 2005a, b), Salt Range (Grant, following the End-Permian mass extinction differed owing to 1970), Nepal (e.g., Waterhouse, 1978), Kashmir (Shimizu, fundamental differences in their physiology and behavior. 1981) and Oman (Chen et al., 2005a, b; Twitchett et al., 2004). Brachiopods and ammonoids are representatives of physiologi- During Olenekian time, one group of Permian relict species cally dissimilar groups of marine organisms with possible continued to exist in Nepal, while another one migrated possi- metabolic and skeletal differences (Knoll et al., 2007). bly to the Mangyshlak and Romanian areas. Brachiopods belong to a group of organisms with a calcium Localities of Induan Mesozoic-type articulated brachio- carbonate skeleton that was often massive with respect to sup- pod are known only in lower to middle-latitude regions: South porting organic tissue and formed from fluids minimally buff- China (e.g., Chen et al., 2005b; Shen and He, 1994); Northwest ered by physiology. Ammonoids, in contrast, belonged to a Caucasus (Dagys, 1974), Himalayan region (Chen et al., 2005a, group of organisms with a calcium carbonate skeleton of mod- b; Bittner, 1899), South Primorye (Dagys, 1974; this study), erate mass and formed from fluids that were relatively well and Idaho (Girty, 1927). However, the most diverse Induan buffered with respect to the factors that govern carbonate pre- brachiopod faunas were reported from South China (14 genera, cipitation. The latter condition is considered to be more sensi- including five Mesozoic-type genera) and South Primorye (six tive to high watermass PCO2 (hypercapnic stress). Mesozoic-type genera) (Fig. S10). No Induan articulated Differences in feeding behaviour may have mattered also. brachiopods have been discovered in high palaeolatitudes. Data on brachiopod sizes changes in the Permian-Triassic tran- Data of the present study show that the most diverse Ole- sition in South China (e.g., Chen et al., 2005a, b) and our addi- nekian articulated brachiopod faunas are present in the former tional evidence on the Late Permian and Early Triassic USSR area (South Primorye and Mangyshlak). These faunas brachiopods mainly of the former USSR (Tables S2–S7 and consist of 21 and 13 genera, respectively (Fig. S11). Other Fig. 7) is in an agreement with the hypothesis of Keller and known localities of Olenekian articulated brachiopods include Abramovich (2009) that high-stress conditions led to both bio- Idaho (Hoover, 1979; Perry and Chatterton, 1979; Girty, 1927), diversity loss and size reduction (‘Lilliput effect’ in Urbanek’s South China (Perry and Chatterton, 1979), Carpathians- (1993) sence). According to Harries and Knorr (2009), this Balkanian area (Jordan, 1993), Tibet (Chen et al., 2005b; Chen, effect represents a pronounced reduction in size of the biota 1983), Japan (Dagys, 1993), Spitsbergen (Dagys, 1993, 1974), associated with the aftermath of mass extinctions. However, by Alps (Jordan, 1993), and New Zealand (MacFarlan, 1992). contrast with Early Triassic brachiopods, survived the End- Among known Early Triassic articulated brachiopods, Permian mass extinction, ammonoids from the same habitats/ only a few Olenekian genera (Obnixia, Hustedtiella? (Dagys, facies in both the Primorye region and Kazakhstan do not show 1974) and Aparimirhynchia (MacFarlan, 1992)) reached high- any trends toward smaller size in the Early Triassic. These latitude areas. In contrast to Early Triassic brachiopods, Induan different outcomes may have been related to differences in (Figs. S12) and Olenekian ammonoid assemblages, according trophic resource availability (just after the End-Permian crisis, to our data, are common for both the tropical-subtropical area notably brachiopods may have suffered from a relative paucity and the high-latitude region of the Boreal realm, which is in of food resources). Epifaunal benthos (suspension-feeding agreement with previous studies on this topic (e.g., Brayard et brachiopods as well as bivalves) dominated Lower–Middle al., 2009a, b, 2006b; Tozer, 1994). Triassic marine environments, but the more modern life habits (e.g., infaunalization associated with burrowing suspension and 4.3 Palaeophysiological and Behavioural Contexts of End- deposit feeders) increased during the Late Triassic (Bonuso and Permian Environmental Stresses Bottjer, 2008). Judging by the abundance and wide distribution The End-Permian–Early Triassic interval is dominated by of nektic ammonites during the Early Triassic, one can infer unusual oceanic and climatic conditions (e.g., oceanic anoxia, that they generally were not specialized feeders, although they expansion of desert belts, declining atmospheric levels of O2 were apparently presumably bottom feeders during the Late but increasing of levels of CO2, a biocalcification crisis; e.g., Palaeozoic and Mesozoic, because optimal temperatures of Galfetti et al., 2007; Payne and Kump, 2007; Kidder and their growth are mainly comparable to those obtained from Worsley, 2004; Isozaki, 1997; Wignall and Hallam, 1993; Baud their co-occurring benthos on the shelf (e.g., Moriya et al., et al., 1989). Oceanic anoxia has been invoked as the main kill 2003; Smyshlyaeva et al., 2002). Thus, Early Triassic ammon- mechanism of the Permian-Triassic extinction event (Wignall oids had greater access to trophic resources and therefore stood and Hallam, 1993), although this hypothesis has been disputed a better chance of success than brachiopods. by Hermann et al. (2011) on the basis of the largely terrigenous Another factor in the differential recovery of these two origin of particulate organic matter in the Lower Triassic of the clades may have been related to differences in larval/juvenile Salt Range and Surghar Range. Nevertheless, chemically survival and development. After hatching, brachiopod larvae and/or physiologically harsh environmental conditions cer- and juvenile ammonoids may be dispersed with the aid of cur- tainly existed at the end of Permian, possibly caused by un- rents. All planktic brachiopod larvae are capable of delaying

12 Yuri D Zakharov and Alexander M Popov settlement until a suitable substratum has been located, al- Early Triassic may have resulted in generally more mobile though the motile larval stage usually lasts only days or hours substrates that were unfavorable for larval brachiopod settle- (Williams et al., 1997). High sedimentation rates during the ment (cf., Algeo and Twitchett, 2010). Ammonoids, in contrast,

40 42 30 (N =100) 14.2 11.8 Terebratulida

() (=11)N (N =18) 27 L 20 (N =147)

0 10 Spiriferinida

() (N =23)

L 9.4 0 33.3 30 (N =177) Mass-extinction

20 11.4

mm (N =43) () Athyritida

L 10

0 13.5 8.7 14.1 (N =32) Rhynchonellida (N =105) 10 (N =114)

()

L 0 47 (N =54) 40

()

L 20 Spiriferida

0 70

60 48.2 (N =107) 69 50 (N =64)

() 38.5

L 30 (N =16)

20 Productida

10 13.7 (N =15) 0 Lower Upper Lower Upper Griesb. Diener. Smithian Spathian Wuchiapingian Changhsingian Induan Olenekian Upper Permian 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 Pri- morye (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.

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 13 are characterised by direct ontogenetic development without a published papers on Early Triassic ammonoids and data on this larval stage and, thus, were not dependent on substrate charac- topic in press, that substantially improved this paper. This work teristics. The survival of planktic brachiopod larvae was sig- is a contribution to UNESCO-IUGS IGCP Project 572 and nificantly lower because of the short duration of their motile financially supported by the Russian grant RFBR (No. 14-05- phase, the necessity of finding a suitable substrate, and the 00011-a). specificity of their trophic needs—factors that worked to their disadvantage during the Early Triassic. The contrast in recov- REFERENCES CITED ery patterns for brachiopods and ammonoids was most evident Algeo, T., Twitchett, R., 2010. Anomalous Early Triassic Sedi- at high palaeolatitudes, most difficult apparently place for ment Fluxes due to Elevated Weathering Rates and Their brachiopod larvae migration. Biological Consequences. Geology, 38(11): 1023–1026 Balini, M., Gavrilova, V. A., Nicora, A., 2000. Biostratigraphical 5 CONCLUSIONS Revision of the Classic Lower Triassic Dolnapa Section 1. Ammonoids exceeded their pre-crisis taxonomic diver- (Mangyshlak, West Kazakhstan). Zentralblatt für Geologie sity and abundance by the late Induan (Dienerian Substage). In und Paläontologie I , 1998: 1441–1462 contrast, brachiopods never recovered their pre-crisis diversity Baud, A., Beauchamp, B., 2001. Proposals for Redefinition of and abundance anytime during the Triassic or later, in spite of the Griesbachian Substage and for the Base of the Triassic the fact that about 13 brachiopod genera survived the End- in the Arctic Regions. Proceedings of the International Permian mass extinction (recent palaeontological and carbon- Symposium “The Global Stratotype of the Permian-Triassic isotopic records allow to assume that only restricted ammonoid Boundary and the Paleozoic-Mesozoic Events”. Changxing. lineages at both the generic and the family levels survived the 26–28 mentioned mass extinction). Baud, A., Magaritz, M., Holser, W. T., 1989. Permian-Triassic of 2. The differential recovery patterns of brachiopods and the Tethys: Carbon Isotope Studies. Geologische Rund- ammonoids seem to be a result of physiological and behav- schau, 78(2): 649–677 ioural differences, reflected in their different life modes and Bittner, A., 1899. Himalayan Fossils. Trias Brachiopoda and distributions in ecospace. Specifically, brachiopods were at a Lamellibranchiata. Palaeontologica Indica, 15(2): 1–76 disadvantage during the Early Triassic because of the short Bjerager, M., Seidler, L., Stemmerik, L., et al., 2006. Ammonoid duration of their motile phases, the need to find a suitable sub- Stratigraphy and Sedimentary Evolution across the strate for larval settlement, and the specificity of their trophic Permian-Triassic Boundary in East Greenland. Geological requirements. Magasine, 143(5): 635–656 3. Brachiopod’s ‘lilliput effect’ developed during the Bogoslovskaya, M. F., Kuzina, L. F., Leonova, T. B., 1999. Clas- Early Triassic, judging from published data and partly con- sification and Distribution of Late Paleozoic Ammonoids. firmed by some original evidences from South Primorye, Man- In: Rozanov, A. Y., Shevyrev, A. A., eds., Fossil Cephalo- gyshlak, North Caucasus and Transcaucasia, may have been a pods: Recent Advances in Their Study. Izdatelstvo response to a relative paucity of food resources for this specific Akademii Nauk SSSR, Moscow. 89–124 (in Russian) benthic organisms of that time. Bonuso, N., Bottjer, D. J., 2008. A Test of Biogeographical, En- 4. Diversification of Early Triassic brachiopods at high vironmental, and Ecological Effect on Middle and Late Tri- latitudes was quite restricted: no articulated brachiopods of assic Brachiopod and Bivalve Abundance Patterns. Palaios, Induan age and only three genera of Olenekian age (Obnixia 23(1): 43–54 and Hustedtiella? (Dagys, 1974) and Aparimirhynchia Brayard, A., Bucher, H., 2008. Smithian (Early Triassic) Am- (MacFarlan, 1992)) have been discovered in the Boreal realm. monoid Faunas from Northwestern Guangxi (South China): However, immediately after the End-Permian extinction event Taxonomy and Biochronology. Fossils and Strata, 56: 1– (and possibly just during it), ammonoids colonized and stably 179 inhabited many regions in the Boreal realm throughout the Brayard, A., Bucher, H., Brühwiler, T., et al., 2007a. Prohar- Triassic. The contrast in recovery patterns for brachiopods and poceras Chao: A New Ammonoid Lineage Surviving the ammonoids was most evident in the high palaeolatitudes, ap- End-Permian Mass Extinction. Lethaia, 40(2): 175–181 parently most difficult place for brachiopod larvae migrations. Brayard, A., Escarguel, G., Bucher, H., 2007b. The Biogeogra- phy of Early Triassic Ammonoid Faunas: Clusters, Gradi- ACKNOWLEDGMENTS ents, and Networks. Geobios, 40(6): 749–765 For help in finding some important Early Triassic Brayard, A., Bucher, H., Escarguel, G., et al., 2006a. The Early brachiopod and ammonoid fossils in South Primorye and regu- Triassic Ammonoid Recovery: Palaeoclimatic Significance lar assistance, we are indebted to Y Shigeta (National Museum of Diversity Gradients. Palaeogeography, Palaeoclimatol- of Nature and Science, Tsukuba), H Maeda, and T Kumagae ogy, Palaeoecology, 239(3–4): 374–395 (Kyoto University) and also to V T S’’edin (Institute of Pacific Brayard, A., Escarguel, G., Bucher, H., 2006b. The Biogeogra- Oceanology, Vladivostok). Our cordial thanks are due to Prof. phy of Early Triassic Ammonoid Faunas: Clusters, Gradi- T A Algeo (University of Cincinnati) for providing valuable ents, and Networks. Geobios, 40: 749–765 editorial comments, Prof. Shuzhong Shen (Nanjing Institute of Brayard, A., Escarguel, G., Bucher, H., et al., 2009a. Smithian Geology and Palaeontology) and anonymous reviewer for and Spathian (Early Triassic) Ammonoid Assemblages from stimulating remarks, as well as for their information on latest Terranes: Paleoceanographic and Paleogeographic Implica-

14 Yuri D Zakharov and Alexander M Popov

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APPENDIX: Original Data on Brachiopod and Ammonoid Remains from the Lower Triassic of South Primorye and Man- gyshlak 1 SOUTH PRIMORYE 1.1.2 Ammonoid occurrences The first geological studies in Russian southern Far East Early Triassic ammonoids are significantly more diverse in (Ussuri region, later named as Primorye) were made by V. P. the mentioned section, as well as in other South Primorye sections, Margaritov, a graduate from the St.-Peterburg University, arriving than associated brachiopods. Induan ammonoids from the Lazur- in Vladivostok in 1880 as a teacher of math. On the western coast naya Bay Formation, exposed on the western coast of Ussuri Gulf, of the Ussuri Gulf, near Shamara Bay (now Lazurnaya) he dis- are represented by a single genus (Tompophiceras), belonging to covered some fossils, including ammonoids and inarticulated the Palaeozoic-type family Dzhulfitidae, and five genera within brachiopods. His collection fell into the hands of the President of the Mesozoic-type families Ophiceratidae, Proptychitidae, Mee- the Russian Academy of Sciences, A. P. Karpinsky, a recognized koceratidae, and Xenoceltitidae?. Early Olenekian (Smithian) expert in Late Palaeozoic ammonoids, who distinguished Early ammonoids from the uppermost portion of the Lazurnaya Bay Triassic ceratitid ammonoids among collected fossils. Later, Early Formation (Ussuriflemingites abrekensis (=“Gyronites separa- and Middle Triassic marine deposits in the environs of Vladi- tus”) beds of the “Hedenstroemia” bosphorensis Zone) and the vostok were studied by D. L. Ivanov, the chief of a geological lower portion of the Tobizin Cape Formation (Euflemingites pry- team making reconnaissance work for the construction of the nadai beds of the mentioned zone) in this section are more di- Trans-Siberian railroad. He collected mollusc remains in the area verse, representing by 22 genera within Mesozoic-type families of the Shamara Bay and articulated brachiopod and mollusc shells Sageceratidae, Hedenstroemiidae, Aspenitidae, Ussuriidae, on Russian Island. On the initiative of A. P. Karpinsky, collec- Paranannitidae, Clypeoceratidae, Arctoceratidae, Meekoceratidae, tions V. P. Margaritov and D. L. Ivanov were forwarded to Aus- Dieneroceratidae, and Flemingitidae (Fig. S1). trian palaeontologists K. Diener and A. Bittner, who identified and described them (Bittner, 1899b; Diener, 1895). Later, Triassic 1.2 Abrek Bay cephalopods were investigated by Kiparisova (1961), Burij and The Abrek Bay Section, located at Strelok Strait and repre- Zharnikova (1981, 1962), Zakharov (1997a, 1978, 1968), sented by the Lazurnaya Bay and Zhitkov Cape formations, is Zharnikova (1985, 1972), Shigeta et al. (2009), Smyshlyaeva and worthy of consideration as one of the reference sections for Zakharov (2011), and Smyshlyaeva (2010). However, after Bitt- Lower Triassic stratigraphy in South Primorye (Shigeta et al., ner’s investigation, only a few Early Triassic brachiopod taxa, 2009; Markevich and Zakharov, 2004; Zakharov et al., 2002; based on material from South Primorye, was described by Dagys Zakharov and Popov, 1999; Kiparisova, 1972, 1961; Burij, 1959). (1974, 1965) and Shigeta et al. (2009). Observed ranges of ammonoids and brachiopods across the 1.2.1 Brachiopod occurrences Induan-Lower Anisian interval in South Primorye (Seryj-Tri In the lower part of the Induan Lazurnaya Bay Formation of Kamnya, Abrek, Konechnyj, Tobizin, Ayax-Balka, Zhitkov, the Abrek Bay Section only inarticulated brachiopods (Lingula Golyj, and Paris), summarized in Figs. S1–S9 and Table S1, help borealis Bittner and Orbiculoidea sp.) have been discovered to understand the main features of their recovery on specific level (Shigeta et al., 2009). Among late Induan brachiopods from the from the End-Permian extinction. Lytophiceras-bearing beds of the Gyronites subdharmus Zone about five genera of articulated brachiopods, including Abrekia, 1.1 Seryj-Tri Kamnya Capes have been recognized (Shigeta et al., 2009; Dagys, 1974). All of The Seryj Cape-Tri Kamnya Cape Section, exposed at the them belong to Mesozoic-type families (Wellerellidae, Pontisii- western Ussuri Gulf, is represented by the Lazurnaya and Tobizin dae, Rhynchonellidae, and Norellidae). Representatives of the formations (Zakharov et al., 2010; Markevich and Zakharov, Abrekia (A. sulcata Dagys) have been discovered also in the 2004; Zakharov, 1997b, 1996, 1968; Kiparisova, 1972, 1961; Lower Olenekian (Smithian) portion of the Zhitkov Cape Forma- Burij, 1959; Korzh, 1959; Diener, 1895). tion (“Hedenstroemia” bosphorensis Zone, presumably Euflem- ingites prynadai beds) (Fig. S2). 1.1.1 Brachiopod occurrences Lissorhynchia sp. (Pontiidae), the first record of this genus 1.2.2 Ammonoid occurrences from South Primorye, seems to be the oldest Induan articulated The Induan ammonoid succession in the Abrek Bay Section brachiopod species found in the region. A single representative of consists of a single genus (Tompophiceras), belonging to a this species was collected by T. Kumagae in the lower part of the Permian-type family Dhulfitidae, and 12 genera within five Lazurnaya Bay Formation (Gyronites subdharmus Zone, about Mesozoic-type families (Proptychitidae, Ophiceratidae, Meeko- 20 m above the top of the underlying early Induan Tompo- ceratidae, Paranoritidae, and ?Paranannitidae). Among Smithian phiceras ussuriense Zone). Other Induan brachiopod fossils from ammonoids of the mentioned section 27 genera belonging to 15 this section are represented by Lingula borealis Bittner. A few Mesozoic-type families have been recognized (Fig. S2). They are articulated brachiopods, mainly rhynchonellids, were collected in following: Sageceratidae, Paranannitidae, Aspenitidae, Xenocelti- the lower part of the Smithian Tobizin Formation (Euflemingites tidae, Proptychitidae?, Clypeoceratidae, Arctoceratidae, Oweniti- prynadai beds) (Fig. S1). dae, Inyoitidae?, Prionitidae, Kashmiritidae, Meekoceratidae, Dieneroceratidae, Flemingitidae, and Palaeophyllitidae.

18 Yuri D Zakharov and Alexander M Popov

1.3 Konechnyj Cape Zakharov, 2004). The Konechnyj Cape, located at Novik Bay on Russian Is- At Konechnuj Cape, only a small portion, about 6 m thick, of land, is represented by the Schmidt Formation (Markevich and 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. Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 19

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 known. The articulated brachiopods (Zeilleriidae? gen. and sp. were recently collected there. The two brachiopod assemblages nov. E and Spiriferinidae gen. and sp. indet.) (Fig. S3) have been have been recognized in the section: (1) Hustedtiella planicosta- collected only from the upper part of the middle Olenekian Spiriferinida-Rhynchonellida-“Fletcherithyris” margaritovi- (early Spathian) Schmidt Cape Formation. Zeilleriidae? below (occurs mainly in blocks) and (2) monospecies-“Fletcherithyris” margaritovi assemblage above. 1.4.2 Ammonoid occurrences Both the assemblages correspond to the Tirolites ussuriensis beds Only early to middle Olenekian (Smithian-early Spathian) of the early Spathian Tirolites-Amphistephanites Zone in other ammonoid succession, represented by 20 genera belonging to 14 sections at the Russian Island (Markevich and Zakharov, 2004). Mesozoic-type families is known there. These families are fol- Representatives of five genera within five Mesozoic-type families lowing: Aspenitidae, Ussuriidae, Paranannitidae, Owenitidae, (Rhynchonellidae, Neoretziidae, Spiriferinidae, Dielasmatidae, and Meekoceratidae, Dieneroceratidae, Prionitidae, Tirolitidae, Me- Zeilleriidae?) were discovered in this locality. lagathiceratidae, Stephanitidae, Dinaritidae, Clypeoceratidae, Arctoceratidae, and Flemingitidae) (Fig. S3). 1.4 Tobizin Cape The Tobizin Cape Formation, exposed on the south part of 1.5 Ayax Bay-Balka Cape Russian Island (Novyj Dzhigit Bay area) is represented by the The Ayax Bay-Balka Cape, exposed on the northern part of Lazurnaya Bay (basal part), Tobizin Cape and Schmidt Cape Russian Island, consists of Lazurnaya Bay, Tobizin Cape and formations (Markevich and Zakharov, 2004; Buryi, 1979; Zak- Schmidt Cape formations (Markevich and Zakharov, 2004; Zak- harov, 1968; Burij, 1959; Korzh, 1959; Diener, 1895). harov, 1997a, 1978, 1968; Buryi, 1979; Kiparisova, 1972, 1961; Korzh, 1959; Diener, 1895). 1.4.1 Brachiopod occurrences From the early Olenekian (Smithian) Tobizin Cape Forma- 1.5.1 Brachiopod occurrences tion only inarticulated brachiopod Lingula borealis Bittner is Only inarticulated brachiopod Lingula borealis Bittner 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 Lazurnaya Cape Formation only bad preserved Gyronites? sp. shells were collected from the lower part (Induan-Smithian) of (Meekoceratidae) has been recognized. Among early to middle the section, which consists of the Lazurnaya Bay and Tobizin Olenekian ammonoids from the uppermost portion of the Cape formations. Articulated brachiopods Lepismatina aff. Lazurnaya Bay Formation, Tobizin Cape and Schmidt Cape mansfieldi (Girty) and spiriferinids belonging to Mesozoic- formations 22 ammonoid genera belonging to 17 Mesozoic- type families Lepismatinidae and Spiriferinidae have been type families have been listed (Zakharov, 1997a, b). Among reported from the lower part of the middle Olenekian (early them are following: Sageceratidae, Hedenstroemiidae, Ussurii- Spathian) Schmidt Cape Formation (Fig. S4). dae, Paranannitidae, Arctoceratidae, Meekoceratidae, Dienero- ceratidae, Owenitidae, Inyotidae, Prionitidae, Xenoceltitidae, 1.5.2 Ammonoid occurrences Tirolitidae, Melagathiceratidae, Stephanitidae, Doricranitidae?, At the top of the Induan conglomerate member of the Dinaritidae, and Palaeophyllitidae (Fig. S4). 22 Yuri D Zakharov and Alexander M Popov

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 S5.

1.6 Schmidt Cape-Tchernyschev Bay Spiriferinidae, Neoretziidae, Dielasmatidae, and Zeilleriidae (?). The Schmidt Cape-Tchernyschev Bay, located on the Earliest Anisian spiriferinid and terebratulid brachiopods are south-eastern part of Russian Island, is represented by the reported from the Ussuriphyllites amurensis Zone of the Schmidt Cape and Zhitkov Cape formations (Markevich and Tchernyschev Bay area (Fig. S5). Zakharov, 2004; Buryi, 1979; Zakharov, 1978, 1968; Kiparisova, 1972, 1961; Burij, 1959; Korzh, 1959). 1.6.2 Ammonoid occurrences Spathian ammonoids from the Schmidt and Zhitkov Cape 1.6.1 Brachiopod occurrences formations of this section are represented by 21 genera belong- Schmidt Cape area is one of the places with abundantearly ing to 12 Mesozoic-type families (Sageceratidae, Paranannitidae, Spathian Mesozoic-type articulated brachiopods (mainly in the Khvalynitidae, Meekoceratidae, Prionitidae, Columbitidae, Ka- Tirolites ussuriensis beds of the Tirolites-Amphistephanites Zone) zakhstanitidae, Xenoceltitidae, Tirolitidae, Dinaritidae, and Pa- (Fig. S5). In this part of the mentioned section, there are five laeophyllitidae) (Fig. S5). genera belonging to the following Mesozoic-type families: 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.. ?Sub- fengshanites multiformis; reg. substage. regional substage. Other designations as in Figs. S1 and S5. 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 (Fig. S6). They are represented by 24 genera belonging to 13 The Zhitkov Cape Section, exposed on the northern part of families (Sageceratidae, Paranannitidae, Khvalynitidae, Iscu- Russian Island, is represented by the Tobizin Cape, Zhitkov litidae, Meekoceratidae, Dieneroceratidae, Columbitidae, Ka- Cape and Karazin Cape formations (Markevich and Zakharov, zakhstanitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, 2004; Zakharov, 1997a, 1978, 1968; Buryi, 1979; Kiparisova, Dinaritidae, and Megaphyllitidae). 1972, 1961; Burij, 1959; Korzh, 1959). 1.8 Golyj (Kom-Pikho-Sakho) Cape 1.7.1 Brachiopod occurrences The Golyj Cape Section, exposed at eastern part of the Majority of articulated brachiopods from this section was Ussuri Gulf, consists of Lazurnaya Bay, Zhitkov Cape and collected in the Tirolites ussuriensis beds of the early Spathian Karazin formations (Markevich and Zakharov, 2004; Buryi, Tirolites-Amphistephanites Zone (Schmidt Formation), others 1979; Zakharov, 1968; Burij, 1959; Korzh, 1959). were collected in the late Spathian Subfengshanites multiformis Zone (Fig. S6). Olenekian portion of the Zhitkov Cape Section 1.8.1 Brachiopod occurrences includes seven genera belonging to six Mesozoic-type families Articulated brachiopods in this section were discovered (Rhynchonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae, only in the middle Olenekian (early Spathian) portion of the Dielasmatidae, and Zeilleriidae?). Zhitkov Cape Formation (Tirolites-Amphistephanites Zone). They are represented by five genera of four Mesozoic-type 1.7.2 Ammonoid occurrences families (Rhynchonellidae, Neoretziidae, Spiriferinidae, and Smithian and Spathian ammonods of this section were Zeilleriidae?) (Fig. S7). collected in the upper portion of the Tobizin Cape Formation (Anasibirites nevolini Zone) and Schmidt Cape (Tirolites- 1.8.2 Ammonoid occurrences Amphistephanites Zone), and Zhitkov Cape (Neocolumbites Representatives of a single genus Gyronites (Mesozoic- insignis and Subfengshanites multiformis zones) formations type family Meekoceratidae) were reported from the Induan 26 Yuri D Zakharov and Alexander M Popov

Lazurnaya Bay Formation (Fig. S7). Early to late Olenekian 1.9.1 Brachiopod occurrences ammonoids were discovered from the lower (“Hedenstroemia” Articulated brachiopods from the Paris Bay Section were bosphorensis Zone), middle (Tirolites-Amphistephanites Zone) collected from the early Spathian Schmidt Formation and upper (Neocolumbites insignis Zone) portions of the Zhit- (Tirolites-Amphistephanites Zone) and uppermost beds of the kov Cape Formation. Olenekian ammonoid assemblage of the late Spathian Zhitkov Formation (Subfengshanites multiformis Golyj Cape section consists of 24 genera belonging to 17 Zone). Spathian assemblage of articulated brachiopods consists Mesozoic-type families: Sageceratidae, Hedenstroemiidae, of six genera belonging to five Mesozoic-type families: Rhyn- Aspenitidae, Ussuriidae, Paranannitidae, Khvalynitidae, Arcto- chonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae, and ceratidae, Owenitidae, Meekoceratidae, Inyoitidae, Prionitidae, Zeilleriidae? (Fig. S8). Columbitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Melagathiceratidae, and Flemingitidae. 1.9.2 Ammonoid occurrences Ammonoids from the Paris Bay Section are originated from 1.9 Paris Bay the upper portion of the Smithian Tobizin Formation (the upper The Paris Bay Section, located on the northern part of part of the Anasibirites nevolini Zone), early Spathian Schmidt Russian Island, is represented by the Tobizin Cape, Schmidt Formation (Tirolites-Amphistephanites Zone), and late Spathian Cape, Zhitkov Cape and Karazin Cape formations (Markevich Zhitkov Formation (Neocolumbites insignis and Subfengshanites and Zakharov, 2004; Buryi, 1979; Zakharov, 1968; Burij, 1959; multiformis zones). Olenekian (mainly Spathian) ammonoids Korzh, 1959). 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, Man- gyshlak. 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. 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 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, Dienero- 3 DISCUSSION ceratidae, Columbitidae, Xenoceltitidae, Keyserlingitidae, Tiro- The Induan of South Primorye (SP) is characterised by litidae, Stephanitidae, Dinaritidae, and Hungaritidae (Fig. S8). seven articulated brachiopod species: early Induan Lissorhynchia sp. and six late Induan species, including Abrekia sulcata and 2 MANGYSHLAK, DOLNAPA (KAZAKHSTAN) new ones (Figs. S1 and S2 and Table S1). Early Olenekian The first Triassic fossils in Mangyshlak were found in 1914 brachiopods from SP show a slight reduction in species number, by M. V. Bayarunas, who identified and described some ammon- largely due to a variety of facial conditions. Middle Olenekian oids among them 22 years later (Bajarunas, 1936). Later, Early brachiopods from SP (about eight species, including new ones) Triassic of Mangyshlak were investigated by Astachova (1960), (Figs. S3–S9 and Table S1) are more diverse than those of Man- Shevyrev (1968), Gavrilova (2007, 1989, 1980), Balini et al. gyshlak. In addition, information on late Olenekian brachiopods (2000) and Zakharov et al. (2008). Before our investigation, Tri- from SP (about five species, including new ones) and Man- assic brachiopods from Mangyshlak were described by Dagys gyshlak (12 species), as well as the published data on Olenekian (1974) on the bases of A. A Shevyrev’s collection. species from Eurasia (e.g., Chen et al, 2005; Xu and Grant, 1994; The reference section for the Lower Triassic in Mangyshlak, Chen, 1983; Xu and Liu, 1983) and North America (e.g., Perry located at the Dolnapa Well area in Kara-Tau, is represented by and Chatterton, 1979; Girty, 1927), illustrates a general trend in the Tartalinskaya, Karadzhatykskaya and Karaduanskaya forma- marked rising of taxonomic diversity of Mesozoic-type brachio- tions. pods from the Lower Induan through the Upper Olenekian. The Induan, lower, middle and upper Olenekian in SP are character- 2.1 Brachiopod Occurrences ised by 20, 11 and 30 species, respectively. Similar changes in Spathian brachiopod assemblage of the mentioned section ammonoid succession, with highest diversification for the early consists of 8–11 genera belonging to eight Mesozoic-type fami- Olenekian, occur in some other Tethyan regions. According to lies (Pontisiidae, Norellidae, Diplospirellidae, Neoretziidae, Le- published data (Chen et al., 2005) 13 articulated brachiopod line- pismatinidae, Dielasmatidae, Antezeilleridae, and Plectoconchi- ages at familial level survived the End-Permian mass extinction. dae) (Fig. S9). As a result, at least 12 Palaeozoic type articulated brachiopod genera occur in the Lower Triassic, mainly in the Lower Griesba- 2.2 Ammonoid Occurrences chian; at the same time the Permian-type Prelissorhynchia at Spathian ammonoids are represented by 23 genera belonging Mangyshlak was found by us in the first time in the upper Ole- to 14 Mesozoic-type families (Sageceratidae, Khvalynitidae, nekian. Dinaritidae, Prionitidae, Columbitidae, Kazakhstanitidae, Kash- However, only four ammonoid lineages at family- miritidae, Xenoceltitidae, Olenikitidae, Tirolitidae, Doricranitidae, superfamily level survived the mentioned mass extinction; appar- Dinaritidae, Palaeophyllitidae, and Megaphyllitidae (Fig. S9). ently only a single Palaeozoic-type ammonoid genus (Episage- ceras) is known from the Lower Triassic (Zakharov, 1978). Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 29

Table S1 Late Permian (Changhsingian) and Early Triassic brachiopods and ammonoids

Number of the Changhsingian articulated brachiopod genera has been calculated by Chen et al. (2005); genera survived the Permian-Triassic boundary are indicated by asterisk; Early Triassic brachiopods from the Abrek Section are illustrated in Shigeta et al. (2009, Figs. 146–148 therein).

Table S1-1 Early Induan (Griesbachian) articulated brachiopod genera

No. Genus Reference No. Genus Reference 1 Acosarina* Cooper & Grant Chen et al., 2005 9 Paryphella* Liao Chen et al., 2005

2 Araxathyris* Grunt Chen et al., 2005 10 Prelissorhynchia* Chen et al., 2005 Xu & Grant 3 Cathaysia* Ching Chen et al., 2005 11 Pustula* Thomas Chen et al., 2005

4 Laevorhynchia Shen & He Shen and He, 1994; 12 ?Retimarginifera* Chen et al., 2005 Mu et al., 2007 Waterhouse 5 Orbicoelia* Chen et al., 2005 13 Spinomarginifera* Chen et al., 2005 Waterhouse & Piyasin Huang 6 Crurithyris* George Yang et al., 1987; 14 “Spiriferina” D’Orbigny Chen et al., 2005 Shen et al., 2006 7 Paracrurithyris* Liao Chen et al., 2005 15 Spirigerella* Waagen Chen et al., 2005

8 Paraspiriferina* Reed Chen et al., 2005 16 Tethyochonetes* Chen et al., 2005 Chen & others

Table S1-2 Late Induan (Dienerian) articulated brachiopod genera

No. Genus Reference No. Genus Reference 1 Abrekia Dagys Shigeta et al., 2009 7 Piarorhynchella? Shigeta et al., 2009 (Figs. 147.1–147-24 therein) Dagys (Figs. 147.29–147.32 therein)

2 Lepismatina Wang Chen et al., 2005 8 Pontisiinae gen. and Shigeta et al., 2009 sp. nov. Figs. 147.41–147.48 therein)

3 Lissorhynchia Dagys, 1974; this study 9 Retzioidea gen. and Chen et al., 2005 Yang & Xu (Figs. 3-1 and 3-2) sp. indet. (=Neowellerella)

4 ?Maorirhynchia MacFarlan, 1992 10 “Spiriferina” Xu and Grant, 1994 MacFarlan D’Orbigny

5 “Mentzelia” Girty, 1927 11 Wellerellidae gen. Shigeta et al., 2009 and sp. nov. A

6 Periallus Hoover Hoover, 1979 12 Wellerellidae gen. Shigeta et al., 2009 and sp. nov. B

30 Yuri D Zakharov and Alexander M Popov

Table S1-3 Olenekian (Smithian and Spathian) articulated brachiopod genera

No. Genus Reference No. Genus Reference 1 Abrekia Dagys Shigeta et al., 2009 23 ?Retimarginifera* Chen et al., 2005 (Figs. 147. 1–147.24) Waterhouse 2 Antezeilleria Xu and Liu, 1983 24 Rhaetina Waagen Hoover, 1979; Dagys, 1974 Xu & Liu 3 Aparimarhynchia MacFarlan, 1992 25 Rhynchonellidae gen. This study (Figs. 3-7–3-10) MacFarlan indet. 4 “Fletcherithyris” Dagys, 1974; this study 26 Rhynchonellidae gen. This study (Figs. 3-11–3-14) nov. A 5 Hustedtiella Dagys Dagys, 1974; this study 27 Rhynchonellidae gen. This study (Figs. 3-23–3-26) nov. 1 6 Lepismatina Wang Dagys, 1974; this study 28 Rhynchonellidae gen. This study (Figs. 3-15–3-18) (Figs. 3-31–3-34) nov. 2 7 Lepismatinidae This study 29 Rhynchonellidae gen. This study gen. and sp. nov. nov. 3 8 Lissorhynchia Original data 30 Rhynchonellidae gen. This study Yang & Xu (Figs. 3-19–3-22) indet. 9 Neoretzia Dagys Chen, 1983 31 “Rhynchonella” Chen et al., 2005 Fischer de Waldheim 10 Nudirostralina Chen, 1983 32 Spirigerellina Dagys Dagys, 1974; this study Yang & Xu (Figs. 3-27–3-30) 11 ?Sinuplicorhynchia This study 33 Spiriferinidae gen. This study Dagys nov. A (Figs. 3-35–3-38) 12 Spirigerellinae This study 34 Spiriferinidae gen. This study gen. and sp. nov. nov. B 13 Obnixia Hoover Hoover, 1979 35 Zeilleriidae gen. nov. This study A 14 Orbicoelia* Chen et al., 2005 36 Zeilleriidae gen. nov. This study Waterhouse & B Piyasin 15 Paranorellina Dagys, 1974 37 Zeilleriidae gen. nov. This study Dagys C

16 Periallus Hoover Chen et al., 2005 38 Zeilleriidae gen. nov. This study D

17 Portneufia Hoover Hoover, 1979 39 Zeilleriidae gen. nov. This study E

18 Piarorhynchella Dagys, 1974 40 Zeilleriidae gen. nov. This study Dagys F

19 Prelissorhynchia Chen et al., 2005; this study 41 Thyratryaria Xu & Xu and Liu, 1983 (?)* Xu & Grant (Figs. 3-3–3-6) Liu

20 Proanadyrella Xu and Liu, 1983 42 Vex Hoover Hoover, 1979 Xu & Liu

21 ?Proanadyrella This study 43 Zeilleriidae gen. This study Xu & Liu indet.

22 Protogusarella Perry and Chatterton, 1979

Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 31

Table S1-4 Changhsingian ammonoid genera

No. Genus Reference No. Genus Reference 1 Abichites Shevyrev Bogoslovskaya et al., 1999 22 Phisonites Shevyrev Bogoslovskaya et al., 1999 2 Avushoceras Bogoslovskaya et al., 1999 23 Pleuronodoceras Bogoslovskaya et al., 1999 Ruzhencev Chao & Liang 3 Changhsingoceras Zhou et al., 1999 24 Pseudogastrioceras Bogoslovskaya et al., 1999 Chao & Liang Spath 4 ?Cyclolobus Waagen Zhou et al., 1999 25 Pseudostephanites Zhou et al., 1999 Chao & Liang 5 Dushanoceras Zhao, Bogoslovskaya et al., 1999 26 Pseudotirolites Sun Bogoslovskaya et al., 1999 Liang & Zheng 6 Dzhulfites Shevyrev Bogoslovskaya et al., 1999 27 Pseudotoceras Bogoslovskaya et al., 1999 Ruzhencev 7 Dzhulfoceras Bogoslovskaya et al., 1999 28 Qinjiangoceras Zhou et al., 1999 Ruzhencev Zhao, Liang & Zheng 8 Episageceras Bogoslovskaya et al., 1999 29 Rongjiangoceras Zhou et al., 1999 Noetling Zhao, Liang & Zheng 9 “Hypophiceras” Shevyrev, 2006; 30 ?Rotaraxoceras Zakharov et al., 2005; Trümpy Yin and Zhang, 1996 Ruzhencev Kotlyar et al., 1983 (=xenodiscid? ammonoid) 10 Huananoceras Bogoslovskaya et al., 1999 31 Rotodiscoceras Bogoslovskaya et al., 1999 Chao & Liang Chao & Liang 11 Iranites Teichert & Bogoslovskaya et al., 1999 32 Schizoloboceras Bogoslovskaya et al., 1999 Kummel Zhao, Liang & Zheng 12 Julfoceras Bogoslovskaya et al., 1999 33 Shevyrevites Bogoslovskaya et al., 1999 Ruzhencev Teichert & Kummel 13 Liuchengoceras Zhou et al., 1999 34 Sinoceltites Chao & Bogoslovskaya et al., 1999 Zhao, Liang & Zheng Liang 14 Longmenshanoceras Zhou et al., 1999 35 Stacheoceras Zhou et al., 1999 Zhao, Liang & Zheng Gemmellaro 15 Meitianoceras Zheng Zhou et al., 1999 36 Sutchanites Zakharov and Oleinikov, Zakharov 1994 16 Mingyuexiaceras Zhou et al., 1999 37 Tapashanites Bogoslovskaya et al., 1999 Zhao, Liang & Zheng Chao & Liang 17 Neoaganides Bogoslovskaya et al., 1999 38 “Tompophiceras” Shevyrev, 2006; Yin and Plummer & Scott Popow (=dzhulfitid? Zhang, 1996 ammonoid) 18 “?Otoceras” Gries- Shevyrev, 2006; 39 Trigonogastrites Zhou et al., 1999 bach (=araxoceratid? Yin and Zhang, 1996 Zhao, Liang & ammonoid) Zheng 19 Pachydiscoceras Zhou et al., 1999 40 ?Vedioceras Zakharov et al., 2005; Chao & Liang Ruzhencev Kotlyar et al., 1983 20 Paratirolites Bogoslovskaya et al., 1999 41 Xenodiscus Waagen Bogoslovskaya et al., 1999 Stoyanow 21 Pentagonoceras Zhou et al., 1999 Zhao, Liang & Zheng

Wuchiapingian-Changhsingian boundary in Transcaucasia and Iran is indicated following Zakharov et al. (2005)—inside of the Vedioceras ventrosulcatum of the Dzhulfian. 32 Yuri D Zakharov and Alexander M Popov

Table S1-5 Early Induan (Griesbachian) ammonoid genera

No. Genus Reference No. Genus Reference 1 Aldanoceras Dagys & Ermakova, 2002 9 Metophiceras Spath Shevyrev, 2000 Ermakova

2 Anotoceras Hyatt Shevyrev, 1999 10 Otoceras Griesbach Tozer, 1994; Shevyrev, 1986 (=Metotoceras Spath)

3 Bernhardites Shevyrev Shevyrev, 1999; 11 Ophiceras Griesbach Tozer, 1994 (=?Shevyrevoceras Waterhouse, 1994 Waterhouse)

4 Bukkenites Tozer Tozer, 1994 12 Proptychites Waagen Waterhouse, 1994; this study (=Pseudoproptychites Bando)

5 Discophiceras Shevyrev, 2000 13 Shalshalia Water- Waterhouse, 1994 Spath house

6 Episageceras* Noetling Waterhouse, 1994 14 Tompophiceras Shevyrev, 1999; Popow Dagys and Ermakova, 1996 (=Zakharovites Waterhouse)

7 Hypophiceras Trümpy Shevyrev, 2000 15 Vishnuites Diener Waterhouse, 1994 8 Lytophiceras Spath Waterhouse, 1994 16 Wordieoceras Tozer Tozer, 1994

Table S1-6 Late Induan (Dienerian) ammonoid genera

No. Genus Reference No. Genus Reference 1 Ambites Waagen Waterhouse, 1994 31 Meekophiceras Tozer Tozer, 1994 2 ?Anaxenaspis Zakharov et al., 2010; 32 Mesokantoa Waterhouse, 1994 Kiparisova Ermakova, 2002 Waterhouse

3 Anotoceras Hyatt Waterhouse, 1994 33 Metophiceras Spath Waterhouse, 1994; this (=?Altoconchella study Waterhouse)

4 Aspiditella Strand Shevyrev, 1986 34 Ophiceras Griesbach Waterhouse, 1994 5 Aspitella Waterhouse Waterhouse, 1996a 35 ?Mullericeras Ware, Ware et al., 2011 Jenks, Hautmann & Bucher

6 Bernhardites Waterhouse, 1994; this study 36 Pachyproptychites Shigeta et al., 2009 Shevyrev Diener (=?Shevyrevoceras Waterhouse)

7 Bukkenites Tozer Tozer, 1994 37 ?Parahedenstroemia Ware et al., 2011 Spath

8 Clypeoceras Smith Spath, 1934; this study 38 Paranorites Waagen Shevyrev, 1986 (=?Aspitella Water- house) 9 Clypites Waagen Brühwiller et al., 2010b 39 Paraspidites Spath Shevyrev, 1986 10 Collignonites Bando Shevyrev, 1986 40 Pleuroambites Tozer Tozer, 1994 11 Dunedinites Tozer Tozer, 1994 41 Pleurogyronites Tozer Tozer, 1994 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 33

Continued

No. Genus Reference No. Genus Reference 12 Eoptychites Spath Brayard et al., 2006a, 42 ?Preflorianites Spath Shevyrev, 1986 (=Waagenoproptychites b; this study Waterhouse)

13 Eovavilovites Ermakova Ermakova, 2002 43 Prejuvenites Shevyrev, 1999; Waterhouse Waterhouse, 1996a

14 Episageceras* Popow Zakharov, 1978 44 Prionolobus Waagen Brühwiler et al., 2008; (=?Lilangia this study Waterhouse)

15 Gyrolecanites Spath Shevyrev, 1986 45 Proptychites Waagen Tozer, 1994; this study (=Pseudoproptychites Bando)

16 Gyronites Waagen Waterhouse, 1994; 46 ?Pseudohedenstroemia Shevyrev, 1986 (=?Nangykangia Waterhouse) this study Kummel

17 Gyrophiceras Spath Mu et al., 2007 47 Pseudosageceras Waterhouse, 1994 Diener

18 ?Hedenstroemia Waagen Zakharov et al., 2009 48 Pseudovishnuites Mu et al., 2007 Zakharov & Mu,

19 Heibergites Tozer Tozer, 1994 49 Radioceras Waterhouse Waterhouse, 1996a 20 Himoceras Waterhouse Waterhouse, 1996a 50 Sakhaites Vozin Ermakova, 2002 21 Discophiceras Spath Waterhouse, 1994; 51 Shangganites Brühwiler et al., 2008 (=?Himophiceras Waterhouse) this study Brühwiler, Brayard, Bucher & Guodun

22 Hubeitoceras Waterhouse Waterhouse, 1994; 52 Subinyoites Spath Shevyrev, 1986 Brühwiler et al., 2008

23 Jieshanites Brühwiler, Brayard, Brühwiler et al., 2008 53 Tompoproptychites Brayard et al., 2006a, b; Bucher & Guodun Vavilov & Zakharov Vavilov and Zakharov, 1976 24 Khangsaria Waterhouse Waterhouse, 1996a 54 Ussuriceras Spath Shevyrev, 1986 25 Kingites Waagen Shevyrev, 1986 55 Vavilovites Tozer Tozer, 1994; this study (=Fuchsites Waterhouse) 26 Koninckites Waagen Bruhwiler et al., 56 Vercherites Brühwiler, Brühwiler, et al., 2010b 2010b Ware, Bucher, Krystyn & Goudemand 27 Kummelia Waterhouse Waterhouse, 1996a; 57 Vishnuites Diener Waterhouse, 1994 (=?Tilihonia Water- this study house; ?Zhaojinkeoceras Waterhouse; ?Mesokantoa Waterhouse) 28 Kymatites Waagen Waterhouse, 1994; 58 Wordieoceras Spath Tozer, 1994 (=?Khangsaria Waterhouse) this study

29 Lilastroemia Waterhouse Waterhouse, 1996a 59 ?Xenoceltites Spath Shevyrev, 1986 30 Lytophiceras Spath Waterhouse, 1994 60 Xenodiscoides Spath Brühwiler et al., 2008

Ammonoid assemblage from the upper Chegaji Member (top 10–30 cm) in Nepal, investigated by Waterhouse (1994), yielding Gy- ronites, seems to be Dienerian in age.

34 Yuri D Zakharov and Alexander M Popov

Table S1-7 Early Olenekian (Smithian) ammonoid genera

No. Genus Reference No. Genus Reference 1 Abrekites Shigeta & Shigeta et al., 2009 69 Metinyoites Spath Shevyrev, 1986 Zakharov

2 Ambites Waagen Shevyrev, 1986 70 Metussuria Spath Zakharov et al., 2010 3 Ambitoides Shigeta & Shigeta et al., 2009 71 Mianwaliites Brühwiler et al., 2012b Zakharov Brühwiler et al.

4 Anaflemingites Brayard and Bucher, 2008 72 Monneticeras Brühwiler et al., 2012b Kummel & Steele Brühwiler et al.

5 Anahedenstroemia Shevyrev, 1986 73 Mudiceras Brühwiler et al., 2010b Hyatt Brühwiler et al.

6 Anakashmirites Spath Shevyrev, 1986 74 Mullericeras Ware, Ware et al., 2011; this study Jenks, Hautmann & Bucher

7 Anasibirites Zakharov et al., 2010 75 Nammalites Brühwiler et al., 2012b Mojsisovics Brühwiler, Bucher & Goudemand

8 Anastephanites Spath Shevyrev, 1986 76 Nanningites Brayard and Bucher, 2008 Brayard & Bucher

9 Anawasatchites Brayard et al., 2006a, b 77 Nilgiria Waterhouse Waterhouse, 1996b McLearn

10 Anaxenaspis Kiparisova Zakharov et al., 2010 78 ?Nordophiceras Waterhouse, 1996b Popow

11 Arctoceras Hyatt Shigeta et al., 2009 79 Nuezelia Brühwiler, Brühwiler et al., 2012a Bucher, Krystyn & Breta

12 Arctoprionites Spath Shevyrev, 1986 80 Nyalamites Brühwiler et al., 2010a Brühwiler, Bucher & Goudemand

13 Aspenites Brayard and Bucher, 2008 81 Owenites Zakharov et al., 2010 Hyatt & Smith Hyatt & Smith

14 Balhaeceras Shigeta et al., 2009 82 Palaeokazakhsta- Markevich and Zakharov, Shigeta & Zakharov nites Zakharov 2004

15 Boewanites Waterhouse Waterhouse, 1996b 83 Parahedenstroemia Shevyrev, 1986 Spath

16 Brayardites Brühwiler et al., 2010a 84 Parakymatites Shevyrev, 1986 Brühwiler, Bucher & Waagen Goudemand

17 Burijites Zakharov Zakharov, 1978 85 Paranannites Brayard and Bucher, 2008 Hyatt & Smith 18 Churkites Okuneva Markevich and Zakharov, 86 Paranorites Waagen Shevyrev, 1986 2004 19 Colchenoceras Waterhouse, 1996b 87 Paraspedites Spath Brayard et al., 2006a, b Waterhouse 20 Cordillerites Hyatt & Brayard and Bucher, 2008 88 Parastephanites Brayard et al., 2006a, b Smith Hyatt Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 35

Continued

No. Genus Reference No. Genus Reference 21 Clypeoceras Smith Brühwiler et al., 2012b; 89 Parussuria Spath Zakharov et al., 2010 Waagen, 1895 22 Clypites Waagen Brayard et al., 2006a, b 90 Pijaconcha Water- Waterhouse, 1996b house 23 Clypeoceratoides Ermakova, 2002 91 Preflorianites Spath Brayard and Bucher, 2008 Dagys & Ermakova

24 Dieneroceras Brayard and Bucher, 2008 92 Prionites Waagen Brühwiler et al., 2010a Spath 25 Dorikranites Hyatt Shevyrev, 1968 93 Prionolobus Waagen Brayard et al., 2006a 26 Dunedinites Tozer Tozer, 1994 94 Procurvoceratites Brayard and Bucher, 2008 Brayard & Bucher 27 Eoptychites Spath Waterhouse, 1996b 95 Proharpoceras Chao Brayard and Bucher, 2008 28 Eoptychitoides Brühwiler et al., 2010c; 96 Prosphingitoides Shevyrev, 1995 Waterhouse Waterhouse, 1996b Shevyrev 29 Epihedenstroemia Shevyrev, 1986 97 Protophiceras Hyatt Brayard and Bucher, 2008 Spath 30 Escargueltites Brühwiler et al., 2012a 98 Pseudoceltites Hyatt Brühwiler et al., 2010a 31 Eukashmirites Shevyrev, 1986 99 Pseudoaspenites Brayard and Bucher, 2008 Kummel Spath 32 Euflemingites Tozer, 1994 100 Pseudaspidites Spath Brayard and Bucher, 2008 Spath 33 Flemingites Brayard and Bucher, 2008 101 Pseudoflemingites Brayard and Bucher, 2008 Waagen Spath 34 Glyptophiceras Brühwiler et al., 2010a 102 Pseudo- Shevyrev, 1986 Spath hedenstroemia Kummel 35 Galfettites Brayard and Bucher, 2008 103 Pseudosageceras Brayard and Bucher, 2008 Brayard & Bucher Diener 36 Guangxiceltites Brayard and Bucher, 2008 104 Pseudokazakhstanites Markevich and Zakharov, Brayard & Bucher Zakharov 2004 37 Guangxiceras Brayard and Bucher, 2008 105 Punjabites Brüchwiler et al., 2012b Brayard & Bucher Brüchwiler et al.. 38 Guodunites Brayard and Bucher, 2008 106 ?Radioceras Brüchwiler et al., 2010b Brayard & Bucher Waterhouse 39 Gurleyites Shevyrev, 1986 107 Radioprionites Shigeta et al., 2009 Mathews Shigeta & Zakharov 40 Gyronites Waagen Tozer, 1994 108 Rohillites Brüchwiler et al., 2012b; Waterhouse Waterhouse, 1996b 41 Hanielites Welter Brayard and Bucher, 2008 109 Sakhaites Vozin Ermakova, 2002 42 Hebeisenites Brayard and Bucher, 2008 110 Shamaraites Shigeta et al., 2009 Brayard & Bucher Shigeta & Zakharov 43 Hedenstroemia Brayard and Bucher, 2008 111 Shigetaceras Brühwiler et al., 2010a Waagen Brühwiler, Bucher & Goudemand 44 Hemiaspenites Brayard et al., 2006a, b 112 Steckites Brühwiler, Brühwiler et al., 2012a Kummel & Steele Bucher & Krystyn 36 Yuri D Zakharov and Alexander M Popov

Continued

No. Genus Reference No. Genus Reference 45 Hemiprionites Brayard and Bucher, 2008 113 Stephanites Waagen Brühwiler et al., 2010a Spath 46 Hochuliites Brühwiler et al., 2012b 114 Subalbanites Zakharov, 1978 Brüchwiler et al. Zakharov 47 Inyoites Brayard and Bucher, 2008 115 Subinyoites Spath Brüchwiler et al., 2012b; Hyatt & Smith Shevyrev, 1986 48 Jinyaceras Brayard and Bucher, 2008 116 Subflemingites Spath Brühwiler et al., 2010a Brayard & Bucher 49 Jolinkia Waterhouse, 1996b 117 Submeekoceras Spath Brayard and Bucher, 2008 Waterhouse 50 Juvenites Smith Brayard and Bucher, 2008 118 Subvishnuites Spath Brayard and Bucher, 2008 51 Kashmirites Brayard and Bucher, 2008 119 Tellerites Mojsisovics Brayard et al., 2006a, b Diener 52 Kashmiritidae gen. Brühwiler et al., 2010b 120 Thermalites Smith Brayard et al., 2006a, b nov. A 53 Kazakhstanites Shevyrev, 1995 121 Truempyceras Brühwiler et al., 2012a Shevyrev Brühwiler et al. 54 Kelteroceras Ermakova, 2002 122 Tulongites Brühwiler, Brühwiler et al., 2010a Ermakova Bucher & Goudemand 55 Kingites Waagen Brühwiler et al., 2010c 123 Tuyangites Zhao Shevyrev, 1986 56 Koiloceras Brühwiler et al., 2012b 124 Urdyceras Brayard & Brayard and Bucher, 2008 Brühwiler et al. Bucher 57 Koninckites Shevyrev, 1986 125 Ussuria Diener Zakharov et al., 2010 Waagen 58 Kraffticeras Brühwiler et al., 2012a 126 Ussuridiscus Shigeta & Shigeta et al., 2009 Brühwiler, Zakharov Bucher & Krystyn

59 Kymatites Waagen Waterhouse, 1996a 127 Ussuriflemingites Shigeta et al., 2009 Shigeta & Zakharov 60 Lanceolites Hyatt Brayard and Bucher, 2008 128 Ussurijuvenites Smyshlyaeva and & Smith Smyshlyaeva & Zakharov Zakharov, 2011 61 Larenites Brayard and Bucher, 2008 129 Vercherites Brühwiler, Brühwiler et al., 2010b Brayard and Ware, Bucher, Krystyn & Bucher Goudemand 62 Latisageceras Waterhouse, 1996b 130 Wasatchites Mathews Tozer, 1994 Ruzhencev 63 Lepiskites Ermakova, 2002 131 Wailiceras Brayard & Brayard and Bucher, 2008 Dagys & Ermakova Bucher 64 Leyeceras Brayard and Bucher, 2008 132 Weitschaticeras Brayard & Brayard and Bucher, 2008 Brayard & Bucher Bucher 65 Lingyunites Chao Brayard and Bucher, 2008 134 Xenoceltites Spath Brühwiler et al., 2010a 66 Meekoceras Hyatt Brühwiler et al., 2012b 135 Xenodiscoides Spath Brühwiler et al., 2011b 67 Melagathiceras Tozer, 1994 136 Xiaoqiaoceras Brühwiler et al., 2010c; Tozer Brayard & Bucher Brayard and Bucher, 2008

68 Mesohedenstro- Brayard and Bucher, 2008 emia Chao Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 37

Table S1-8 Late Olenekian (Spathian) ammonoid genera

No. Genus Reference No. Genus Reference 1 Albanites Arthaber Shevyrev, 1986 66 Neomeekoceras Tozer Tozer, 1994 2 Anakashmirites Spath Shevyrev, 1986 67 Nordophiceras Popow Zakharov, 1978 3 Anaxenaspis Kiparisova Zakharov et al., 2010 68 Nordophiceratoides Guex et al., 2010 4 Arctomeekoceras Popow Zakharov, 1978 69 Nuetzelia Brühwiler, Brühwiler et al., 2012a Bucher & Krysyn 5 Arctoprionites Spath Zakharov, 1978 70 Olenekoceras Dagys & Ermakova, 2002 Ermakova 6 Arianites Arthaber Shevyrev, 1986 71 Olenikites Hyatt Zakharov, 1978 7 Arnautoceltites Diener Markevich and 72 Keyserlingites Hyatt Zakharov, 1978 Zakharov, 2004 8 Bajarunia Dagys Ermakova, 2002 73 Palaeophyllites Welter Guex et al., 2010 9 Balkanites Ganev Brayard et al., 2006a, b 74 Mianwallites Brühwiler et al., 2012b Brühwiler et al. 10 Bandoites Zakharov Markevich and 75 Nyalamites Brühwiler, Brühwiler et al., 2010a Zakharov, 2004 Bucher & Goudemand 11 Beatites Arthaber Brayard et al., 2006a, b 76 Pallasites Renz & Renz Brayard et al., 2006a, b

12 Beneckeia Mojsisovics Shevyrev, 1986 77 Paradinarites Zhao Shevyrev, 1986 13 Beyrichites Waagen Shevyrev, 1986 78 Paragoceras Arthaber Guex et al., 2010 14 Boreoceras Dagys & Guex et al., 2010; Er- 79 Parahedenstroemia This study Ermakova makova, 2002; Spath (=Rudolftruempyiceras this study Guex et al.) 15 Boreomeekoceras Popow Ermakova, 2002 80 Paranannites Shevyrev, 1986 Hyatt & Smith 16 Burijites Zakharov Zakharov, 1978 81 Paranorites Waagen Brayard et al., 2006a, b

17 Carteria Guex et al. Guex et al., 2010 82 Paranoritoides Guex Shevyrev, 1986 18 Ceccaisculitoides Guex et al. Guex et al., 2010 83 Parasibirites Popow Zakharov, 1978 19 Chioceras Renz & Renz Shevyrev, 1986 84 Parussuria Spath Zakharov et al., 2010 20 Chiotites Renz & Renz Shevyrev, 1986 85 Popovites Tozer Tozer, 1994 21 Columbites Hyatt & Smith Zakharov, 1978 86 Columbitidae gen. nov. Brayard et al., 2006a, b 22 Cordillerites Hyatt & Smith Guex et al., 2010 87 Praesibirites Ermakova, 2002 Dagys & Ermakova 23 Coscaites Guex et al. Guex et al., 2010 88 Preflorianites Spath Shevyrev, 1986 24 Cowboyiceras Guex et al., 2010 89 Prefloarianitoides Brayard et al., 2006a, b Guex et al. Wang 25 Dagnoceras Athaber Shevyrev, 1986 90 Prenkites Arthaber Zakharov, 1978 26 Dalmatites Kittl Shevyrev, 1986 91 Procarnites Arthaber Zakharov et al., 2010 27 Diaplococeras Hyatt Brayard et al., 2006a, b 92 Procolumbites Markevich and Zakharov, Astachova 2004 28 Dieneroceras Spath Zakharov et al., 2010 93 Prohungarites Spath Guex et al., 2010 29 Digitophyllites Zhao Brayard et al., 2006a, b 94 Prosphingites Zakharov, 1978 Mojsisovics 30 Dinarites Mojsisovics Shevyrev, 1986 95 Protropites Arthaber Shevyrev, 1986 31 Doaboceras Boulin, Bouyx, Brayard et al., 2006a, b 96 Proptychitoides Spath Brayard et al., 2006a, b Termier & Termier 32 Dorikranites Hyatt Shevyrev, 1986 97 Pseudoaspidites Spath Shevyrev, 1986 33 Eodanubites Wang Brayard et al., 2006a, b 98 Pseudodinarites Hyatt Brayard et al., 2006a, b

34 Eogymnites Spath Shevyrev, 1986 99 Pseudoceltites Hyatt Shevyrev, 1986 38 Yuri D Zakharov and Alexander M Popov

Continued

No. Genus Reference No. Genus Reference 35 Eophyllites Spath Shevyrev, 1986 100 Pseudoharpoceras Brayard et al., 2006a, b Waagen 36 Epiceltites Arthaber Shevyrev, 1986 101 Pseudokymatites Spath Brayard et al., 2006a, b 37 Epiceltitoides Guex Guex, 1978 102 Pseudoprosphingites Markevich and Zakharov, Shevyrev 2004 38 Epiceltitoides Guex Guex, 1978 103 Pseudosageceras Zakharov et al., 2002 Diener 39 Eschericerafites Guex et al. Guex et al., 2010 104 Pseudodinarites Hyatt Brayard et al., 2006a, b 40 Evenites Dagys & Ermakova Ermakova, 2002 105 Pseudosvalbardiceras Ermakova, 2002 Dagys & Ermakova 41 Fengshanites Zhao Brayard et al., 2006a, b 106 Silberlingeria Guex et al., 2010 Guex et al. 42 Gaudemerites Guex et al. Guex et al., 2010 107 Stacheites Kittl Shevyrev, 1986 43 Glabercolumbites Guex et al. Guex et al., 2010 108 Subcolumbites Spath Shevyrev, 1986 44 Hellenites Renz & Renz Zakharov, 1978 109 Subfengshanites Zakharov et al., 2008 Zakharov 45 Hemilecanites Spath Brayard et al., 2006a, b 110 Subinyoites Spath Brayard et al., 2006a, b 46 Hyrcanites Shevyrev Shevyrev, 1986 111 Subolenekites Zakharov Zakharov, 1978 47 Idahocolumbites Guex et al. Guex et al., 2010 112 Subvishnuites Spath Shevyrev, 1986 48 Isculitoides Spath Brayard et al., 2006a, b 113 Sulioticeras Tozer Tozer, 1994 49 Ivesgalleticeras Guex et al. Guex et al., 2010 114 Svalbardiceras Frebold Zakharov, 1978 50 Jeanbesseiceras Guex et al. Guex et al., 2010 115 Tapponnierites Guex et al., 2010 Guex et al. 51 Kazakhstanites Shevyrev Shevyrev, 1968 116 Tardicolumbites Guex et al., 2010 Guex et al. 52 Keyserlingites Hyatt Zakharov, 1978 117 Tchernyschevites Markevich and Zakharov, Zakharov 2004 53 Kiparisovites Astachova Shevyrev, 1986 118 Timoceras Dagys & Ermakova, 2002 Ermakova 54 Koninckitoides Dagys & Ermakova, 2002 119 Tirolites Mojsisovics Shevyrev, 1986 Ermakova 55 Khvalynites Shevyrev Shevyrev, 1986 120 Tjururpites Shevyrev Shevyrev, 1986 56 Leiophyllites Diener Shevyrev, 1986 121 Tozericeras Guex Guex, 1978 57 Mangyshlakites Shevyrev Shevyrev, 1986 122 Tunglanites Zhao Shevyrev, 1986 58 Marcouxia Guex et al. Guex et al., 2010 123 Ussuriidae gen. nov. Guex et al., 2010; Kummel, (=“Ussurites Hyatt”) 1969; this study 59 Meropella Renz & Renz Brayard et al., 2006a, b 124 Vickohlerites Kummel Brayard et al., 2006a, b 60 Metadagnoceras Tozer Tozer, 1994 125 Xenoceltites Spath Brayard and Bucher, 2008 61 Metahedenstroemia Spath Shevyrev, 1986 126 ?Xenodiscoides Spath Brayard et al., 2006a, b 62 Metasageceras Renz & Renz Brayard et al., 2006a, b 127 Zenoites Renz & Renz Brayard and Bucher, 2006a 63 Metinyoites Spath Waterhouse, 1996b 128 Zhitkovites Zakharov Markevich and Zakharov, 2004 64 Monacanthites Tozer Tozer, 1994 129 Ziyunites Wang Brayard et al., 2006a, b 65 Neocolumbites Zakharov Zakharov, 1978

Following Hyatt and Smith’s (1905) original proposal, we continue to consider that the Neopopanoceras haugi Zone is earliest Ani- sian, in spite of the recent tendency to expect it to be latest Olenekian (Shevyrev, 2006; Tozer, 1994; Bucher, 1989). Ammonoid genera Deweveria, Neopopanoceras, Goricanites, Subhungarites, Pseudacrochordiceras, Inyoceras, and Courtilloticeras, known from of the Neopopanoceras haugi Zone of the American tropical-subtropical realm (Guex et al., 2010; Bucher, 1989), have not been in- cluded in Table S1, because this zone seems to be earliest Anisian in age (in spite of such a fact that these ammonoids are associated with Keyserlingites and Pseudosvalbardiceras, common for the uppermost Olenekian in the Boreal realm (Zakharov et al., 2008). Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis 39

Table S2 Measurements for brachiopod length selected for analysis of miniaturization: Order Productida

Stage Substage, formation, Length (mm) Region, reference/ zone Max. Min. Mean collection authority Induan Griesbachian 11 1.5 4.2 South China (N=9); Liao, 1980 Griesbachian 13.7 6.3 8.84 South China (N=15); Chen et al., 2006 Upper Changh- Urushten Fm. 69.0 2.0 18.4 Urushten Reef, North Caucasus singian (N=64); Licharev, 1937 Lower Changh- Nikitin Fm. 38.5 7.0 22.7 Nikitino, North Caucasus (N=16); singian Licharev, 1937 Lower Araxoceras 37.1 7.2 12.7 Dorasham, Transcaucasia (N=97); Wuchapingian latissimum B.V. Koczyrkevicz’s coll. Pseudodunbarula 48.2 6.5 14.7 Dorasham, Transcaucasia (N=107); arpaensis-Araxilevis B.V. Koczyrkevicz’s coll. intermedius N. number of specimens used for our observation

Table S3 Measurements for brachiopod length selected for test of miniaturization: Order Spiriferida

Stage Zone Length (mm) Region, reference/ Max. Min. Mean collection authority Lower Araxoceras latissimum 47.0 4.6 22.4 Dorasham, Transcaucasia (N=54); Wuchiapingian B.V. Koczyrkevicz’s coll. Lower Araxoceras latissimum and 42.0 8.0 25.6 Dorasham, Transcaucasia (N=15); Wuchiapingian Pseudodunbarula arpaensis- B.V. Koczyrkevicz’ coll. Araxilevis intermedius

Disignations as in Table S2.

Table S4 Measurements for brachiopod length selected for analysis of miniaturization: Order Rhynchonellida (e.g., Wellerella, Abrekia, Piarorhynchella, and Lissorhynchia)

Stage Substage, forma- Length (mm) Region, reference/ tion, zone Max. Min. Mean collection authority Olenekian Spathian 10.1 3.5 7.3 Dolnapa, Mangyshlak (N=57); Y. D. Zakharov’s coll. Spathian 8.7 3.3 6.17 Paris Bay, Primorye (N=105); A. M. Popov’s coll. Smithian 9.2 5.5 6.86 Shamara Bay, Primorye (N=53); A. M. Popov’s coll. Upper Induan Dienerian 7.8 3.5 6.2 Abrek Bay, Primorye (N=41); A. M. Popov’s coll. Upper Urushten Fm. 13.5 5.3 8.58 Urushten Reef, North Caucasus; Changhsingian B. V. Koczyrkevicz”s coll. Upper Wuchiap- Vedioceras 14.1 3.5 7.57 Dorasham and Ogbin, Transcaucasia ingian ventrosulcatum (N=114); B. V. Koczyrkevicz’s coll. Lower Wuchiap- Araxoceras latis- 8.5 3.8 5.91 Dorasham and Ogbin, Transcaucasia ingian simum (N=71); B. V. Koczyrkevicz’s coll.

Disignations as in Table S2

40 Yuri D Zakharov and Alexander M Popov

Table S5 Measurements for brachiopod length selected for analysis of miniaturization: Order Athyridida (e.g., Araxathyris, Spirigerellina, and Hustedtiella)

Stage Substage, zone Length (mm) Region, reference/ Max. Min. Mean collection authority Olenekian Spathian 12.1 3.4 8.2 Dolnapa, Mangyshlak (N=14); Y. D. Zakharov’s coll. Spathian 11.4 4 8.1 Paris Bay and Schmidt Cape, Primorye (N=43); A. M. Popov’s coll. Lower Wuchiap- Araxoceras 33.3 4.9 15.0 Dorasham, Transcaucasia (N=177); ingian latissimum B. V. Koczyrkevicz’s coll.

Disignations as in Table S2.

Table S6 Measurements for brachiopod length selected for test of miniaturization: Order Spiriferinida (Lepismaina)

Stage Substage Length (mm) Region, refence/ Max. Min. Mean collection authority Olenekian Spathian 9.4 3.7 5.9 Dolnapa, Mangyshlak (N=23); Y. D. Zakharov’s coll. Spathian 12.2 5.7 8.47 Paris Bay and Schmidt Cape, Primorye (N=15); A. M. Popov’s coll.

Disignations as in Table S2.

Table S7 Measurements for brachiopod length selected for test of miniaturization: Order Terebratulida (e.g., Notothyris and Gefonia)

Stage Substage, zone Length, mm Region, reference/ (beds) Max. Min. Mean collection authority Olenekian Spathian 27 6.9 18.7 Shmidt Bay, Primorye (N=147); A. M. Popov’s coll. 28 1.8 8.9 Western USA (N=46); Hoover, 1979 Upper Changh- Urushten Fm. 42 4.3 10.9 Urushten Reef, North Caucasus singian (N=100); Likharew, 1937 Upper Vedioceras 11.8 2.5 6.96 Dorasham and Ogbin, Transcaucasia Wuchiapingian ventrosulcatum (N=18); B. V. Koczyrkevicz’s coll. Lower Araxoceras 12.7 5.6 8.4 Dorasham and Ogbin, Transcaucasia Wuchiapingian latissimum (N=4); B. V. Koczyrkevicz’s coll. Pseudodunbarula 14.2 7.4 10.5 Dorasham and Ogbin, Transcaucasia arpaensis- (N=11); B. V. Koczyrkevicz’s coll. Araxilevis intermedius Disignations as in Table S2.

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