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 Recovery of Brachiopod and Ammonoid Faunas Following the End-Permian Crisis: Additional Evidence from the Lower Triassic of the Russian Far East and Kazakhstan Yuri D Zakharov*, Alexander M Popov Far Eastern Geological Institute, Far Eastern Branch, Russian Academy of Sciences, Stoletiya Prospect 159, Vladivostok 690022, Russia ABSTRACT: After the End-Permian mass extinction, ammonoids reached levels of taxonomic diver- sity 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 set- tlement. 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 Razdol naia ° 60° 80° N 80° N 60° 44 N ’ ()a USSURIYSK Rakovka Rakovka ° hka ac 40 N Komarovkani Iv 2 Barsukovka ° Kaimanovka 40 N aia 1 Perevozn Study area 1 000 km Borisovka Razdol'noye Razdol'naya 70° 90° 110° E a Zavodskoy k Bol. Kiparisovka ov yom rt A Artyom Vol’no-Nadezhdinskoe Zanadvorovka Steklyanukha Tavrichanka Uglovoe Smolyaninovo Bogataya Sukhodol Vladivostok Bol’shoi Kamen’ 1 G Amur Gulf 5 Ussuri ulf (b) 6 43° 43° 3 8 Popov Isl. Russian Island Fokino 4 2 Reyneke Isl. 7 Dunai Sea of Japan Putyatin Isl. 0 15 km Askol’d Isl. 132°E Figure 1. (a) Location of study areas in the former USSR. 1. South Primorye and 2. Mangyshlak. (b) Study sections in South 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