Recovery of brachiopod and ammonoid faunas following the endPermian crisis: additional evidence from the Lower Triassic of the
Russian Far East and Kazakhstan
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Yuri D. Zakharov*, Alexander M. Popov
Far Eastern Geological Institute, Far Eastern Branch, Russian Academy of Sciences, Stoletiya Prospect 159,
Vladivostok, 690022 Russia
Abstract
The present paper provides information on recovery of brachiopod and ammonoid faunas following endPermian crisis, based on original evidence from the southern Russian Far East and Kazakhstan (Middle Asia) as well
as the published data on Eurasia and North America. After the end-Permian mass extinction, ammonoids reached
levels of taxonomic diversity higher than in the Changhsingian by the Dienerian substage of the Induan (early Early
Triassic). However, brachiopods exhibit a prolonged delay in recovery, and their taxonomic diversity had not
recovered to Late Permian levels even by the Olenekian (late Early Triassic). 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 (‘Lilliput
effect’) 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
vulnerabilities associated with their larval stage: brachiopod larvae had a short-duration motile stage of development
during which they had to locate a suitable substrate for settlement. In contrast, ammonoids had no larval stage and
juveniles may have been dispersed widely into favorable habitats. These factors may account for differences in the
relative success of ammonoids and brachiopods at high-latitude regions following the end-Permian mass extinction:
ammonoids successfully recolonized the Boreal region during the Early Triassic whereas brachiopods did not.
Keywords: Lower Triassic; Former USSR; Brachiopods; Ammonoids; Biotic recovery.
*
Corresponding author. Far Eastern Geological Institute, Russian Academy of Sciences (Far Eastern Branch).
Prospect Stoletiya 159, Vladivostok, 690022 Russia. Tel.: (4232) 317567; fax: (4232) 317847.
E-mail address: yurizakh@mail.ru (Y.D. Zakharov)
1. Introduction
The Permian-Triassic boundary is associated with the largest extinction event of
Phanerozoic time (e.g., Erwin, 1994; Shen and Shi, 2002; Shen et al., 2006; Yin and Zhang,
1996). Early Triassic successions of both benthonic and nektonic marine organisms (e.g.,
brachiopods and ammonoids) contain important information about the postextinction biotic
recovery interval. However, in spite of significant progress in the past decades in investigation of
Early Triassic brachiopods (e.g., Dagys, 1974, 1993; Feng and Jiang, 1978; Hoover, 1979; Perry
and Chatterton, 1979; Sun et al., 1981; Chen, 1983; Xu and Liu, 1983; MacFarlan, 1992; Jordan,
1993; Xu and Grant, 1994; Shen and He, 1994; Shen and Shi, 1996; Shen and Shi, 1996; Chen et
al., 2002, 2005a,b; 2006, 2009; Ruban, 2006a,b, 2009; Shigeta et al., 2009) and ammonoids (e.g.,
Shevyrev, 1986, 2006; Zakharov, 1974, 1977, 1978, 1983, 1996, 1997; Guex, 1978, 2010;
Bando, 1980; Gavrilova, 1980, 2007; Korchinskaya and Vavilov, 1987; Dagys and Ermakova,
1988, 1996; Tozer, 1994; Krystyn and Orchard, 1996; Ermakova, 2002; Krystyn et al., 2003;
Brayard et al., 2006, 2007a,b, 2009a,b,c; Brayard and Bucher, 2008; Mu et al., 2007; Brühwiler
et al., 2008, 2010a,b,c, 2011a,b; Ware et al., 2011; Smyshlyaeva and Zakharov, 2011), our
knowledge of these important invertebrate groups, especially Mesozoic-type Brachiopoda,
remains incomplete. Early Triassic (mainly Olenekian) ammonoids are abundant and diverse in
many regions of the world, but the most diverse Olenekian brachiopod faunas have been
discovered in the former USSR area (South Primorye and Mangyshlak). The goal of this work is
to analyze taxonomic diversity patterns of Early Triassic articulated brachiopods and ammonoids
from the former USSR (Fig. 1, see the Power Point) in order to reconstruct biotic recovery
patterns after the end-Permian mass extinction.
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2. Materials
Original materials on Early Triassic invertebrates (brachiopods and ammonoids) used for
our investigation were obtained by Y.D. Zakharov and A.M. Popov from Induan and Olenekian
sections of South Primorye (Seryj-Tri Kamnya, Abrek, Konechnyj, Tobizin, Ayax-Balka,
Schmidt-Tchernyschev, Zhitkov, Golyj, and Paris) during the last five years. Investigation of
Olenekian brachiopods from Mangyshlak (Dolnapa), Kazakhstan was made on the basis of Y.D.
Zakharov’s collection from his Kara-Tau expedition in 1978.
Systematic determinations of brachiopods and ammonoids follow classifications
established by Carter and Johnson (2006) for Spiriferinida, Lee et al. (2006) for Terebratulida,
Savage et al. (2002) for Rhynchonellida, Alvarez and Rong (2002) for Athyridida,
Bogoslovskaya et al. (1999) for Prolecanitida and Goniatitida, and Zakharov (1978), Shevyrev
(1986), Tozer (1994); Waterhouse (1994, 1996a,b), Brayard et al. (2006a,b), Brayard and Bucher
(2008), Shigetabet al. (2009), Brühwiler et al. (2010a,b, 2011a,b), Guex et al. (2010), and Ware
et al. (2011) for Ceratitida.
3. Lower Triassic ammonoid and brachiopod biostratigraphy
3.1. South Primorye (Russian Far East)
The Early Triassic epiplatformal terrigenous assemblage of South Primorye (Fig. 1) lies
unconformably on the Upper Palaeozoic and more ancient marine and continental sedimentary
and volcano-sedimentary deposits, volcanics and granitoids (Zakharov, 1968, 1978; Zakharov et
al., 2010). In many areas of South Primorye (e.g., Russian Island, Ussuri and Amur gulfs, Abrek
Bay and Artyom area), the bulk of the Induan or Induan to earliest Olenekian Lazurnaya Bay
Formation consists of shallow-marine shelf sediments. These are predominantly conglomerate
and sandstone with lenses of sandy limestone-coquina dominated by bivalves. The lower part of
the Induan is represented by the Tompophiceras ussuriense Zone, its upper part corresponds to
the Gyronites subdharmus Zone (Figs. S1 and S2; see figures in the Power Point and textAppendix).
The Tobizin Cape Formation, of early Olenekian (Smithian) age, is present on Russian
Island and on the western coast of Ussuri Gulf (between the Seryj and Tri Kamnya capes), where
it conformably overlies the Lazurnaya Bay Formation and corresponds to the lower part of the
Ayaxian regional substage. It is represented by sandstone with lenses of coquinoid calcareous
sandstone with mainly cephalopod and bivalve remains. On Russian Island, the following zones
can be recognized: “Hedenstroemia” bosphorensis below and Anasibirites nevolini above (Figs.
S3 and S4).
The Schmidt Cape Formation, of middle Olenekian (early Spathian) age, corresponds to
the upper part of the Ayaxian regional substage. It is exposed only on Russian Island and
consists of sandstone with thick lenses of sandy limestone-coquina (mainly brachiopod-bivalve
one). It is represented by a single zone, the Tirolites-Amphistephanites Zone (Figs. S3 and S6).
The Zhitkov Cape Formation, of Early Olenekian to Late Olenekian or (locally) only Late
Olenekian age, represents deeper shelf facies characterised by the dominance of mudstone and
siltstone with numerous calcareous-marly nodules and lenses and thin, irregular beds of
sandstone. The lower part of the formation, which is found in the West Ussuri Gulf-Strelok Strait
area (Figs. S1 and S7), is represented by the “Hedenstroemia” bosphorensis and Anasibirites
nevolini zones and is early Olenekian (Smithian) in age. The upper part of the formation was
formed in this area during the middle (Tirolites-Amphistephanites Zone) and late (Neocolumbites
insignis and Subfengshanites multiformis zones) Olenekian. On Russian Island (Figs. S5-S7) and
the western coast of Amur Gulf (Zakharov et al., 2005), only the upper Olenekian Zhitkov Cape
Formation is present However, the Olenekoceras-bearing part of the Olenekian on the western
coast of Amur Gulf is characterised mainly by fine-grained, striped sandstone (Atlasov Cape
Formation).
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The Karazin Cape Formation represents the Anisian everywhere in South Primorye. No
benthic forms of allochthonous origin have been found in this formation and therefore it seems to
have accumulated under offshore environmental conditions. It composed mainly of fucoid
sandstone with large calcareous septarian nodules. The Lower Anisian in South Primorye is
represented by the Ussuriphyllites amurensis and Leiophyllites pradyumna zones (Figs. S5-S7).
The detailed lithological succession and distribution of brachiopod and ammonoid
species in the Lower Triassic of South Primorye are shown in some figures (Figs. S1 and S8 see Online Supplement). Figure 2 illustrates the stratigraphic distribution of brachiopods and
ammonoids at the family and order levels (within the Upper Changhsingian-Upper Olenekian
interval).
3.1.1. Tompophiceras ussuriense Zone (lower Induan)
The ammonoid assemblage of the Lower Induan (Griesbachian) Tompophiceras
ussuriense Zone (at least 18 m thick) is characterised by the presence of two ceratitid ammonoid
genera: Tompophiceras (T. ussuriense (Zakharov)), occurred together with inarticulated
brachiopods (Lingula sp.) and some bivalves, and Lythophiceras (Lythophiceras sp.), the latter
occurring in the upper part of the zone. For detailed ammonoid distribution data, see Fig. S1.
However, no Griesbachian articulated brachiopods were found in the Far East apparently
because of their extreme rarity there. A remarkable feature of the Griesbachian ceratitid
ammonoid assemblage from South Primorye is that Tompophiceras is a representative of the
Palaeozoic-type family (Dzhulfitidae) (Fig. 2). However, Lythophiceras belongs to a Mesozoictype family (Ophiceratidae).
3.1.2. Gyronites subdharmus Zone (upper Induan)
The Upper Induan (Dienerian) Gyronites subdharmus Zone (45-110 m thick) is marked
by the first appearance datum of the ceratitid ammonoid genus Gyronites (G. subdharmus
Kiparisova), used here for recognition of the base of the Dienerian. Other ceratitid ammonoid
genera in the lower part of the zone are Lythophiceras (L. eusakuntala Zakharov), Proptychites
(P. hiemalis (Diener)) and possibly Tompophiceras (Tompophiceras sp.), in the upper part –
Ussuridiscus (U. varaha (Diener)), Wordieoceras (W. cf. wordiei (Spath)), Dunedinites (D.
magnumbilicatus (Kiparisova)), Melagathiceratidae (Melagathiceratidae gen. et sp. indet.), and
Pachiproptychites (P. otoceratoides (Diener)). All these ceratitid ammonoids of the Gyronites
subdharmus Zone, with the exception of the Dzhulfitidae (Tompophiceras) and Proptychitidae
(e.g., Proptychites and Pachiproptychites), are representatives of typical Mesozoic-type families
(Ophiceratidae, Meekoceratidae, Melagathiceratidae, and possibly Paranannitidae). The
Palaeozoic-type Dzhulfitidae was discussed earlier, but the ancestral group of the Proptychitidae
is unknown at present (Fig. 2).
The articulated brachiopod assemblage of the Dienerian Gyronites subdharmus Zone is
characterised by the presence of four families of the order Rhynchonellida (Figs. 2 and 3):
Pontisiidae (Lissorhynchia sp., Pontisiinae gen. and sp. nov.), Wellerellidae (Wellerellidae gen.
and sp. nov. A, Wellerellidae gen. and sp. nov. B), Norellidae (Piarorhynchella? sp. nov.) and
Rhynchonellidae (Abrekia sulcata Dagys, Rhynchonellidae gen. and sp. nov.). For detailed
biostratigraphic distribution data, see Online Supplement (Figs. S1 and S2). Among these
brachiopod families, the Pontisiidae and the Wellerellidae are Palaeozoic-type ones, but the
Norellidae and the Rhynchonellidae seem to have originated during the Early Triassic (Induan).
Newly found Dienerian brachiopod shells in South Primorye, including specimens of
Abrekia, first described by A.S. Dagys (see Online Supplement), are generally very small. Their
maximal shell length is only 10 to 11 mm. In addition, some Late Permian rhynchonellid
(Wellerellidae) and terebratulid (Labaiidae) species are also significantly smaller than other Late
Permian brachiopods (e.g., Orthida, Spiriferida, Strophomenida and Productida shells, which
reach lengths up to 44, 46, 47 and 70 mm, respectively).
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3.1.3. “Hedenstroemia” bosphorensis Zone (lower Olenekian)
The early Olenekian (early Smithian) “Hedenstroemia” bosphorensis Zone has been
subdivided into a lower unit (“Gyronites separatus” (= Ussuriflemingites abrekensis) Beds) and
an upper unit (Euflemingites prynadai Beds) (Zakharov, 1997). The 16- to 50-m-thick
“Gyronites separatus” Beds are characterised by the occurrence of the prolecanitid ammonoids
Aspenitidae (Parahedensroemia sp., P. kiparisovae) and Sageceratidae (Pseudosageceras cf.
multilobatum Noetling). Numerous ceratitid ammonoids also occur: Meekoceratidae (Ambitoides
orientalis Shigeta and Zakharov), Inyoitidae (Subvishnuites? sp.), Prionitidae (Radioprionites
abrekensis Shigeta and Zakharov, Gurleyites sp.), Flemingitidae (Ussuriflemingites abrekensis
Shigeta and Zakharov), Xenoceltitidae (Preflorianites cf. radiatus Chao), Proptychitidae
(Paranorites varians (Waagen), Kummelia? sp.), Arctoceratidae (Arctoceras septentrionale
(Diener)), Clypeoceratidae (Clypeoceras spitiense (Krafft)), and Columbitidae (Proharpoceras
carinatitabulatum Chao).
The Euflemingites prynadai Beds, which are at least 106 m thick, are also characterised
by the presence of prolecanitid ammonoids Hedenstroemiidae (“Hedenstroemia” bosphorensis
Zakharov), Aspenitidae (Parahedenstroemia conspicienda Zakharov), Sageceratidae (e.g.,
Pseudosageceras multilobatum Noetling), Ussuriidae (e.g., Ussuria schamarae Diener), and
numerous ceratitid ammonoids: Meekoceratidae (e.g., Meekoceras subcristatum Kiparisova,
Abrekites edites Shigeta and Zakharov, A. planus Shigeta and Zakharov), Dieneroceratidae (e.g.,
Dieneroceras chaoi Kiparisova), Inyoitidae (Inyoites spicini Zakharov), Prionitidae
(Radioprionites abrekensis Shigeta and Zakharov), Flemingitidae (Euflemingites prynadai
(Kiparisova), E. artyomensis Smyshlyaeva, Flemingites radiatus Waagen, F. aff. glaber
Waagen, Balhaeceras balhaense Shigeta and Zakharov, Rohillites laevis Shigeta and Zakharov,
Palaeokazakhstanites ussuriensis (Zakharov)), Palaeophyllitidae (Anaxenaspis orientalis
(Diener), Eophyllites sp., E. ascoldiensis Zakharov), Xenoceltitidae (Preflorianites cf. radiatus
Chao), Arctoceratidae (e.g., Arctoceras septentrionale (Diener)), Clypeoceratidae (Clypeoceras
timorense (Wanner)), Melagathiceratidae (Juvenites simplex (Chao), J. dieneri Hyatt and Smith,
J. cf. septentrionalis (Smith), J. aff. sinuosus Kiparisova, Ussurijuvenites popovi Smyshlyaeva
and Zakharov, U. artyomensis Smyshlyaeva and Zakharov), Paranannitidae (e.g., Paranannites
novikensis Zakharov, Prosphingitoides ovalis Kiparisova), and Owenitidae (Owenites koeneni
Hyatt and Smith). The early Smithian ammonoid succession is dominated by ceratitid
ammonoids, all of which seem to be Mesozoic-type ones, as well as by four other prolecanitid
ammonoid families.
No articulated brachiopods were found in the lower part of the “Hedenstroemia”
bosphorensis Zone. The articulated brachiopod assemblage of the Euflemingites prynadai Beds
is characterised by species of the families Wellerellidae (Wellerellidae gen. and sp. nov.),
Norellidae (Piarorhynchella? sp. nov.), and Rhynchonellidae (Abrekia sulcata Dagys). For
detailed biostratigraphic distribution data, see Online Supplement (Figs. S1 and S4). Among
these three early Smithian articulated brachiopod families, only the Wellerellidae is a
Palaeozoic-type one (Fig. 2). The ‘Lilliput effect’ continuous to be evident among Early
Smithian brachiopods of these families, which have maximal shell lengths of only 7 to 8 mm.
3.1.4. Anasibirites nevolini Zone (lower Olenekian)
The early Olenekian (late Smithian) Anasibirites nevolini Zone (20-68 m thick) is marked
by the first appearance datum of the genus Anasibirites. The assemblage is characterised by rare
occurrences of the prolecanitid ammonoids Sageceratidae (Pseudosageceras sp. indet.) and
Aspenitidae (Parahedenstroemia nevolini Burij and Zharnikova), but by numerous ceratitid
ammonoids, represented by Meekoceratidae (e.g., Meekoceras subcristatum Kiparisova),
Tirolitidae (Bandoites elegans Zakharov), Prionitidae (e.g., Hemiprionites contortus Burij and
Zharnikova, Shigetaceras dunajense (Zakharov), Arctoprionites ovalis Burij and Zharnikova,
Anasibirites nevolini Burij and Zharnikova, Wasatchites sikhotealinensis Zakharov, Gurleyites
maichensis Burij and Zharnikova), Palaeophyllitidae (Burijites skorochodi Burij and Zharnikova,
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Anaxenaspis sp. nov.), Xenoceltitidae (Preflorianites? sp.), Arctoceratidae (e.g., Arctoceras
labogense (Zharnikova), Churkites syaskoi Zakharov and Shigeta), Melagathiceratidae (Juvenites
simplex (Chao)), Paranannitidae (Paranannites minor Kiparisova, Prosphingitoides ovalis
(Kiparisova)), and Owenitidae (Owenites koeneni Hyatt and Smith). For detailed ammonoid
distribution data, see Online Supplement (Figs. S3 and S4). The late Smithian ammonoid
succession is dominated by ceratitid ammonoids, all of which seem to be Mesozoic-type ones, as
well as by two other prolecanitid ammonoid families (Fig. 2). Very rare and small rhynchonellid
brachiopods (up to 6 mm in length) were found in the zone (Table S4), but their preservation is
often moderate to poor in clayey facies.
3.1.5. Tirolites-Amphistephanites Zone (upper Olenekian)
The early Olenekian (early Spathian) Tirolites-Amphistephanites Zone (40-50 m thick) is
marked by the first appearance datum of the genera Tirolites and Amphistephanites, used here for
recognition of the base of the Spathian. The assemblage is characterised by the prolecanitid
ammonoids Sageceratidae (Pseudosageceras sp.) and Aspenitidae (Parahedenstroemia sp.) and
by ceratitid ammonoids, represented by Meekoceratidae (Bajarunia dagysi Zakharov),
Tirolitidae (Tirolites usuriensis Zharnikova, T. subcassianus Zakharov, Bandoites elegans
Zakharov), Dinaritidae (Tchernyschewites costatus Zakharov), Olenikitidae (e.g., Kazakhstanites
sonticus Zakharov), Stephanitidae (Amphistephanites parisensis (Zakharov)) and Flemingitidae
(Guangxiceras tobisinense (Zakharov)). The early Spathian ammonoids assemblages of the
Tirolites-Amphistephanites Zone are dominated by Ceratitida, whereas Prolecanitida are very
rare. All recognized ammonoid families seem to be Mesozoic-type ones.
This time interval studied in South Primorye is characterised by the presence of abundant
articulated brachiopods, represented by the next families of the orders Rhynchonellida,
Athyritida, Spiriferinida, and Terebratulida: Rhynchonellidae (e.g., Rhynchonellidae gen and sp.
nov. A), Neoretziidae (e.g., Hustedtiella planicosta Dagys), Lepismatinidae (Lepismatina aff.
mansfieldi (Girty)), Diplospirellidae (Spirigerinella pygmaea Dagys), Dielasmatidae
(“Fletcherithyris” margaritovi Dagys), Zeilleriidae? (Zeilleriidae? gen. and sp. indet.), and
Zeilleriidae? (e.g., Zeilleriidae? gen. and sp. nov. A). For detailed biostratigraphic distribution
data, see Online Supplement (Figs. S3 and S8). Observed ranges across the TirolitesAmphistephanites Zone are summarized in Figure 2. However, of the five brachiopod families
known from this stratigraphic interval, two (Neoretziidae and Dielasmatidae) are Palaeozoic-type
ones. The shell sizes of all these early Spathian brachiopod families are quite reduced (<10 mm
in length), with the exception of some dielasmatid brachiopods (e.g., “Fletcherithyris”
margaritovi) that are up to 25 mm long.
3.1.6. Neocolumbites insignis Zone (upper Olenekian)
The late Olenekian (middle Spathian) Neocolumbites insignis Zone (66-95 m thick) is
marked by the first appearance datum of the genus Neocolumbites. The ammonoid assemblage is
characterised by species of both the Prolecanitida and Ceratitida orders. Ceratitid ammonoids are
represented by Meekoceratidae (Svalbardiceras zhitkoviense Zakharov), Tirolitidae (Tirolites cf.
subcassianus Zakharov), Palaeophyllitidae (Burijites skorochodi (Burij and Zharnikova),
Leiophyllites praematurus Kiparisova), Keyserlingitidae (e.g., Olenekoceras meridianus
Zakharov), Columbitidae (e.g., Neocolumbites insignis Zakharov, Columbites ussuriensis Burij
and Zharnikova, Procolumbites subquadratus Burij and Zharnikova, Hellenites inopinatus
Kiparisova), and Procarnitidae (Procarnites sp.). The prolecanitid ammonoids Sageceratidae
(Pseudosageceras sp.) and Khvalynitidae (Khvalynites unicus (Kiparisova)) are present in lower
abundance. The ammonoid assemblages of this zone are dominated by Ceratitida. All middle
Spathian ammonoid families seem to be Mesozoic-type ones. Small spiriferinid brachiopod
shells, up to 12 mm in length, were found in this zone.
3.1.7. Subfengshanites multiformis Zone (upper Olenekian)
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The late Olenekian (late Spathian) Subfengshanites multiformis Zone (16-18 m thick) is
marked by the first appearance datum of the genus Subfengshanites. This zone is characterised
by the occurrence of abundant ceratitid and very rare prolecanitid (Pseudosageceras sp.)
ammonoid species. The main ceratitid ammonoid taxa known from this level are the
Dieneroceratidae (Dieneroceras karasini Kummel and Teichert), Palaeophyllitidae
(Palaeophyllites superior Zakharov, Leiophyllites sp.), Paranannitidae (e.g., Zhitkovites insularis
(Kiparisova), Pseudoprosphingites globosus (Kiparisova), Isculitoides? suboviformis
(Kiparisova)), Columbitidae (Subfengshanites multiformis Kiparisova, Arnautoceltites gracilis
(Kiparisova), and Prenkites aff. timorensis Spath). All recognized ammonoid families seem to be
Mesozoic-type ones.
The Subfengshanites multiformis Zone is characterised also by the occurrence of
Rhynchonellidae (Rhynchonellidae gen. et sp. nov. 1, 2 and 3), Lepismatinidae (Lepismatinidae
gen. and sp. nov.), and Neoretziidae (Hustedtiella planicosta Dagys). For detailed
biostratigraphic distribution data and brachiopod shell size, see Online Supplement (Figs. S5 and
S8). Observed ranges across the Subfengshanites multiformis Zone are summarized in Figure 2.
Among brachiopod families only the Neoretziidae crossed the Permian-Triassic boundary.
3.2. Mangyshlak (Kazakhstan)
The upper Olenekian in Mangyshlak has been investigated in detail in the Dolnapa Well
reference section (Bajarunas, 1936; Astachova, 1960; Shevyrev, 1968; Gavrilova, 1980, 1989,
2007; Balini et al., 2000; Zakharov et al., 2008), located in the Kara-Tau area. Two
lithostratigraphic units, corresponding to the Olenekian Tjururpiniaya Series, can be recognized
in this section: the Tartalinskaya Fm. (Tirolites cassianus-Kiparisovites carinatus and
Columbites parisianus-Procolumbites zones), which is composed of mudstone, siltstone and
fine-grained sandstone with calcareous interlayers, and the Karadzhatykskaya Fm.
(Arnautoceltites bajarunasi-Stacheites undatus Zone and Eumorphotis Beds), which consists of
intercalations of mudstone with calcareous boulders, siltstone with plant detritus, and finegrained sandstone with interlayers of limestone (Fig. S9).
3.2.1. Tirolites cassianus-Kiparisovites carinatus Zone (upper Olenekian)
The ammonoid assemblage of the Upper Olenekian (Lower Spathian) Tirolites cassianusKiparisovites carinatus Zone (which is at least 113 m thick) is characterised by the presence of
two ammonoid orders: the Prolecanitida (Sageceratidae - Pseudosageceras longilobatum
Kiparisova) and the Ceratitida. The latter consists of Dinaritidae (“Dinarites” asiaticus
Shevyrev, “D.” orientalis Shevyrev), Olenikitidae (Tjururpites cf. costatus Shevyrev,
Kiparisovites carinatus Astachova, Hyrcanites nodosus Shevyrev, Kazakhstanites dolnapaensis
Shevyrev), Tirolitidae (e.g., Tirolites cf. cassianus Quenstedt), ?Prionitidae (Albanites gracilis
(Kiparisova)) and Doricranitidae (Doricranites) (Fig. 4). The early Spathian ammonoid
assemblages of the Tirolites cassianus-Kiparisovites carinatus Zone are dominated by Ceratitida,
and no Palaeozoic-type ammonoid families were identified.
Articulated brachiopods are represented by three orders (Rhynchonellida, Spiriferida, and
Terebratulida) and several families: Norellidae (Piarorhynchella mangyshlakensis Dagys),
Lepismatinidae (Lepismatina sp. nov.), and Dielasmatidae (“Fletcherithyris” margaritovi
Dagys) (Fig. 4). No Palaeozoic-type brachiopod families were discovered in this zone. As in the
case with the Lower Triassic of South Primorye, size reduction (‘Lilliput effect’) has been
observed in many brachiopods from the Tirolites cassianus-Kiparisovites carinatus Zone. Shell
length is mostly between 6 and 10 mm, with the exception for Dielasmatidae (up to 20 mm in
length).
3.2.2. Columbites parisianus-Procolumbites Zone (upper Olenekian)
The ammonoid assemblage of the Upper Olenekian (middle Spathian) Columbites
parisianus-Procolumbites Zone (274 m thick) is characterised by the presence of prolecanitid
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ammonoids Sageceratidae (Pseudosageceras longilobatum Kiparisova) and Khvalynitidae
(Khvalynites mangyshlakensis Shevyrev) and abundant ceratitid ammonoids: Dinaritidae
(“Dinarites” asiaticus Shevyrev, “D.” orientalis Shevyrev), Xenoceltitidae (Xenoceltites
mangyshlakensis Shevyrev, X. bajarunasi Shevyrev, Preflorianites kiparisovae Shevyrev),
Kashmiritidae (Kashmirites subdimorphus Kiparisova, K. popowi Shevyrev, Eukashmirites
subdimorphus (Kiparisova)), Olenikitidae (Tjururpites costatus Shevyrev, Kiparisovites ovalis
Shevyrev, Kazakhstanites dolnapaensis Shevyrev), Tirolitidae (Tirolites armatus Shevyrev, T.
longilobatus Shevyrev, T. rossicus Kiparisova), Doricranitidae (Doricranites acutus
Mojsisovics, D. bogdoanus (Buch)), ?Prionitidae (Albanites gracilis Kiparisova),
Palaeophyllitidae (Leiophyllites exacudus Shevyrev), Procarnitidae (Procarnites kokeni
Arthaber), and Columbitidae (Columbites cf. parisianus Hyatt and Smith, C. ventroangustus
Shevyrev, Procolumbites karatauchicus Astachova, Hellenites kazakhstanicus Shevyrev,
Mangyshlakites mirificus Shevyrev) (Fig. 4). The early Spathian ammonoid assemblages of the
Columbites parisianus-Procolumbites Zone are dominated by Ceratitida, and no Palaeozoic-type
ammonoid families were recognized.
Articulated brachiopods are represented by four orders (Rhynchonellida, Athyridida,
Spiriferinida, and Terebratulida), among which a number of families were recognized:
Pontisiidae (Prelissorynchia (?) sp. nov., Lissorhynchia sp. nov., Piarorhynchella
mangyshlakensis Dagys), Neoretziidae (Hustedtiella planicosta Dagys), Diplospirellidae
(Spirigerellina sp., Spirigerellinae gen. and sp. indet.), Lepismatinidae (Lepismatina sp. nov.),
Antezeilleriidae (Antezeilleria sp.), and an uncertain family (Proanadyrella (?) sp., Thyratryaria
sp. nov. A, Thyarotryaria sp. nov. B) (Fig. 4). The genus Prelissorynchia (Pontisiidae) possibly
crossed the Permian-Triassic boundary; formerly, this genus was considered to be only Late
Palaeozoic in age. Brachiopod miniaturization is also recorded for this level, with shell lengths
for all of the families above measuring between 5.3 and 10 mm.
3.2.3. Arnautoceltites bajarunasi-Stacheites undatus Zone (upper Olenekian)
The ammonoid assemblage of the Upper Olenekian (late Spathian) Arnautoceltites
bajarunasi-Stacheites undatus Zone (172 m thick) is characterised by the presence of
prolecanitid ammonoids Sageceratidae (Pseudosageceras sp.) and some ceratitid ammonoid
families: Dinaritidae (“Dinarites”orientalis Shevyrev, Stacheites undatus (Astachova), S.
concavus Shevyrev), Xenoceltitidae (Preflorianites sp.), Olenikitidae (Kazakhstanites
dolnapaensis Shevyrev, Tjururpites costatus Shevyrev, Kiparisovites ovalis Shevyrev),
Tirolitidae (Tirolites armatus Shevyrev), Prionitidae (Arctoprionites sp., Albanites gracilis
(Kiparisova), and Columbitidae (Arnautoceltites bajarunasi (Astachova)). The late Spathian
ammonoid assemblages of the Arnautoceltites bajarunasi-Stacheites undatus Zone are
dominated by Ceratitida, and no Palaeozoic-type ammonoid families were recognized.
Three articulated brachiopod orders (Rhynchonellida, Spiriferinida, and Terebratulida)
were found in the Arnautoceltites bajarunasi-Stacheites undatus Zone. They are represented by
the families Pontisiidae (Lissorhynchia sp. nov., Piarorhynchella mangyshlakites Dagys),
Lepismatinidae (Lepismatina sp. nov.), and an uncertain family (Thyratryaria aff, pertumida Xu
and Liu). Among these brachiopod families, only a single Palaeozoic-type one (Pontisiidae) has
been discovered. The ‘Lilliput effect’ is also prevalent among the brachiopod fauna of the
Arnautoceltites bajarunasi-Stacheites undatus Zone. Lissorhynchia exhibits shell lengths to a
maximum of 5.3 mm, and other latest Spathian brachiopods are only marginally larger (up to 710 mm).
3.2.4. Eumorphotis Beds (upper Olenekian)
In the uppermost part of the Olenekian in the Dolnapa Well section, the ~151-m-thick
Eumorphotis Beds yielded very rare prolecanitid ammonoids (Pseudosageceras sp.) together
with bivalves, gastropods, and nautiloids. However, ceratitid ammonoids and articulated
brachiopods have not been discovered here.
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4. Discussion
4.1. Overview of Early Triassic brachiopod and ammonoid faunas
4.1.1. Palaeozoic-type taxa
About 13 families of articulated brachiopods survived the end-Permian mass extinction.
Among them, 13 Palaeozoic-type brachiopod genera have been identified in the Lower Triassic:
Cathaysia, Paryphella, Spinomarginifera, Retimarginifera (?), Acosarina, Prelissorhynchia (?),
Araxathyris, Spirigerella, Orbicoella, Crurithyris, Paracrurithyris, Paraspiriferina, and
Tethyochonetes (Chen et al., 2005b; Shen et al., 2006; present study) (Table S1). They belong to
the Paraspiriferinidae, Schizophoriidae, Productellidae, Ambocoellidae, Pontisiidae, and
Athyrididae families. All these genera, with the exception of Orbicoelia, Prelissorhynchia (?)
and possibly Retimarginifera, were discovered only in the lower Griesbachian. However,
Retimarginifera (?) was reported from the lower Olenekian (Chen et al., 2005b),
Prelissorhynchia (?) from both the Griesbachian (Chen et al., 2005b) and the upper Olenekian
(this study), and Orbicoelia only from the upper Olenekian (Chen et al., 2005b). Besides, at least
six additional Permian-type brachiopod genera belonging to six other families (Diplospirellidae,
Neoretziidae, Laballidae, Pennospiriferinidae, Dielasmatidae, and Angustothyrididae) also may
have survived into the Lower Triassic (Fig. 5).
However, only four ammonoid lineages (representing different taxonomic levels)
survived the mass extinction: (1) Xenodiscidae and Dzhulfitidae (at the family level), (2)
Otoceratoidea (at the superfamily level), and (3) Episageceras (at the generic level). Only a
single Palaeozoic-type genus (Episageceras) is for certain known from the Lower Triassic (Fig.
5). In Siberia and the Russian Far East, representatives of Episageceras were collected from the
lower and upper parts of the Induan (Popow, 1961; Zakharov, 1978; Dagys and Ermakova,
1996). About 18 Mesozoic-type genera have been reported from the Induan (Table S1).
A few ammonoid genera have been recorded in the Upper Changhsingian of the Meishan
section in South China: Pseudogastrioceras, Otoceras (?), Hypophiceras, Metophiceras,
Tompophiceras, “Glyptophiceras”, and Pseudosageceras (Hypophiceras fauna) (Yang et al.,
1996; Yin et al., 1996). In the South China region, at least three genera crossed the PermianTriassic boundary (e.g., Yang et al., 1996; Yin et al., 1996; McGovan, 2005; Brayard et al.,
2009a,b). McGovan and Smith (2007) have shown in their Fig. 7 that Aldanoceras, as well as
Otoceras, Xenodiscus, Hypophiceras, Tompophiceras, Metophiceras and Episageceras, crossed
the Permian-Triassic boundary (established stratigraphic ranges are shown there as solid lines),
using Dagys and Ermakova’s (1996) data on the Verkhoyansk area. However, the age of the
Otoceras boreale Zone in the Boreal realm is under debate (e.g., Kozur, 1998; Shevyrev, 2006).
Furthermore, as was recently emphasized by Shevyrev (2006), ammonoids of the so-called
Hypophiceras fauna in Meishan represent a mixed assemblage of Permian and Triassic forms.
The composition of the assemblage at Meishan is in doubt because of poor preservation, where
ammonoid shells are strongly deformed and suture lines are usually unobservable (Shevyrev,
2006). In this situation, some Permian araxoceratid ammonoids could have been mistaken for
Triassic Otoceras, Permian Paratirolites or Pseudotirolites for Triassic Tompophiceras, and
Permian Xenodiscus for Triassic Hypophiceras.
According to Brayard et al. (2007a), the Smithian genus Proharpoceras probably was
derived from the Wuchiapingian Anderssonoceratidae (Otoceratina), thus implying that an
additional lineage among Ceratitida crossed the Permian-Triassic boundry. However, this
inference concerning two taxa from distant stratigraphic levels needs to be tested through further
morphological analysis and additional fossil finds.
4.1.2. Mesozoic-type taxa
About 15 Mesozoic-type articulated brachiopod genera were found in the Induan (Table
S1), some of which are known from Russia (e.g., Lissorhynchia, Abrekia, Rhynchonellidae gen.
nov. 4, Piarorhynchella (?), Wellerellidae gen. nov. A, Wellerellidae gen. nov. B, and
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Pontisiinae gen. nov.). However, Chen et al. (2005a) reported only 7 Mesozoic-type genera from
the Induan of South China (Shen and He, 1994; Shen et al., 2006; Chen et al., 2002, 2005b),
Idaho (Girty, 1927), Himalayas (Bittner, 1899), and North Caucasus (Gefo and Sunduki areas)
(Dagys, 1974). The taxonomic diversity of brachiopods during the Dienerian was only about
11% of that for the Changhsingian (Fig. 6).
Among Olenekian articulated brachiopods, only three Palaeozoic-type genera
(Retimarginifera (?), Orbicoelia, and Prelissorhynchia (?)) but 40 Mesozoic-type genera are
known. They are identified as Maorirhynchia (?), Rhynchonellidae gen. nov. 1, 2 and 3,
Nudirostralina,
Abrekia,
Sinuplicorhynchia
(?),
Paranorellina,
Piarorhynchella,
Aparimarhynchia, Lissorhynchia, Spirigerellina, Spirigerellinae gen. nov., Neoretzia,
Hustedtiella, Lepismatina, Spiriferinidae gen. nov. A, Spiriferinidae gen. nov. B, Obnixia,
“Fletcherithyris”, Rhaetina, Portneufia, Antezeilleria, Zeilleriidae? gen. nov. A, B, C, D and F,
Periallus, Protogusarella, Thyratryaria, Proanadyrella, and Vex, of which twelve are new
genera (Table S1). Chen et al. (2005b) reported only 18 Mesozoic-type brachiopod genera in the
Olenekian on the basis of data from South China (Sun et al., 1981; Xu and Liu, 1983; Chen,
1983), southwest Japan (Dagys, 1974), Idaho (Girty, 1927; Hoover, 1979; Perry and Chatterton,
1979), Himalayas (Bittner, 1899), Tibet (Sun et al., 1981; Chen, 1983), Dobrogea (Jordan,
1993), New Zealand (MacFarlan, 1992), and Spitsbergen (Dagys, 1974). The taxonomic
diversity of brachiopods during the Smithian and Spathian was only about 16 and 19% of that for
the Changhsingian, respectively (Fig. 6).
Induan ammonoids are apparently represented by only a single Palaeozoic-type genus
(Episageceras) but 67 Mesozoic-type genera (Fig. S12). Only Mesozoic-type ammonoid genera
have been reported from the Smithian and Spathian which have yielded 136 and 129 genera,
respectively (Fig. 6; Table S1). McGovan (2005) inferred that ammonoids did not fully recover
until the Spathian or Anisian. However, recent data (Brayard et al., 2009a,b; Brühwiler et al.,
2010a,b,c, 2011a,b; Guex et al., 2010; this study) show that the recovery of ammonoid faunas
proceeded significantly more rapidly. Brayard et al. (2009a,b) conclude that ammonoid diversity
level was higher than the maximum level recorded during all the Permian by the Smithian.
According to our data, based on recently published and original evidences, ammonoids reached
levels of taxonomic diversity during the Dienerian that were 146% higher than in the
Changhsingian, representing a much faster recovery than that for brachiopods (the diversity of
brachiopods during the Dienerian was only about 8% of that for the Changhsingian ) (Fig. 6).
Maximum taxonomic diversity of Permian ammonoids is associated with the Wordian-Roadian
(Bogoslovskaya et al., 1999). However, the diversity of ammonoids during the Smithian (Table
S1) reached not less than 200% of that for the Wordian.
4.2. Patterns of geographical differentiation of Early Triassic brachiopod and ammonoid faunas
Early Triassic ammonoid diversification produced about 290 genera, while Early Triassic
articulated brachiopod diversification produced only about 68 genera (Table S1). Griesbachian
relict articulated brachiopods have been recovered from South China (Sheng et al., 1984; Xu and
Grant, 1994; Liao, 1979a,b, 1980, 1984, 1987; Yang et al., 1983; Chen and Shi, 1999; Chen et
al., 2000, 2005a,b), Salt Range (Grant, 1970; Kummel and Teichert, 1970), Nepal (Waterhouse,
1978, 1994; Waterhouse and Shi, 1991), Kashmir (Shimizu, 1981) and Oman (Twitchett et al.,
2004; Chen et al., 2005a,b). During Olenekian time, one group of Permian relict species
continued to exist in Nepal, while another one migrated possibly to the Mangyshlak and
Romanian areas. At the beginning of the Induan, just after the Late Permian mass extinction, a
single Palaeozoic-type ammonoid Episageceras migrated from the Tethys to the Boreal realm
(Russian North-East) (Popow, 1961; Zakharov, 1978; Dagys and Ermakova, 1996), together with
a Mesozoic-type ammonoid group. The latter included apparently ancestral forms of the
Otoceras (Otoceratidae) and Tompophiceras (Dzhulfitidae) (Zakharov et al., 2005, 2008). We
hypothesize that these ammonoids migrated into higher northern latitudes to restore their food
supply that had been disrupted by major temperature fluctuations (Zakharov et al., 2001, 2008;
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Kozur, 2007) and oceanic anoxia (Hallam, 1994; Isozaki, 1997; Wignall and Twitchett, 2002)
associated with the Permian-Triassic boundary crisis.
Localities of Induan Mesozoic-type articulated brachiopod are known only in lower to
middle-latitude regions: South China (Xu and Grant, 1994; Shen and He, 1994; Chen et al.,
2002, 2005b); northwest Caucasus (Dagys, 1974), Himalayan region (Bittner, 1899; Waterhouse
and Shi, 1991; Chen et al., 2005a,b), South Primorye (Dagys, 1974; this study), and Idaho (Girty,
1927). However, the most diverse Induan brachiopod faunas were reported from South China (14
genera, including five Mesozoic-type genera) and South Primorye (six Mesozoic-type genera)
(Fig. S10). No Induan articulated brachiopods have been discovered in high palaeolatitudes.
Data of the present study show that the most diverse Olenekian articulated brachiopod
faunas are present in the former USSR area (South Primorye and Mangyshlak). These faunas
consist of 21 and 13 genera, respectively (Fig. S11). Other known localities of Olenekian
articulated brachiopods include Idaho (Girty, 1927; Hoover, 1979; Perry and Chatterton, 1979),
South China (Perry and Chatterton, 1979), Carpathians-Balkanian area (Jordan, 1993), Tibet
(Sun et al., 1981; Chen, 1983; Chen et al., 2005b), Japan (Dagys, 1993), Spitsbergen (Dagys,
1974, 1993), Alps (Jordan, 1993), and New Zealand (MacFarlan, 1992).
Among known Early Triassic articulated brachiopods, only a few Olenekian genera
(Obnixia, Hustedtiella? (Dagys, 1974) and Aparimirhynchia (MacFarlan, 1992)) reached highlatitude areas. In contrast to Early Triassic brachiopods, Induan (Figs. S12) and Olenekian
ammonoid assemblages, according to our data, are common for both the tropical-subtropical area
and the high-latitude region of the Boreal realm, which is in agreement with previous studies on
this topic (e.g., Tozer, 1994; Brayard et al., 2006b, 2009a,b).
4.3. Palaeophysiological and -behavioural contexts of end-Permian environmental stresses
The end-Permian-Early Triassic interval is dominated by unusual oceanic and climatic
conditions (e.g., oceanic anoxia, expansion of desert belts, declining atmospheric levels of O2 but
increasing of levels of CO2, a biocalcification crisis; Baud et al., 1989; Holser et al., 1989;
Wignall and Hallam, 1993; Hallam, 1994; Kidder and Worsley, 2004; Galfetti et al., 2007; Payne
and Kump, 2007). Oceanic anoxia has been invoked as the main kill mechanism of the PermianTriassic extinction event (Wignall and Hallam, 1993; Wignall and Twitchett, 2002), although
this hypothesis has been disputed by Hermann et al. (2011) on the basis of the largely terrigenous
origin of particulate organic matter in the Lower Triassic of the Salt Range and Surghar Range.
Nevertheless, chemically and/or physiologically harsh environmental conditions certainly existed
at the end of Permian, possibly caused by unusual oceanic and climatic conditions that affected
the marine biota for several million years (Fraiser et al., 2005; Algeo and Twitchett, 2010; Algeo
et al., 2011). The largest effects were during the Griesbachian, which represents a ‘survival
phase’ for brachiopods and ammonoids following the end-Permian ecological crisis.
The patterns of recovery of brachiopods and ammonoids following the end-Permian mass
extinction differed owing to fundamental differences in their physiology and behavior.
Brachiopods and ammonoids are representatives of physiologically dissimilar groups of marine
organisms with possible metabolic and skeletal differences (Knoll et al., 2007). Brachiopods
belong to a group of organisms with a calcium carbonate skeleton that was often massive with
respect to supporting organic tissue and formed from fluids minimally buffered by physiology.
Ammonoids, in contrast, belonged to a group of organisms with a calcium carbonate skeleton of
moderate mass and formed from fluids that were relatively well buffered with respect to the
factors that govern carbonate precipitation. The latter condition is considered to be more
sensitive to high watermass PCO2 (hypercapnic stress).
Differences in feeding behaviour may have mattered also. Data on brachiopod sizes
changes in the Permian-Triassic transition in South China (Liao, 1979; Shen and He, 1994; He et
al., 2007; Chen et al., 2005a,b) and our additional evidence on the post-extinction
miniaturization of brachiopods from the Lower Triassic of the former USSR (Tables S2-S7) is in
an agreement with the hypothesis of Keller and Abramovich (2009) that high-stress conditions
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led to both biodiversity loss and size reduction (‘Lilliput effect’ in Urbanek’s (1993) sence).
According to Harries and Knorr (2007), this effect represents a pronounced reduction in size of
the biota associated with the aftermath of mass extinctions. However, by contrast with Early
Triassic brachiopods, survived the end Permian mass extinction, ammonoids from the same
habitats/facies in both the Primorye region and Kazakhstan do not show any trends toward
smaller size in the Early Triassic. These different outcomes may have been related to differences
in trophic resource availability (just after the end-Permian crisis, notably brachiopods may have
suffered from a relative paucity of food resources). Epifaunal benthos (suspension-feeding
brachiopods as well as bivalves) dominated Lower-Middle Triassic marine environments, but the
more modern life habits (e.g., infaunalization associated with burrowing suspension and deposit
feeders) increased during the Late Triassic (Bonuso and Bottjer, 2008). Judging by the
abundance and wide distribution of nektic ammonites during the Early Triassic, one can infer
that they generally were not specialized feeders, although they were apparently presumably
bottom feeders during the late Palaeozoic and Mesozoic, because optimal temperatures of their
growth are mainly comparable to those obtained from their co-occurring benthos on the shelf
(Smyshlyaeva et al., 2002; Moriya et al., 2003; Zakharov et al., 2003, 2006, 2011). Thus, Early
Triassic ammonoids had greater access to trophic resources and therefore stood a better chance
of success than brachiopods.
Another factor in the differential recovery of these two clades may have been related to
differences in larval/juvenile survival and development. After hatching, brachiopod larvae and
juvenile ammonoids may be dispersed with the aid of currents. All planktic brachiopod larvae
are capable of delaying settlement until a suitable substratum has been located, although the
motile larval stage usually lasts only days or hours (Williams et al., 1997). High sedimentation
rates during the Early Triassic may have resulted in generally more mobile substrates that were
unfavorable for larval brachiopod settlement (cf. Algeo and Twitchett, 2010). Ammonoids, in
contrast, are characterised by direct ontogenetic development without a larval stage and, thus,
were not dependent on substrate characteristics. The survival of planktic brachiopod larvae was
significantly lower because of the short duration of their motile phase, the necessity of finding a
suitable substrate, and the specificity of their trophic needs--factors that worked to their
disadvantage during the Early Triassic. The contrast in recovery patterns for brachiopods and
ammonoids was most evident at high palaeolatitudes, most difficult apparently place for
brachiopod larvae migration.
5. Conclusions
Ammonoids exceeded their pre-crisis taxonomic diversity and abundance by the late
Induan (latest Dienerian). In contrast, brachiopods never recovered their pre-crisis diversity and
abundance anytime during the Triassic or later. The differential recovery patterns of brachiopods
and ammonoids seem to be a result of physiological and behavioural differences, reflected in
their different life modes and distributions in ecospace. Specifically, brachiopods were at a
disadvantage during the Early Triassic because of the short duration of their motile phases, the
need to find a suitable substrate for larval settlement, and the specificity of their trophic
requirements. The contrast in recovery patterns was most evident at high palaeolatitudes.
Diversification of Early Triassic brachiopods at high latitudes was quite restricted: no articulated
brachiopods of Induan age, and only three genera of Olenekian age (Obnixia and Hustedtiella?
(Dagys, 1974) and Aparimirhynchia (MacFarlan, 1992)) have been discovered in the Boreal
realm. However, immediately after the end-Permian extinction event, ammonoids colonized and
stably inhabited many regions in the Boreal realm throughout the Triassic. The contrast in
recovery patterns for brachiopods and ammonoids was most evident at high palaeolatitudes.
Acknowledgments
For help in finding some important Early Triassic brachiopod and ammonoid fossils in
South Primorye and regular assistance, we are indebted to Y. Shigeta (National Museum of
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Nature and Science, Tokyo), H. Maeda, and T. Kumagae (Kyoto University) and also to V.T
S’’edin (Institute of Pacific Oceanology, Vladivostok). Our cordial thanks are due to Prof. T.A.
Algeo (University of Cincinnati) for providing valuable editorial comments, Prof. Shuzhong
Shen (Nanjing Institute of Geology and Palaeontology) and anonymous reviewer for stimulating
remarks, as well as for their information on latest published papers on Early Triassic ammonoids
and data on this topic in press, that substantially improved this paper. This work is a contribution
to UNESCO-IUGS IGCP Project 572 and financially supported by the Russian grants RFBR
(11-05-98538-R_vostok_a and 11-05-000785-a).
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951
952
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969
970
971
972
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973
20
974
975
976
977
978
979
980
981
982
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. (2005b); genera survived the Permian-Triassic b
oundary are indicated by asterisk; Early Triassic brachiopods from the Abrek section are illustrated in Sigeta et al., 2009 – figs. 146-148).
1. Early Induan (Griesbachian) articulated brachiopod genera.
N
Genus
Main source of information
N
Genus
1
Acosarina* Cooper Chen et al., 2005
9
Paryphella* Liao
& Grant
2
Araxathyris*Grunt Chen et al., 2005
10
Prelissorhynchia*
Xu & Grant
Cathaysia* Ching
Chen et al., 2005
11
Pustula* Thomas
3
4
Laevorhynchia
Shen and He, 1994; Mu et
12
?Retimarginifera*
Shen & He
al., 2007
Waterhouse
5
Orbicoelia*
Chen et al., 2005
13
Spinomarginifera*
Waterhouse &
Huang
Piyasin
6
Crurithyris *
Yang et al., 1987; Shen et
14
“Spiriferina”
George
al., 2006
D’Orbigny
7
Paracrurithyris*
Chen et al., 2005
15
Spirigerella*
Liao
Waagen
8
Paraspiriferina*
Chen et al., 2005
16
Tethyochonetes*
Reed
Chen & others
2. Late Induan (Dienerian) articulated brachiopod genera
N
Genus
Main source of information
N
1
Abrekia Dagys
Shigeta et al., 2009
7
(Figs. 147.1-147-24)
2
Lepismatina Wang Chen et al., 2005
8
3
Dagys, 1974; this study
(Figs. 3-1 and 3-2)
9
MacFarlan, 1992
10
5
Lissorhynchia
Yang & Xu
(=Neowellerella”)
?Maorirhynchia
MacFarlan
“Mentzelia”
Girty, 1927
11
6
Periallus Hoover
Hoover, 1979
12
4
983
984
Reference
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Chen et al., 2005
Genus
Piarorhynchella?
Dagys
Pontisiinae gen.
and sp. nov.
Retzioidea gen. and
sp. indet.
Main source of information
Shigeta et al., 2009
(Figs. 147.29-147.32)
Shigeta et al., 2009
Figs. 147.41-147.48)
Chen et al., 2005
“Spiriferina”
D’Orbigny
Wellerellidae gen.
and sp. nov. A
Wellerellidae gen.
and sp. nov. B
Xu and Grant, 1994
3. Olenekian (Smithian and Spathian) articulated brachiopod genera
N
Genus
Main source of information
N
Genus
1
Abrekia Dagys
Shigeta et al., 2009
23
?Retimarginifera*
(Figs. 147. 1-147.24)
Waterhouse
2
Antezeilleria Xu & Xu and Liu, 1983
24
Rhaetina Waagen
Liu
3
Aparimarhynchia
MacFarlan, 1992
25
Rhynchonellidae
MacFarlan
gen. indet.
4
“Fletcherithyris”
Dagys, 1974; this study
26
Rhynchonellidae
gen. nov. A
5
Hustedtiella Dagys Dagys, 1974; this study
27
Rhynchonellidae
(Plate I, figs. 23-26)
gen. nov. 1
6
Lepismatina Wang Dagys, 1974; this study
28
Rhynchonellidae
(Figs. 3-31--3-34)
gen. nov. 2
7
Lepismatinidae
This study
29
Rhynchonellidae
gen. and sp. nov.
gen. nov. 3
8
Lissorhynchia
Original data (Plate I, figs.
30
Rhynchonellidae
Yang & Xu
19-22)
gen. indet.
9
Neoretzia Dagys
Chen, 1983
31
“Rhynchonella”
Fischer de
Waldheim
Shigeta et al., 2009
Shigeta et al., 2009
Main source of information
Chen et al., 2005
Dagys, 1974; Hoover, 1979
This study (Figs. 3-7—310)
This study (Figs. 3-11—314)
This study
This study (Figs.3-15—318)
This study
This study
Chen et al., 2005
21
10
Nudirostralina
Yang & Xu
?Sinuplicorhynchia
Dagys
Spirigerellinae gen.
and sp. nov.
Obnixia Hoover
Chen, 1983
32
This study
33
This study
34
Hoover, 1979
35
Chen et al., 2005
36
Dagys, 1974
37
16
Orbicoelia*
Waterhouse &
Piyasin
Paranorellina
Dagys
Periallus Hoover
Chen et al., 2005
38
17
Portneufia Hoover
Hoover, 1979
39
18
Piarorhynchella
Dagys
Prelissorhynchia
(?)* Xu & Grant
Proanadyrella Xu
& Liu
?Proanadyrella Xu
& Liu
Protogusarella
Dagys, 1974
40
Chen et al., 2005; this study
(Figs. 3-3--3-6)
Xu and Liu, 1983
41
This study
43
11
12
13
14
15
19
20
21
22
985
986
987
988
42
Spirigerellina
Dagys
Spirferiinidae gen.
nov. A
Spirferiinidae gen.
nov. B
Zeilleriidae gen.
nov. A
Zeilleriidae gen.
nov. B
Dagys, 1974; this study
(Figs. 3.27-3.30)
This study (Figs. 3-35—338)
This study
Zeilleriidae gen.
nov. C
Zeilleriidae gen.
nov. D
Zeilleriidae gen.
nov. E
Zeilleriidae gen.
nov. F
Thyratryaria Xu &
Liu
Vex Hoover
This study
Zeilleriidae gen.
indet.
This study
This study
This study
This study
This study
This study
Xu and Liu, 1983
Hoover, 1979
Perry and Chatterton, 1979
4. Changhsingian ammonoid genera
(Wuchiapingian-Changhsingian boundary in Transcaucasia and Iran is indicated following Zakharov et al. (2005) – inside of the Vedioceras
ventrosulcatum of the Dzhulfian )
N
Genus
Main source of information
N
Genus
1
Abichites Shevyrev
Bogoslovskaya et al., 1999
22
2
Avushoceras
Ruzhencev
Changhsingoceras
Chao & Liang
? Cyclolobus Waagen
Bogoslovskaya et al., 1999
23
Zhou et al., 1999
24
Zhou et al., 1999
25
Dushanoceras Zhao,
Liang & Zheng
Dzhulfites Shevyrev
Bogoslovskaya et al., 1999
26
Bogoslovskaya et al., 1999
27
7
Dzhulfoceras
Ruzhencev
Bogoslovskaya et al., 1999
28
8
Episageceras
Noetling
Bogoslovskaya et al., 1999
29
9
“Hypophiceras"
Trümpy (=
xenodiscid?
ammonoid)
Huananoceras Chao
& Liang
Iranites Teichert &
Kummel
Yin et al., 1996; Shevyrev,
2006)
30
Phisonites
Shevyrev
Pleuronodoceras
Chao & Liang
Pseudogastrioceras Spath
Pseudostephanites
Chao & Liang
Pseudotirolites
Sun
Pseudotoceras
Ruzhencev
Qinjiangoceras
Zhao, Liang &
Zheng
Rongjiangoceras
Zhao, Liang &
Zheng
?Rotaraxoceras
Ruzhencev
Bogoslovskaya et al., 1999
31
Bogoslovskaya et al., 1999
32
12
Julfoceras
Ruzhencev
Bogoslovskaya et al., 1999
33
13
Liuchengoceras
Zhou et al., 1999
34
3
4
5
6
10
11
Rotodiscoceras
Chao & Liang
Schizoloboceras
Zhao, Liang &
Zheng
Shevyrevites
Teichert &
Kummel
Sinoceltites Chao
Main source of
information
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
Zhou et al., 1999
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
Zhou et al., 1999
Zhou et al., 1999
Kotlyar et al., 1983;
Zakharov et al., 2005
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
Bogoslovskaya et al., 1999
22
14
15
16
17
18
19
20
21
989
990
Zhou et al., 1999
35
Zhou et al., 1999
36
Mingyuexiaceras
Zhao, Liang & Zheng
Neoaganides
Plummer & Scott
Zhou et al., 1999
37
Bogoslovskaya et al., 1999
38
“?Otoceras”
Griesbach (=
araxoceratid?
ammonoid)
Pachydiscoceras
Chao & Liang
Paratirolites
Stoyanow
Pentagonoceras
Zhao, Liang & Zheng
Yin et al., 1996; Shevyrev,
2006)
39
Zhou et al., 1999
40
Bogoslovskaya et al., 1999
41
4
5
6
7
8
& Liang
Stacheoceras
Gemmellaro
Sutchanites
Zakharov
Tapashanites Chao
& Liang
“Tompophiceras”
Popow (=
dzhulfitid?
ammonoid)
Trigonogastrites
Zhao, Liang &
Zheng
Zhou et al., 1999
Zakharov and Oleinikov,
1994
Bogoslovskaya et al., 1999
Yin et al., 1996;
Shevyrev, 2006)
Zhou et al., 1999
?Vedioceras
Ruzhencev
Xenodiscus
Waagen
Kotlyar et al., 1983;
Zakharov et al., 2005
Bogoslovskaya et al., 1999
Genus
Metophiceras
Spath
Otoceras
Griesbach
(=Metotoceras
Spath)
Ophiceras
Griesbach
Main source of information
Shevyrev, 2000
Zhou et al., 1999
5. Early Induan (Griesbachian) ammonoid genera
N
Genus
Main source of information
1
Aldanoceras Dagys Ermakova, 2002
& Ermakova
2
Anotoceras Hyatt
Shevyrev, 1999
3
991
992
993
Zhao, Liang & Zheng
Longmenshanoceras
Zhao, Liang & Zheng
Meitianoceras Zheng
N
9
10
Tozer, 1994; Shevyrev,
1986
Bernhardites
Shevyrev
(=?Shevyrevoceras
Waterhouse)
Bukkenites Tozer
Waterhouse, 1994;
Shevyrev, 1999
11
Tozer, 1994
Tozer, 1994
12
Proptychites
Waagen
(=Pseudoproptychi
tes Bando)
Waterhouse, 1994; this
study
Discophiceras
Spath
Episageceras*
Noetling
Shevyrev, 2000
13
Waterhouse, 1994
Waterhouse, 1994
14
Hypophiceras
Trümpy
Lytophiceras Spath
Shevyrev, 2000
15
Shalshalia
Waterhouse
Tompophiceras
Popow
(=Zakharovites
Waterhouse)
Vishnuites Diener
Waterhouse, 1994
16
Wordieoceras
Tozer
Tozer, 1994
Dagys and Ermakova,
1996; Shevyrev, 1999
Waterhouse, 1994
6. Late Induan (Dienerian) ammonoid genera (ammonoid assemblage from the upper Chegaji Member (top 10-30 cm) in Nepal,
investigated by Waterhouse (1994), yielding Gyronites, seems to be Dienerian in age).
N
Genus
Main source of information
N
Genus
1
2
Ambites Waagen
?Anaxenaspis
Kiparisova
Anotoceras Hyatt
Waterhouse, 1994
Ermakova, 2002; Zakharov
et al., 2010
Waterhouse, 1994
31
32
Aspiditella Strand
Aspitella
Waterhouse
Shevyrev, 1986
Waterhouse, 1996a
34
35
Meekophiceras Tozer
Mesokantoa
Waterhouse
Metophiceras Spath
(=?Altoconchella
Waterhouse)
Ophiceras Griesbach
?Mullericeras Ware,
Jenks, Hautmann &
3
4
5
33
Main source of
information
Tozer, 1994
Waterhouse, 1994
Waterhouse, 1994; this
study
Waterhouse, 1994
Ware et al., 2011
23
6
7
Bernhardites
Shevyrev
(=?Shevyrevoceras
Waterhouse)
Bukkenites Tozer
Waterhouse, 1994; this
study
36
Tozer, 1994
37
Clypeoceras Smith
(=?Aspitella
Waterhouse)
Clypites Waagen
Collignonites
Bando
Dunedinites Tozer
Spath, 1934; this study
Bucher
Pachyproptychites
Diener
Shigeta et al., 2009
Ware et al., 2011
38
?Parahedenstroemia
Spath
Paranorites Waagen
Brühwiller et al., 2010b
Shevyrev, 1986
39
40
Paraspidites Spath
Pleuroambites Tozer
Shevyrev, 1986
Tozer, 1994
Tozer, 1994
41
Tozer, 1994
Eoptychites Spath
(=Waagenoproptychites
Waterhouse)
Eovavilovites
Ermakova
Episageceras*
Popow
Brayard et al., 2006; this
study
42
Pleurogyronites
Tozer
?Preflorianites Spath
Ermakova, 2002
43
Zakharov, 1978
44
Waterhouse, 1996a;
Shevyrev, 1999
Brühwiler et al., 2008;
this study
15
Gyrolecanites
Spath
Shevyrev, 1986
45
16
Gyronites Waagen
(=?Nangykangia
Waterhouse)
Gyrophiceras
Spath
?Hedenstroemia
Waagen
Heibergites Tozer
Waterhouse, 1994; this
study
46
Mu et al., 2007
47
Zakharov et al., 2009a
48
Tozer, 1994
49
Himoceras
Waterhouse
Discophiceras
Spath
(=?Himophiceras
Waterhouse)
Hubeitoceras
Waterhouse
Jieshanites
Brühwiler,
Brayard, Bucher &
Guodun
Khangsaria
Waterhouse
Kingites Waagen
Waterhous, 1996a
50
Prejuvenites
Waterhouse
Prionolobus Waagen
(=?Lilangia
Waterhouse)
Proptychites Waagen
(=Pseudoproptychites
Bando)
?Pseudohedenstroemia
Kummel
Pseudosageceras
Diener
Pseudovishnuites
Zakharov & Mu,
Radioceras
Waterhouse
Sakhaites Vozin
Waterhouse, 1994; this
study
51
Shangganites
Brühwiler, Brayard,
Bucher & Guodun
Brühwiler et al., 2008
Waterhouse, 1994;
Brühwiler et al., 2008
Brühwiler et al., 2008
52
Subinyoites Spath
Shevyrev, 1986
53
Tompoproptychites
Vavilov & Zakharov
Vavilov and Zakharov,
1976; Brayard et al.,
2006
Waterhouse, 1996a
54
Ussuriceras Spath
Shevyrev, 1986
Shevyrev, 1986
55
Tozer, 1994; this study
26
Koninckites
Waagen
Bruhwiler et al., 2010b
56
27
Kummelia
Waterhouse
(=?Tilihonia
Waterhouse;
?Zhaojinkeoceras
Waterhouse;
Waterhouse, 1996a; this
study
57
Vavilovites Tozer
(=Fuchsites
Waterhouse)
Vercherites
Brühwiler, Ware,
Bucher, Krystyn &
Goudemand
Vishnuites Diener
8
9
10
11
12
13
14
17
18
19
20
21
22
23
24
25
Shevyrev, 1986
Shevyrev, 1986
Tozer, 1994; this study
Shevyrev, 1986
Waterhouse, 1994
Mu et al., 2007
Waterhouse, 1996a
Ermakova, 2002
Brühwiler, et al., 2010b
Waterhouse, 1994
24
28
29
30
994
995
?Mesokantoa
Waterhouse)
Kymatites Waagen
(=?Khangsaria
Waterhouse)
Lilastroemia
Waterhouse
Lytophiceras Spath
Waterhouse, 1994; this
study
58
Wordieoceras Spath
Tozer. 1994
Waterhouse, 1996a
59
?Xenoceltites Spath
Shevyrev, 1986
Waterhouse, 1994
60
Xenodiscoides Spath
Brühwiler et al., 2008
N
69
Genus
Metinyoites Spath
Main source of information
Shevyrev, 1986
70
71
Metussuria Spath
Mianwaliites
Brühwiler et al.
Monneticeras
Brühwiler et al.
Mudiceras
Brühwiler et al.
Mullericeras Ware,
Jenks, Hautmann &
Bucher
Nammalites
Brühwiler, Bucher
& Goudemand
Nanningites
Brayard & Bucher
Nilgiria
Waterhouse
?Nordophiceras
Popow
Nuezelia
Brühwiler, Bucher,
Krystyn & Breta
Nyalamites
Brühwiler, Bucher
& Goudemand
Owenites Hyatt &
Smith
Palaeokazakhstanites Zakharov
Zakharov et al., 2010
Brühwiler et al., 2011b
Parahedenstroemia
Spath
Parakymatites
Waagen
Shevyrev, 1986
Paranannites Hyatt
& Smith
Paranorites
Waagen
Paraspedites Spath
Brayard and Bucher, 2008
Parastephanites
Hyatt
Parussuria Spath
Brayard et al., 2006
Pijaconcha
Waterhouse
Preflorianites
Spath
Waterhouse, 1996b
7. Early Olenekian (Smithian) ammonoid genera
N
Genus
Main source of information
1
Abrekites Shigeta
Shigeta et al., 2009
& Zakharov
2
Ambites Waagen
Shevyrev, 1986
3
Ambitoides Shigeta Shigeta et al., 2009
& Zakharov
4
Anaflemingites
Brayard and Bucher, 2008
Kummel & Steele
5
Anahedenstroemia
Shevyrev, 1986
Hyatt
6
Anakashmirites
Shevyrev, 1986
Spath
72
73
74
7
Anasibirites
Mojsisovics
Zakharov et al., 2010
75
8
Anastephanites
Spath
Anawasatchites
McLearn
Anaxenaspis
Kiparisova
Arctoceras Hyatt
Shevyrev, 1986
76
Brayard et al., 2006
77
Zakharov et al., 2010
78
Shigeta et al. 2009
79
12
Arctoprionites
Spath
Shevyrev, 1986
80
13
Brayard and Bucher, 2008
81
Shigeta et al., 2009
82
Waterhouse, 1996b
83
Brühwiler et al., 2010a
84
17
Aspenites Hyatt &
Smith
Balhaeceras
Shigeta &
Zakharov
Boewanites
Waterhouse
Brayardites
Brühwiler, Bucher
& Goudemand
Burijites Zakharov
Zakharov, 1978
85
18
Churkites Okuneva
86
19
Brayard and Bucher, 2008
88
21
Colchenoceras
Waterhouse
Cordillerites Hyatt
& Smith
Clypeoceras Smith
Markevich and Zakharov,
2004
Waterhouse, 1996b
89
22
Clypites Waagen
Waagen, 1895; Brühwiler
et al., 2011b
Brayard et al., 2006
23
Clypeoceratoides
Dagys &
Ermakova
Ermakova, 2002
91
9
10
11
14
15
16
20
87
90
Brühwiler et al., 2011b
Brühwiler et al., 2010b
Ware, Jenks, Hautmann &
Bucher, 2011; this study
Brühwiler et al., 2011b
Brayard and Bucher, 2008
Waterhouse, 1996b
Waterhouse, 1996b
Brühwiler et al., 2011a
Brühwiler et al., 2010a
Zakharov et al., 2010
Markevich and Zakharov,
2004
Shevyrev, 1986
Shevyrev, 1986
Brayard et al., 2006
Zakharov et al., 2010
Brayard and Bucher, 2008
25
24
Brayard and Bucher, 2008
92
Prionites Waagen
Brühwiler et al., 2010a
25
Dieneroceras
Spath
Dorikranites Hyatt
Shevyrev, 1968
93
Brayard and Bucher, 2006a
26
Dunedinites Tozer
Tozer, 1994
94
27
Eoptychites Spath
Waterhouse, 1996b
95
28
Eoptychitoides
Waterhouse
Epihedenstroemia
Spath
Escargueltites
Waterhouse, 1996b;
Brühwiler et al., 2010c
Shevyrev, 1986
96
Brühwiler et al., 2011a
98
Eukashmirites
Kummel
Euflemingites
Spath
Flemingites
Waagen
Glyptophiceras
Spath
Shevyrev, 1986
99
Tozer, 1994
100
Brayard and Bucher, 2008
101
Brühwiler et al., 2010a
102
Galfettites Brayard
& Bucher
Guangxiceltites
Brayard & Bucher
Guangxiceras
Brayard & Bucher
Guodunites
Brayard & Bucher
Gurleyites
Mathews
Brayard and Bucher, 2008
103
Brayard and Bucher, 2008
104
Brayard and Bucher, 2008
105
Brayard and Bucher, 2008
106
Shevyrev, 1986
107
40
Gyronites Waagen
Tozer, 1994
108
41
42
Hanielites Welter
Hebeisenites
Brayard & Bucher
Brayard and Bucher, 2008
Brayard and Bucher, 2008
109
110
43
Hedenstroemia
Waagen
Brayard and Bucher, 2008
111
44
Hemiaspenites
Kummel & Steele
Brayard et al., 2006
112
45
Hemiprionites
Spath
Hochuliites
Brüchwiler et al.
Inyoites Hyatt &
Smith
Brayard and Bucher, 2008
113
Prionolobus
Waagen
Procurvoceratites
Brayard & Bucher
Proharpoceras
Chao
Prosphingitoides
Shevyrev
Protophiceras
Hyatt
Pseudoceltites
Hyatt
Pseudoaspenites
Spath
Pseudaspidites
Spath
Pseudoflemingites
Spath
Pseudohedenstroemia
Kummel
Pseudosageceras
Diener
Pseudokazakhstani
tes Zakharov
Punjabites
Brüchwiler et al..
?Radioceras
Waterhouse
Radioprionites
Shigeta &
Zakharov
Rohillites
Waterhouse
Sakhaites Vozin
Shamaraites
Shigeta &
Zakharov
Shigetaceras
Brühwiler, Bucher
& Goudemand
Steckites
Brühwiler, Bucher
& Krystyn
StephanitesWaagen
Brüchwiler et al., 2011b
114
Zakharov, 1978
Brayard and Bucher, 2008
115
Subalbanites
Zakharov
Subinyoites Spath
48
Jinyaceras Brayard
& Bucher
Brayard and Bucher, 2008
116
Subflemingites
Spath
Brühwiler et al., 2010a
49
Waterhouse, 1996b
117
Brayard and Bucher, 2008
118
51
Kashmirites Diener
Brayard and Bucher, 2008
119
Submeekoceras
Spath
Subvishnuites
Spath
Tellerites
Brayard and Bucher, 2008
50
Jolinkia
Waterhouse
Juvenites Smith
29
30
31
32
33
34
35
36
37
38
39
46
47
97
Brayard and Bucher, 2008
Brayard and Bucher, 2008
Shevyrev, 1995
Brayard and Bucher, 2008
Brühwiler et al., 2010a
Brayard and Bucher, 2008
Brayard and Bucher, 2008
Brayard and Bucher, 2008
Shevyrev, 1986
Brayard and Bucher, 2008
Markevich and Zakharov,
2004
Brüchwiler et al., 2011b
Brüchwiler et al., 2010b
Shigeta et al., 2009
Waterhouse, 1996b;
Brüchwiler et al., 2011b
Ermakova, 2002
Shigeta et al., 2009
Brühwiler et al., 2010a
Brühwiler et al., 2011a
Brühwiler et al., 2010a
Shevyrev, 1986;
Brüchwiler et al., 2011b
Brayard and Bucher, 2008
Brayard et al., 2006
26
52
Kashmiritidae gen.
nov. A
Kazakhstanites
Shevyrev
Kelteroceras
Ermakova
Brühwiler et al., 2010b
120
Shevyrev, 1995
121
Ermakova, 2002
122
Kingites Waagen
Koiloceras
Brühwiler et al.
Koninckites
Waagen
Kraffticeras
Brühwiler,Bucher
& Krystyn
Kymatites Waagen
Brüchwiler et al., 2010c
Brühwiler et al., 2011b
123
124
Shevyrev, 1986
125
Brühwiler et al., 2011a
126
Waterhouse, 1996a
127
60
Lanceolites Hyatt
& Smith
Brayard and Bucher, 2008
128
61
Larenites Brayard
and Bucher
Brayard and Bucher, 2008
129
62
Latisageceras
Ruzhencev
Lepiskites Dagys &
Ermakova
Leyeceras Brayard
& Bucher
Lingyunites Chao
Meekoceras Hyatt
Waterhouse, 1996b
130
Ermakova, 2002
131
Brayard and Bucher, 2008
132
Brayard and Bucher, 2008
Brühwiler et al., 2011b
134
135
Melagathiceras
Tozer
Mesohedenstroemia Chao
Tozer, 1994
136
53
54
55
56
57
58
59
63
64
65
66
67
68
996
997
998
999
Mojsisovics
Thermalites Smith
Brayard et al., 2006
Truempyceras
Brühwiler et al.
Tulongites
Brühwiler, Bucher
& Goudemand
Tuyangites Zhao
Urdyceras Brayard
& Bucher
Ussuria Diener
Brühwiler et al., 2011a
Ussuridiscus
Shigeta &
Zakharov
Ussuriflemingites
Shigeta &Zakharov
Ussurijuvenites
Smyshlyaeva &
Zakharov
Vercherites
Brühwiler,Ware,
Bucher , Krystyn &
Goudemand
Wasatchites
Mathews
Wailiceras Brayard
& Bucher
Weitschaticeras
Brayard & Bucher
Xenoceltites Spath
Xenodiscoides
Spath
Xiaoqiaoceras
Brayard & Bucher
Shigeta et al., 2009
Brühwiler et al., 2010a
Shevyrev, 1986
Brayard and Bucher, 2008;
Brühwiler et al., 2010a
Zakharov et al., 2010
Shigeta et al., 2009
Smyshlyaeva and
Zakharov, 2011
Brühwiler et al., 2010b
Tozer, 1994
Brayard and Bucher, 2008
Brayard and Bucher, 2008
Brühwiler et al., 2010a
Brühwiler et al., 2011b
Brayard and Bucher, 2008;
Brühwiler et al., 2010c
Brayard and Bucher, 2008
8. Late Olenekian (Spathian) ammonoid genera. (Following Hyatt and Smith’s (1905) original proposal, we
continue to consider that the Neopopanoceras haugi Zone is earliest Anisian, in spite of the recent tendency to
expect it to be latest Olenekian (Bucher, 1989; Tozer, 1994; Shevyrev, 2006).
N
Genus
Main source of
N
Genus
Main source of
information
information
1
Albanites Arthaber
Shevyrev, 1986
66
Neomeekoceras
Tozer, 1994
Tozer
2
Anakashmirites Spath
Shevyrev, 1986
67
Nordophiceras
Zakharov, 1978
Popow
3
Anaxenaspis Kiparisova Zakharov et al., 2010
68
Nordophiceratoides
Guex et al., 2010
4
Arctomeekoceras
Zakharov, 1978
69
Nuetzelia Brühwiler,
Brühwiler et al., 2011a
Popow
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
Guex et al., 2010
Welter
9
Balkanites Ganev
Brayard et al., 2006
74
Mianwallites .
Brühwiler et al., 2011b
Brühwiler et al.
10 Bandoites Zakharov
Markevich and
Zakharov, 2004
11 Beatites Arthaber
Brayard et al., 2006
75
Nyalamites
Brühwiler et al., 2010a
27
Brühwiler, Bucher &
Goudemand,
Pallasites Renz &
Renz
Paradinarites Zhao
Paragoceras
Arthaber
12
Beneckeia Mojsisovics
Shevyrev, 1986
76
13
14
Beyrichites Waagen
Boreoceras Dagys &
Ermakova
(=Rudolftruempyiceras
Guex et al.)
Boreomeekoceras
Popow
Burijites Zakharov
Carteria Guex et al.
Shevyrev, 1986
Ermakova, 2002; Guex
et al., 2010; this study
77
78
Ermakova, 2002
79
Parahedenstroemia
Spath
This study
Zakharov, 1978
Guex et al., 2010
80
Shevyrev, 1986
Ceccaisculitoides Guex
et al.
Chioceras Renz &
Renz
Chiotites Renz & Renz
Columbites Hyatt &
Smith
Cordillerites Hyatt &
Smith
Coscaites Guex et al.
Guex et al., 2010
81
Paranannites Hyatt
& Smith
Paranorites Waagen
Shevyrev, 1986
82
Paranoritoides Guex
Shevyrev, 1986
Shevyrev, 1986
Zakharov, 1978
83
84
Parasibirites Popow
Parussuria Spath
Zakharov, 1978
Zakharov et al., 2010
Guex et al., 2010
85
Popovites Tozer
Tozer, 1994
Guex et al., 2010
86
Brayard et al., 2006
Guex et al., 2010
87
25
26
Cowboyiceras Guex et
al.
Dagnoceras Athaber
Dalmatites Kittl
Shevyrev, 1986
Shevyrev, 1986
88
89
27
28
29
Diaplococeras Hyatt
Dieneroceras Spath
Digitophyllites Zhao
Brayard et al., 2006
Zakharov et al., 2010
Brayard et al., 2006
90
91
92
30
31
Shevyrev, 1986
Brayard et al., 2006
93
94
32
33
Dinarites Mojsisovics
Doaboceras Boulin,
Bouyx, Termier &
Termier
Dorikranites Hyatt
Eodanubites Wang
Columbitidae gen.
nov.
Praesibirites Dagys
& Ermakova
Preflorianites Spath
Prefloarianitoides
Wang
Prenkites Arthaber
Procarnites Arthaber
Procolumbites
Astachova
Prohungarites Spath
Prosphingites
Mojsisovics
Shevyrev, 1986
Brayard et al., 2006
95
96
Shevyrev, 1986
Brayard et al., 2006
34
Eogymnites Spath
Shevyrev, 1986
97
35
Eophyllites Spath
Shevyrev, 1986
98
36
37
Epiceltites Arthaber
Epiceltitoides Guex
Shevyrev, 1986
Guex, 1978
99
100
38
Epiceltitoides Guex
Guex, 1978
101
Protropites Arthaber
Proptychitoides
Spath
Pseudoaspidites
Spath
Pseudodinarites
Hyatt
Pseudoceltites Hyatt
Pseudoharpoceras
Waagen
Pseudokymatites
Spath
Pseudoprosphingites
Shevyrev
Pseudosageceras
Diener
Pseudodinarites
Hyatt
Pseudosvalbardiceras Dagys
& Ermakova
Silberlingeria Guex
et al.
Stacheites Kittl
15
16
17
18
19
20
21
22
23
24
102
39
40
41
42
43
Eschericerafites Guex
et al.
Evenites Dagys &
Ermakova
Fengshanites Zhao
Guex et al., 2010
103
Ermakova, 2002
104
Brayard et al., 2006
105
Gaudemerites Guex et
al.
Glabercolumbites Guex
et al.
Guex et al., 2010
106
Guex et al., 2010
107
Brayard et al., 2006
Shevyrev, 1986
Guex et al., 2010
Brayard et al., 2006
Ermakova, 2002
Shevyrev, 1986
Brayard et al., 2006
Zakharov, 1978
Zakharov et al., 2010
Markevich and
Zakharov, 2004
Guex et al., 2010
Zakharov, 1978
Shevyrev, 1986
Brayard et al., 2006
Shevyrev, 1986
Brayard et al., 2006
Brayard et al., 2006
Markevich and
Zakharov, 2004
Zakharov et al., 2002
Brayard et al., 2006
Ermakova, 2002
Guex et al., 2010
Shevyrev, 1986
28
44
45
Hellenites Renz & Renz
Hemilecanites Spath
Zakharov, 1978
Brayard et al., 2006
108
109
46
47
Hyrcanites Shevyrev
Idahocolumbites Guex
et al.
Isculitoides Spath
Ivesgalleticeras Guex et
al.
Jeanbesseiceras Guex
et al.
Kazakhstanites
Shevyrev
Keyserlingites Hyatt
Shevyrev, 1986
Guex et al., 2010
110
111
Brayard et al., 2006
Guex et al., 2010
112
113
Guex et al., 2010
114
Shevyrev, 1968
115
Zakharov, 1978
116
48
49
50
51
52
54
55
56
57
58
59
60
61
62
1000
1001
1002
1003
1004
1005
1006
1007
1008
1009
1010
1011
1012
1013
1014
1015
1016
1017
1018
1019
1020
1021
Zakharov, 1978
Brayard et al., 2006
Zakharov, 1978
Shevyrev, 1986
Tozer, 1994
Kiparisovites
Astachova
Koninckitoides Dagys
& Ermakova
Khvalynites Shevyrev
Leiophyllites Diener
Mangyshlakites
Shevyrev
Marcouxia Guex et al.
Shevyrev, 1986
118
Ermakova, 2002
119
Shevyrev, 1986
Shevyrev, 1986
Shevyrev, 1986
120
121
122
Tjururpites Shevyrev
Tozericeras Guex
Tunglanites Zhao
Shevyrev, 1986
Guex, 1978
Shevyrev, 1986
Guex et al., 2010
123
Meropella Renz &
Renz
Metadagnoceras Tozer
Brayard et al., 2006
124
Kummel, 1969; Guex et
al., 2010; this study
Brayard et al., 2006
Tozer, 1994
125
Ussuriidae gen. nov.
(=“Ussurites Hyatt”)
Vickohlerites
Kummel
Xenoceltites Spath
Metahedenstroemia
Spath
Metasageceras Renz &
Renz
Shevyrev, 1986
126
?Xenodiscoides Spath
Brayard et al., 2006
127
Zenoites Renz &
Renz
Zhitkovites Zakharov
128
63
64
65
Shevyrev, 1986
Zakharov et al., 2008
Svalbardiceras
Frebold
Tapponnierites Guex
et al.
Tardicolumbites
Guex et al.
Tchernyschevites
Zakharov
Timoceras Dagys &
Ermakova
Tirolites Mojsisovics
117
53
Subcolumbites Spath
Subfengshanites
Zakharov
Subinyoites Spath
Subolenekites
Zakharov
Subvishnuites Spath
Sulioticeras Tozer
Guex et al., 2010
Guex et al., 2010
Markevich and
Zakharov, 2004
Ermakova, 2002
Shevyrev, 1986
Brayard and Bucher,
2008
Brayard et al., 2006
Brayard and Bucher,
2006a
Markevich and
Zakharov, 2004
Brayard et al., 2006
Metinyoites Spath
Waterhouse, 1996b
129 Ziyunites Wang
Monacanthites Tozer
Tozer, 1994
Neocolumbites
Zakharov, 1978
Zakharov
Ammonoid genera Deweveria, Neopopanoceras, Goricanites, Subhungarites, Pseudacrochordiceras, Inyoceras, and
Courtilloticeras, known from of the Neopopanoceras haugi Zone of the American tropical-subtropical realm
(Bucher, 1989; Guex et al., 2010), have not been included in Table 1, 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).
APPENDIX
Original data on brachiopod and ammonoid remains from the Lower Triassic of South Primorye and
Mangyshlak
1. South Primorye
The first geological studies in Russian southern Far East (Ussuri region, later named as Primorye) were
made by V.P. Margaritov, a graduate from the St.-Peterburg University, arriving in Vladivostok in 1880 as a teacher
of math. On the western coast of the Ussuri Gulf, near Shamara Bay (now Lazurnaya) he discovered some fossils,
including ammonoids and inarticulated brachiopods. His collection fell into the hands of the President of the
Russian Academy of Sciences, A.P. Karpinsky, a recognized expert in Late Palaeozoic ammonoids, who
distinguished Early Triassic ceratitid ammonoids among collected fossils. Later, Early and Middle Triassic marine
deposits in the environs of Vladivostok were studied by D.L. Ivanov, the chief of a geological team making
reconnaissance work for the construction of the Trans-Siberian railroad. He collected mollusc remains in the area of
the Shamara Bay and articulated brachiopod and mollusc shells on Russian Island. On the initiative of A.P.
Karpinsky, collections V.P. Margaritov and D.L. Ivanov were forwarded to Austrian palaeontologists K. Diener and
A. Bittner, who identified and described them (Diener, 1895; Bittner, 1899b). Later, Triassic cephalopods were
investigated by Kiparisova (1961), Burij and Zharnikova (1962, 1981), Zakharov (1968,1978,1997a), Zharnikova
29
1022
1023
1024
1025
1026
1027
1028
1029
1030
1031
1032
1033
1034
1035
1036
1037
1038
1039
1040
1041
1042
1043
1044
1045
1046
1047
1048
1049
1050
1051
1052
1053
1054
1055
1056
1057
1058
1059
1060
1061
1062
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(1972, 1985), Shigeta and Zakharov (Shigeta et al., 2009), and Smyshlyaeva (Smyshlyaeva, 2010; Smyshlyaeva and
Zakharov, 2011). However, after Bittner’s investigation, only a few Early Triassic brachiopod taxa, based on
material from South Primorye, was described by Dagys (1965, 1974) and Popov (Shigeta et al., 2009).
Observed ranges of ammonoids and brachiopods across the Induan-Lower Anisian interval in South
Primorye (Seryj-Tri Kamnya, Abrek, Konechnyj, Tobizin, Ayax-Balka, Zhitkov, Golyj, and Paris), summarized in
figures S1-S9 and Table S1, help to understand the main features of their recovery on specific level from the endPermian extinction.
1.1. Seryj-Tri Kamnya capes
The Seryj Cape-Tri Kamnya Cape section, exposed at the western Ussuri Gulf, is represented by the
Lazurnaya and Tobizin formations (Diener, 1895; Burij, 1959; Korzh, 1959; Kiparisova, 1961, 1972; Zakharov,
1968; 1996; Zakharov, 1997b; Markevich and Zakharov, 2004; Zakharov et al., 2010).
1.1.1. Brachiopod occurrences
Lissorhynchia sp. (Pontiidae), the first record of this genus from South Primorye, seems to be the oldest
Induan articulated brachiopod species found in the region. A single representative of this species was collected by T.
Kumagae in the lower part of the Lazurnaya Bay Formation (Gyronites subdharmus Zone, about 20 m above the top
of the underlying Early Induan Tompophiceras ussuriense Zone). Other Induan brachiopod fossils from this section
are represented by Lingula borealis Bittner. A few articulated brachiopods, mainly rhynchonellids, were collected in
the lower part of the Smithian Tobizin Formation (Euflemingites prynadai beds) (Fig. S1).
1.1.2. Ammonoid occurrences
Early Triassic ammonoids are significantly more diverse in the mentioned section, as well as in other South
Primorye sections, than associated brachiopods. Induan ammonoids from the Lazurnaya Bay Formation, exposed on
the western coast of Ussuri Gulf, are represented by a single genus (Tompophiceras), belonging to the Palaeozoictype family Dzhulfitidae, and five genera within the Mesozoic-type families Ophiceratidae, Proptychitidae,
Meekoceratidae, and Xenoceltitidae?. Early Olenekian (Smithian) ammonoids from the uppermost portion of the
Lazurnaya Bay Formation (Ussuriflemingites abrekensis (=”Gyronites separatus”) beds of the “Hedenstroemia”
bosphorensis Zone) and the lower portion of the Tobizin Cape Formation (Euflemingites prynadai beds of the
mentioned zone) in this section are more diverse, representing by 22 genera within Mesozoic-type families
Sageceratidae, Hedenstroemiidae, Aspenitidae, Ussuriidae, Paranannitidae, Clypeoceratidae, Arctoceratidae,
Meekoceratidae, Dieneroceratidae, and Flemingitidae (Fig. S1).
1.2. Abrek Bay
The Abrek Bay section, located at Strelok Strait and represented by the Lazurnaya Bay and Zhitkov Cape
formations, is worthy of consideration as one of the reference sections for Lower Triassic stratigraphy in South
Primorye (Burij, 1959; Kiparisova, 1961, 1972; Zakharov and Popov, 1999; Markevich and Zakharov, 2004;
Zakharov et al., 2002; Shigeta et al., 2009).
1.2.1. Brachiopod occurrences
In the lower part of the Induan Lazurnaya Bay Formation of the Abrek Bay section only inarticulated
brachiopods (Lingula borealis Bittner and Orbiculoidea sp.) have been discovered (Shigeta et al., 2009). Among
Late Induan brachiopods from the Lytophiceras-bearing beds of the Gyronites subdharmus Zone about five genera
of articulated brachiopods, including Abrekia, have been recognized (Dagys, 1974; Shigeta et al., 2009). All of them
belong to Mesozoic-type families (Wellerellidae, Pontisiidae, Rhynchonellidae, and Norellidae). Representatives of
the Abrekia (A. sulcata Dagys) have been discovered also in the lower Olenekian (Smithian) portion of the Zhitkov
Cape Formation (“Hedenstroemia” bosphorensis Zone, presumably Euflemingites prynadai beds) (Fig. S2).
1.2.2. Ammonoid occurrences
The Induan ammonoid succession in the Abrek Bay section consists of a single genus (Tompophiceras),
belonging to a Permian-type family Dhulfitidae, and 12 genera within five Mesozoic-type families (Proptychitidae,
Ophiceratidae, Meekoceratidae, Paranoritidae, and ?Paranannitidae). Among Smithian ammonoids of the mentioned
section 27 genera belonging to 15 Mesozoic-type families have been recognized (Fig. S2). They are following:
Sageceratidae, Paranannitidae, Aspenitidae, Xenoceltitidae, Proptychitidae?, Clypeoceratidae, Arctoceratidae,
Owenitidae, Inyoitidae?, Prionitidae, Kashmiritidae, Meekoceratidae, Dieneroceratidae, Flemingitidae, and
Palaeophyllitidae.
1.3. Konechnyj Cape
The Konechnyj Cape, located at Novik Bay on Russian Island, is represented by the Schmidt Formation (Markevich
and Zakharov, 2004).
At Konechnuj Cape, only a small portion, about 6 m thick, of the middle Olenekian (early Smithian)
Schmidt Cape Formation is exposed. No ammonoids, but abundant of articulated brachiopods were recently
collected there. The two brachiopod assemblages have been recognized in the section: (1) Hustedtiella planicostaSpiriferinida-Rhynchonellida-“Fletcherithyris” margaritovi-Zeilleriidae? below (occurs mainly in blocks) and (2)
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monospecies - “Fletcherithyris” margaritovi assemblage above. Both the assemblages correspond to the Tirolites
ussuriensis beds of the early Spathian Tirolites-Amphistephanites Zone in other sections at the Russian Island
(Markevich and Zakharov, 2004). Representatives of five genera within five Mesozoic-type families
(Rhynchonellidae, Neoretziidae, Spiriferinidae, Dielasmatidae, and Zeilleriidae?) were discovered in this locality.
1.4. Tobizin Cape
The Tobizin Cape Formation, exposed on the south part of Russian Island (Novyj Dzhigit Bay area) is
represented by the Lazurnaya Bay (basal part), Tobizin Cape and Schmidt Cape formations (Diener, 1895; Burij,
1959; Korzh, 1959; Zakharov, 1968; Buryi, 1979; Markevich and Zakharov, 2004).
1.4.1. Brachiopod occurrences
From the early Olenekian (Smithian) Tobizin Cape Formation only inarticulated brachiopod Lingula
borealis Bittner is known. The articulated brachiopods (Zeilleriidae? gen. and sp. nov. E and Spiriferinidae gen. and
sp. indet.) (Fig. S3) have been collected only from the upper part of the middle Olenekian (early Spathian) Schmidt
Cape Formation.
1.4.2. Ammonoid occurrences
Only early to middle Olenekian (Smithian-early Spathian) ammonoid succession, represented by 20 genera
belonging to 14 Mesozoic-type families is known there. These families are following: Aspenitidae, Ussuriidae,
Paranannitidae, Owenitidae, Meekoceratidae, Dieneroceratidae, Prionitidae, Tirolitidae, Melagathiceratidae,
Stephanitidae, Dinaritidae, Clypeoceratidae, Arctoceratidae, and Flemingitidae) (Fig. S3).
1.5. Ayax Bay-Balka Cape
The Ayax Bay-Balka Cape, exposed on the northern part of Russian Island, consists of Lazurnaya Bay,
Tobizin Cape and Schmidt Cape formations (Diener, 1895; Korzh, 1959; Kiparisova, 1961, 1972; Zakharov, 1968,
1978; 1997a; Buryi, 1979; Markevich and Zakharov, 2004).
1.5.1. Brachiopod occurrences
Only inarticulated brachiopod Lingula borealis Bittner and fragments of rhynchonellid and terebratulid
brachiopod shells were collected from the lower part (Induan-Smithian) of the section, which consists of the
Lazurnaya Bay and Tobizin Cape formations. Articulated brachiopods Lepismatina aff. mansfieldi (Girty) and
spiriferinids belonging to Mesozoic-type families Lepismatinidae and Spiriferinidae have been reported from the
lower part of the middle Olenekian (early Spathian) Schmidt Cape Formation(Fig. S4).
1.5.2. Ammonoid occurrences
At the top of the Induan conglomerate member of the Lazurnaya Cape Formation only bad preserved
Gyronites? sp. (Meekoceratidae) has been recognized. Among early to middle Olenekian ammonoids from the
uppermost portion of the Lazurnaya Bay Formation, Tobizin Cape and Schmidt Cape formations 22 ammonoid
genera belonging to 17 Mesozoic-type families have been listed (Zakharov, 1997). Among them are following:
Sageceratidae, Hedenstroemiidae, Ussuriidae, Paranannitidae, Arctoceratidae, Meekoceratidae, Dieneroceratidae,
Owenitidae, Inyotidae, Prionitidae, Xenoceltitidae, Tirolitidae, Melagathiceratidae, Stephanitidae, Doricranitidae?,
Dinaritidae, and Palaeophyllitidae (Fig. S4).
1.6. Schmidt Cape-Tchernyschev Bay
The Schmidt Cape-Tchernyschev Bay, located on the south-eastern part of Russian Island, is represented by
the Schmidt Cape and Zhitkov Cape formations (Burij, 1959; Korzh, 1959; Kiparisova, 1961, 1972; Zakharov,
1968, 1978; Buryi, 1979; Markevich and Zakharov, 2004).
1.6.1. Brachiopod occurrences
Schmidt Cape area is one of places with abundant early Spathian Mesozoic-type articulated brachiopods
(mainly in the Tirolites ussuriensis beds of the Tirolites-Amphistephanites Zone) (Fig. S5). In this part of the
mentioned section, there are five genera belonging to the following Mesozoic-type families: Spiriferinidae,
Neoretziidae, Dielasmatidae, and Zeilleriidae(?). Earliest Anisian spiriferinid and terebratulid brachiopods are
reported from the Ussuriphyllites amurensis Zone of the Tchernyschev Bay area (Fig. S5).
1.6.2. Ammonoid occurrences
Spathian ammonoids from the Schmidt and Zhitkov Cape formations of this section are represented by 21
genera belonging to 12 Mesozoic-type families (Sageceratidae, Paranannitidae, Khvalynitidae, Meekoceratidae,
Prionitidae, Columbitidae, Kazakhstanitidae, Xenoceltitidae, Tirolitidae, Dinaritidae, and Palaeophyllitidae) (Fig.
S5).
1.7. Zhitkov Cape
The Zhitkov Cape section, exposed on the northern part of Russian Island, is represented by the Tobizin
Cape, Zhitkov Cape and Karazin Cape formations (Burij, 1959; Korzh, 1959; Kiparisova, 1961, 1972; Zakharov,
1968, 1978; 1997a; Buryi, 1979; Markevich and Zakharov, 2004).
1.7.1. Brachiopod occurrences
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Majority of articulated brachiopods from this section was collected in the Tirolites ussuriensis beds of the
early Spathian Tirolites-Amphistephanites Zone (Schmidt Formation), others were collected in the late Spathian
Subfengshanites multiformis Zone (Fig. S6). Olenekian portion of the Zhitkov Cape section includes seven genera
belonging to six Mesozoic-type families (Rhynchonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae,
Dielasmatidae, and Zeilleriidae?).
1.7.2. Ammonoid occurrences
Smithian and Spathian ammonods of this section were collected in the upper portion of the Tobizin Cape
Formation (Anasibirites nevolini Zone) and Schmidt Cape (Tirolites-Amphistephanites Zone), and Zhitkov Cape
(Neocolumbites insignis and Subfengshanites multiformis zones) formations (Fig. S6). They are represented by 24
genera belonging to 13 families (Sageceratidae, Paranannitidae, Khvalynitidae, Isculitidae, Meekoceratidae,
Dieneroceratidae, Columbitidae, Kazakhstanitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Dinaritidae, and
Megaphyllitidae).
1.8. Golyj (Kom-Pikho-Sakho) Cape
The Golyj Cape section, exposed at eastern part of the Ussuri Gulf, consists of Lazurnaya Bay, Zhitkov
Cape and Karazin formations (Burij, 1959; Korzh, 1959; Zakharov, 1968; Buryi, 1979; Markevich and Zakharov,
2004).
1.8.1. Brachiopod occurrences
Articulated brachiopods in this section were discovered only in the middle Olenekian (early Spathian)
portion of the Zhitkov Cape Formation (Tirolites-Amphistephanites Zone). They are represented by five genera of
four Mesozoic-type families (Rhynchonellidae, Neoretziidae, Spiriferinidae, and Zeilleriidae?) (Fig. S7).
1.8.2. Ammonoid occurrences
Representatives of a single genus Gyronites (Mesozoic-type family Meekoceratidae) were reported from
the Induan Lazurnaya Bay Formation (Fig. S7). Early to late Olenekian ammonoids were discovered from the lower
(“Hedenstroemia” bosphorensis Zone), middle (Tirolites-Amphistephanites Zone) and upper (Neocolumbites
insignis Zone) portions of the Zhitkov Cape Formation. Olenekian ammonoid assemblage of the Golyj Cape section
consists of 24 genera belonging to 17 Mesozoic-type families: Sageceratidae, Hedenstroemiidae, Aspenitidae,
Ussuriidae, Paranannitidae, Khvalynitidae, Arctoceratidae, Owenitidae, Meekoceratidae, Inyoitidae, Prionitidae,
Columbitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Melagathiceratidae, and Flemingitidae.
1.9. Paris Bay
The Paris Bay section, located on the northern part of Russian Island, is represented by the Tobizin Cape, Schmidt
Cape, Zhitkov Cape and Karazin Cape formations (Burij, 1959; Korzh, 1959; Zakharov, 1968; Buryi, 1979;
Markevich and Zakharov, 2004).
1.9.1. Brachiopod occurrences
Articulated brachiopods from the Paris Bay section were collected from the early Spathian Schmidt
Formation (Tirolites-Amphistephanites Zone) and uppermost beds of the Late Spathian Zhitkov Formation
(Subfengshanites multiformis Zone). Spathian assemblage of articulated brachiopods consists of six genera
belonging to five Mesozoic-type families: Rhynchonellidae, Neoretziidae, Lepismatinidae, Spiriferinidae, and
Zeilleriidae? (Fig. S8).
1.9.2. Ammonoid occurrences
Ammonoids from the Paris Bay section are originated from the upper portion of the Smithian Tobizin
Formation (the upper part of the Anasibirites nevolini Zone), early Spathian Schmidt Formation (TirolitesAmphistephanites Zone), and late Spathian Zhitkov Formation (Neocolumbites insignis and Subfengshanites
multiformis zones). Olenekian (mainly Spathian) ammonoids from this section are represented by 18 genera within
12 Mesozoic-type families: Khvalynitidae, ?Proptychitidae (Proptychitoides), Arctoceratidae, Meekoceratidae,
Dieneroceratidae, Columbitidae, Xenoceltitidae, Keyserlingitidae, Tirolitidae, Stephanitidae, Dinaritidae, and
Hungaritidae (Fig. S8).
2. Mangyshlak, Dolnapa (Kazakhstan)
The first Triassic fossils in Mangyshlak were found in 1914 by M.V. Bayarunas, who identified and
described some ammonoids among them 22 years later (Bajarunas, 1936). Later, Early Triassic of Mangyshlak were
investigated by Astachova (1960), Shevyrev (1968), Gavrilova (1980, 1989, 2007), Balini (Balini et al., 2000) and
Zakharov (Zakharov et al., 2008). Before our investigation, Triassic brachiopods from Mangyshlak were described
by Dagys (1974) on the bases of A.A Shevyrev’s collection.
The reference section for the Lower Triassic in Mangyshlak, located at the Dolnapa Well area in Kara-Tau,
is represented by the Tartalinskaya, Karadzhatykskaya and Karaduanskaya formations.
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2.1. Brachiopod occurrences
Spathian brachiopod assemblage of the mentioned section consists of 8-11 genera belonging to eight
Mesozoic-type families (Pontisiidae, Norellidae, Diplospirellidae, Neoretziidae, Lepismatinidae, Dielasmatidae,
Antezeilleridae, and Plectoconchidae) (Fig. S9).
2.2. Ammonoid occurrences
Spathian ammonoids are represented by 23 genera belonging to 14 Mesozoic-type families (Sageceratidae,
Khvalynitidae, Dinaritidae, Prionitidae, Columbitidae, Kazakhstanitidae, Kashmiritidae, Xenoceltitidae,
Olenikitidae, Tirolitidae, Doricranitidae, Dinaritidae, Palaeophyllitidae, and Megaphyllitidae (Fig. S9)..
3. Discussion (Appendix)
The Induan of South Primorye (SP) is characterised by seven articulated brachiopod species: early Induan
Lissorhynchia sp. and six late Induan species, including Abrekia sulcata and new ones (Figs S1 and S2 and Table
S1). Early Olenekian brachiopods from SP show a slight reduction in species number, largely due to a variety of
facial conditions. Middle Olenekian brachiopods from SP (about eight species, including new ones) (Figs. S3-S9
and Table S1) are more diverse than those of Mangyshlak. In addition, information on late Olenekian brachiopods
from SP (about five species, including new ones) and Mangyshlak (12 species), as well as the published data on
Olenekian species from Eurasia (e.g., Chen, 1983; Xu and Liu, 1983; Xu and Grant, 1994; Chen et al, 2005) and
North America (e.g., Girty, 1927; Perry and Chatterton, 1979), illustrates a general trend in marked rising of
taxonomic diversity of Mesozoic-type brachiopods from the lower Induan through the upper Olenekian. The Induan,
lower, middle and upper Olenekian in SP are characterised by 20, 71, 11 and 30 species, respectively. Similar
changes in ammonoid succession, with highest diversification for the early Olenekian, occur in some other Tethyan
regions. According to published data (Chen et al., 2005) 13 articulated brachiopod lineages at familial level survived
the end-Permian mass extinction. As a result, at least 12 Palaeozoic type articulated brachiopod genera occur in the
Lower Triassic, mainly in the lower Griesbachian; at the same time the Permian-type Prelissorhynchia at
Mangyshlak was found by us in the first time in the upper Olenekian.
However, only four ammonoid lineages at family-superfamily level survived the mentioned mass
extinction; apparently only a single Palaeozoic-type ammonoid genus (Episageceras) is known from the Lower
Triassic (Zakharov, 1978).
References (Appendix)
Astachova, T.V., 1960. New Early Triassic ceratites of Mangyshlak. In: Markovskiy, B.P. (Ed.): Novyje vidy
drevnikh rasteniy i bespozvonochnykh SSSR (New species of fossil plants and invertebrates of the USSR).
Gosgeoltekhizdat, Moscow, pp. 139-159 (in Russian).
Bajarunas, M.V., 1936. Age of the Doricranites beds. Izvestiya Akademii Nauk SSSR, Seriya Geologicheskaya 4,
539-546 (in Russian).
Balini, M., Gavrilova, V.A., Nicora, A., 2000. Biostratigraphical revision of the classic Lower Triassic Dolnapa
section (Mangyshlak, west Kazakhstan). Zentralblatt für Geologie und Paläontologie I (1998), 1441-1462.
Bittner, A., 1899a. Himalayan fossils. Trias Brachiopoda and Lamellibranchiata. Palaeontologica Indica, 15 series 3
(2), 1-76.
Bittner, A., 1899b. Versteinerungen aus den Trias-Ablagerungen des Süd-Ussuri-Gebietes in der ostsibirischen
Küstenprovinz. Mémores du Comité Géologique 7 (4), 1-35.
Bogoslovskaya, M.F., Kuzina, L.F., Leonova, T.B., 1999. Classification and distribution of Late Paleozoic
ammonoids. In: Rozanov, A.Y. and Shevyrev, A.A. (Eds.), Fossil Cephalopods: Recent Advances in Their
Study. Izdatelstvo Akademii Nauk SSSR, Moscow, pp. 89-124 (in Russian).
Brayard, A., Bucher, H., 2008. Smithian (Early Triassic) ammonoid faunas from northwestern Guangxi (South
China): taxonomy and biochronology. Fossils and Strata 56, 1-179.
Brayard, A., Bucher, H., Escarguel, G., Fluteau, F., Bourquin, S., Galfetti,T., 2006a. The Early Triassic ammonoid
recovery: palaeoclimatic significance of diversity gradients. Palaeogeography, Palaeoclimatology,
Palaeoecology 239, 374-395.
Brayard, A., Escarguel, G., Bucher, H., 2006b. The biogeography of Early Triassic ammonoid faunas: clusters,
gradients, and networks. Geobios 40, 749-765.
Brühwiler, T., Brayard, A., Bucher, H., Guodun, K., 2008. Griesbachian and Dienerian (Early Triassic) ammonoid
faunas from northwestern Guangxi and southern Guizhou (South China). Palaeontology 51 (5), 1151-1180.
Brühwiler, T., Bucher, H., Goudemand, N., 2010a. Smithian (Early Triassic) ammonoids from Tulong, South Tibet.
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Brühwiler, T., Ware, D., Bucher, H., Krystyn, L., Goudemand, N., 2010b. New Early Triassic ammonoid faunas
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Brühwiler, T., Bucher, H., Krystin, L., 2011a. Middle and late Smithian (Early Triassic)ammonoids from Spiti
(India). Palaeontology, in press.
Brühwiler, T., Bucher, H., Ware, D., Hermann, E., Hochuli, P.A., Roohi, G., Rehman, K., Yaseen, A., 2011b.
Smithian (Early Triassic) ammonoids from the Salt Range, Pakistan. Palaeontology, in press.
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Bucher, H., 1989. Lower Anisian ammonoids from the northern Humboldt Range (northern Nevada, USA) and their
bearing upon the Lower-Middle Triassic boundary. Eclogae Geological Helvetiae 82 (3), 945-1002.
Buryj, G.I., 1979. Nizhnetriasovye konodonty Yuzhnogo Primorya (Lower Triassic conodonts of South Primorye).
Nauka, Moscow, 143 pp. (in Russian).
.Burij, I.V., 1959. Stratigrafiya triasovykh otlozhenij yuzhnogo Primorya (Stratigraphy of Triassic sediments of
south Primorye). Trudy Dalnevostochnogo Politekhnicheskogo Instituta 54 (1), 3-34 (in Russian).
Burij, I.V., Zharnikova, N.K., 1962. New Triassic ceratitid species of Far East. In: Shvedov, N.A. (Ed.), Sbornik
statei po paleontologii I biostratigrafii (Collected papers on palaeontology and biostratigraphy). NauchnoIssledovatelskij Institut Geologii Arktiki Ministerstva Geologii I Okhrany Nedr SSSR, pp. 78-92 (in Russian).
Burij, I.V., Zharnikova, N.K., 1981. Ammonoids from the Tirolites Zone of South Primorye. Paleontologicheskij
Zhurnal 3, 61-69 (in Russian).
Chen,Y.M., 1983. New advance in the study of Triassic brachiopods in Tulong district of Nyalam Country, Tibet.
Contribution to the Geology of the Qinghai-Xizang Plateau 11, 145-156.
Chen, Z.Q., Shi, G.R., 1999. Revision of Prelissorhynchia Xu and Grant, 1994 (Brachiopoda) from the Upper
Permian of South China. Proceeding of the Royal Society of Victoria 111, 15-26.
Chen, Z.Q., Shi, G.R., Shen, S.Z., Archbold, N.W., 2000. Tethyochonetes gen. nov. (Chonetida, Brachiopoda) from
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