Early Triassic recovery of the brachiopod faunas from the end-Permian mass extinction: A global review

Early Triassic recovery of the brachiopod faunas from the end-Permian mass extinction: A global review

Palaeogeography, Palaeoclimatology, Palaeoecology 224 (2005) 270 – 290 www.elsevier.com/locate/palaeo Early Triassic recovery of the brachiopod fauna...

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Palaeogeography, Palaeoclimatology, Palaeoecology 224 (2005) 270 – 290 www.elsevier.com/locate/palaeo

Early Triassic recovery of the brachiopod faunas from the end-Permian mass extinction: A global review Zhong-Qiang Chena,*, Kunio Kaihob, Annette D. Georgea a

School of Earth and Geographical Sciences, University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia b Institute of Geology and Paleontology, Tohoku University, Sendai 980-8578, Japan Received 22 December 2003; received in revised form 23 November 2004; accepted 23 March 2005

Abstract Brachiopod faunas in Lower Triassic deposits from Spitzbergen, Primorye of Russia, Japan, Mangyslak of Kazakhstan, Alpine Europe, the Caucasus, the Middle East, the Himalaya, South China, New Zealand, and western USA are taxonomically and stratigraphically reviewed. They comprise survivors of the end-Permian extinction and taxa originating in the Early Triassic. Origination of the Mesozoic-type taxa represents the recovery of Early Triassic brachiopods. The initial recovery of brachiopod faunas began in the late Griesbachian, and the time interval between the end of extinction and the onset of the recovery of brachiopods is much shorter than previously suspected. The Early Triassic recovery of brachiopod faunas is characterized by widespread brachiopod dispersal, multiprovincialism, and the presence of rare Lazarus genera at that time. Taxonomic selectivity of the recovery brachiopod faunas favors the rhynchonellids. The re-population of postextinction brachiopods varies geographically: there is a preference for regions either previously barren of latest Permian taxa or regions where the pre-extinction Changhsingian and surviving brachiopods are very rare. This unique biogeographic selectivity is probably partly responsible for the impoverished nature of brachiopod faunas in the Triassic (even Mesozoic) oceans. Five intervals of extinction, survival, survival–recovery, recovery–dispersal, and radiation are recognized based on variations in brachiopod faunas from the latest Permian to the Early Triassic. A dramatic reduction in brachiopod diversity at the end-Permian mass extinction is followed by several stepwise declines in diversity in the survival interval, the time during which the surviving brachiopods are dominated by geographically widespread generalist faunas that adapted to a wide variety of environments. The survival interval was followed by a slow re-population of new lineages dominated by progenitor taxa. D 2005 Elsevier B.V. All rights reserved. Keywords: Early Triassic; End-Permian mass extinction; Recovery; Brachiopod faunas

1. Introduction * Corresponding author. Tel.: +61 893801924; fax: +61 893801037. E-mail address: [email protected] (Z.-Q. Chen). 0031-0182/$ - see front matter D 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2005.03.037

The end-Permian mass extinction has been considered as the most widespread biotic crisis of the

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Phanerozoic, eliminating about 90% or more of all marine species (Erwin, 1993, 1994). Many questions remain about the true magnitude and causes of this dramatic extinction. Similarly, interpretation of the delayed faunal recovery following this extinction remains highly disputed. The final recovery of postPaleozoic marine faunas is generally believed to have occurred in the early Middle Triassic (Anisian), about 5 million years after the extinction event (Erwin, 1993, 1994, 1998). A number of recent studies of this recovery, from the US (Schubert and Bottjer, 1995) and northern Italy (Twitchett and Wignall, 1996; Twitchett, 1999), also showed similar patterns, which are also supported by gastropod data (Erwin, 1996, 1998; Erwin and Pan, 1996). All these studies concluded that simple, cosmopolitan, opportunistic generalists, and low-diversity paleocommunities were characteristic of the Early Triassic oceans. However, these patterns have been mainly derived from gastropod data (Erwin, 1996; Erwin and Pan, 1996), reef-building organisms (Hallam, 1991), crinoids (Foote, 1996), and ichnofossils (Twitchett, 1999). So far there has not been any study on the post-extinction recovery of the global brachiopod faunas. Delayed recovery patterns have been generally applied to brachiopod faunas which have shown extreme delay in their full recovery in the aftermath of the event (Carlson, 1991; Hallam, 1991; Erwin, 1993, 2001). However, timing of the onset of the brachiopod recovery has long remained unknown, although more recently Chen et al. (2002) estimated that its initial phase took place within one million years after the end-Permian extinction, based on new evidence from the well-dated Lower Triassic succession of the Meishan section, South China. In addition, Rong and Shen (2002) carried out a comparative analysis of the end-Permian and end-Ordovician brachiopod mass extinctions and recoveries in South China that provided some insights into regional post-extinction recovery patterns of the Triassic brachiopod faunas, but their data on Early Triassic brachiopods remain incomplete. For example, Feng and Jiang (1978) illustrated TPiarorhynchellar gujiaoensis from the Olenekian of the Guizhou Province. Xu and Grant (1994) reported Spiriferina sp. from the Dienerian (late Induan) from the Xiushan section of Daye City, Hubei Province. Chen et al. (2002) described a new brachiopod genus with clear Mesozoic aspects from

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the late Griesbachian of the Meishan section of Zhejiang Province (discussed further below). Our study reveals that Rong and Shen’s (2002) conclusion that the South Chinese brachiopod faunas recovered in the Middle Triassic after a long bleak interval during the Early Triassic does not present a complete view of post-extinction recovery of Triassic brachiopods in South China. Thus, the global recovery pattern of brachiopod faunas after the greatest extinction recorded in geologic history remains obscure. Also, published data and current available materials tend to indicate that the recovery pattern derived from brachiopod shows some clear differences from other fossil groups (e.g., bivalves, gastropods, and echinoids). (1) Brachiopods were the most diverse fauna in the Permian oceans, but were also the second largest group of victims among the main marine fossil groups that suffered the end-Permian mass extinction (Raup, 1979; Carlson, 1991; Erwin, 1993). The brachiopod recovery pattern should therefore have its own unique feature. (2) Brachiopods had the most abundant representatives that survived the end-Permian mass extinction event (Sheng et al., 1984). The relationships among the pre-extinction brachiopods, survivors and post-extinction recovery and radiating faunas are crucial to understand the process and mechanisms of brachiopod recovery after the event. As such, the brachiopod faunas are able to more precisely indicate biotic factors at extinction, survival, and recovery intervals summarized by Kauffman and Erwin (1995). (3) There are rare Lazarus genera (Jablonski, 1986) in marked contrast to a few Lazarus genera in other groups (e.g. gastropods), which commonly reappear near the close of the recovery interval (Erwin, 1998). Consequently, the recovery model of brachiopods is much less influenced by Lazarus effect at generic level, although the Lazarus effect affected this fossil group at familial level to some extent (also see below). (4) Early Triassic brachiopods mainly comprise endemic elements and appear to record strong provinciality, in sharp contrast to the current hypothesis based on the observation of other fossil groups.

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The aims of this study are: (i) to review Early Triassic brachiopod faunas worldwide by revising and updating their taxonomy and biostratigraphy; (ii) to provide taxonomic and paleobiogeographic perspectives on the Triassic recovery of brachiopods by analyzing taxonomic and biogeographic changes in the faunas through the Early Triassic; and (iii) generate a model summarizing the changes in diversity of brachiopod faunas in the extinction, survival, and recovery intervals.

2. Global view of Early Triassic brachiopods The Early Triassic was generally unfavorable for brachiopods because many of the major groups that existed in the Permian suffered virtual extinction in the end-Permian extinction event (Carlson, 1991). Lower Triassic brachiopod faunas are, therefore, rare and sparse worldwide (Dagys, 1974; Hoover, 1979; Ager, 1988; Ager and Sun, 1988). Despite the magnitude of the extinction, however, relict

faunas of Permian affinity did survive, and Triassic opportunistic species flourished locally (Fig. 1; Tables 1 and 2). As such, Lower Triassic brachiopods consist of two types, the TMesozoic-typer (Ager, 1988) also designated TTriassic-liker (Liao, 1979, 1980; Sheng et al., 1984), and the TPaleozoictyper (Ager, 1988), also termed TPermian-typer (Waterhouse, 1976) or TPermian-liker (Liao, 1979, 1980; Sheng et al., 1984). The latter types usually extend from the strata below the end-Permian mass extinction horizon, calibrated to the base of Bed 25 in the Meishan section (Jin et al., 2000; Kaiho et al., 2001), which coincides with the base of the conodont Clarkina meishanensis Zone or the lower Otoceras Zone in the Meishan section in South China (Mei et al., 1998; Yin et al., 2001), and persist into the postextinction Changhsingian and Lower Triassic strata (Table 1). These brachiopod taxa are also treated here as survivors of the end-Permian mass extinction (for more details see Chen et al., 2005-this volume). Nonetheless, since the latest definition of the P/T boundary uses the first occurrence of the conodont

1 2

4 3

250 Ma Early Triassic

12

15

South China

Paleotethys

13 11

5

14 Pangea

10 9 6 8

Surviving brachiopods

7

16

Mesozoic-type brachiopods

Fig. 1. Biogeographic distribution of the recovery faunas in the Early Triassic (base map after Ziegler et al., 1998). 1 = Spitsbergen; 2 = Primorye of Russia; 3 = Southwest Japan; 4 = northeastern Japan; 5 = South China; 6 = Selong, Tulong, and Lhasa sections of Tibet; 7 = Nepal; 8 = Kashmir; 9 = Salt Rang, Pakistan; 10 = the Central Oman Montains of Oman; 11 = the Caucasus of Armenia; 12 = Mangyslak region of Kazakhstan; 13 = Dobrogea of Romania and Stara Planina of Bulgaria; 14 = Dolomites of northern Italy; 15 = Idaho of western USA; 16 = Perth Basin of Western Australia. The Paleo-Tethys Margin Province includes Primorye of Russia, southwestern Japan, Mangyslak of Kazakhstan, Romania– Bulgaria of Alpine Europe and the Himalayan region. South China and western USA–Spitzbergen represent independent provinces, respectively.

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Table 1 Global distributions of the Permian relict brachiopods in the Early Triassic Species

Family

Superfamily

Age

Locality

Principal authors

Lingula

Lingulidae

Linguloidea

Induan–Olenekian

South China

Lingula

Lingulidae

Linguloidea

Olenekian

Western USA

Lingula Lingula Lingula Lingula Orbiculoidea Orbiculoidea Acosarina

Lingulidae Lingulidae Lingulidae Lingulidae Discinidae Discinidae Schizophoriidae

Linguloidea Linguloidea Linguloidea Linguloidea Discinioidea Discinioidea Enteletoidea

Griesbachian Griesbachian Olenekian Olenekian Griesbachian Olenekian Griesbachian

Salt Range Northern Italy Western Australia northeast Japan South China northeast Japan South China

Tethychonetes

Rugosochonetidae

Chonetoidea

Griesbachian

South China

Paryphella Cathaysia Spinomarginifera

Productellidae Productellidae Productellidae

Productelloidea Productelloidea Productelloidea

Griesbachian Griesbachian Griesbachian

South China South China South China

Spinomarginifera

Productellidae

Productelloidea

Griesbachian

Salt Range

Retimarginifera Retimarginifera Pustula Orbicoelia Orbicoelia Orbicoelia Orbicoelia Paracrurithyris

Productellidae Productellidae Echinoconchidae Ambocoellidae Ambocoellidae Ambocoellidae Ambocoellidae Ambocoellidae

Productelloidea Productelloidea Productelloidea Ambocoellioidea Ambocoellioidea Ambocoellioidea Ambocoellioidea Ambocoellioidea

Smithian Griesbachian Griesbachian Griesbachian Spathian Griesbachian Griesbachian Griesbachian

Nepal Kashmir Kashmir South China Romania Oman Salt Range South China

Paraspiriferina Prelissorhynchia

Paraspiriferinidae Pontisiidae

Pennospiriferinoidea Wellerelloidea

Griesbachian Griesbachian

South China South China

Prelissorhynchia Araxathyris Spirigerella

Pontisiidae Athyrididae Athyrididae

Wellerelloidea Athyridoidea Athyridoidea

Griesbachian Griesbachian Griesbachian

Oman South China Nepal

Liao, 1979, 1980; Yang et al., 1987; Xu and Grant, 1994 Newell and Kummel, 1942; Rodland and Bottjer, 2001 Rowell, 1970 Broglio Loriga et al., 1988 Dickins and McTavish, 1963 Murata, 1973 Liao, 1980 Murata, 1973 Sheng et al., 1984; Xu and Grant, 1994 Liao, 1979, 1980, Chen et al., 2000 Liao, 1980, 1984 Chen, unpublished data Liao, 1979, 1987; Chen and Shi, 1999 Kummel and Teichert, 1970; Grant, 1970 Waterhouse, 1978, 1994 Shimizu, 1981 Shimizu, 1981 Liao, 1980; Yang et al., 1987 Iordan, 1993 Twitchett et al., 2004 Grant, 1970 Liao, 1979, 1987; Xu and Grant, 1994 Xu and Grant, 1994 Liao, 1979, 1987; Chen and Shi, 1999 Chen, unpublished data Yang et al., 1987 Waterhouse, 1978; Waterhouse and Shi, 1991

Hindeodus parvus as a chronostratigraphic marker (Yin et al., 2001), it implies that the current P/T boundary is higher than the horizon of the ammonoid Otoceras Zone, previously used as a marker representing the beginning of the Triassic (e.g., Sheng et al., 1984; Broglio Loriga et al., 1988). Thus, in accordance with the re-definition of the P/T boundary (Yin et al., 2001), many faunas previously thought to be of earliest Triassic are now Ttechnicallyr of latest Permian (Chen et al., 2005-this volume). In such case, the relict Permian brachiopods in Lower Triassic strata are sparse, except for the extraordinarily abundant, and widespread disaster taxon Lingula (Rodland and

Bottjer, 2001). Specimens of this genus are mainly distributed in the Lower Triassic of northeastern Japan, South China, the Himalayan region (Salt Range, Nepal, Kashmir), Western Australia, Oman, Alpine Europe (Romania and Bulgaria), northern Italy, and western USA (Fig. 1). Stratigraphically, the surviving brachiopods are often associated with the conodont Hindeodus parvus and Isarcicella isarcica Zones or the ammonoid Otoceras and Ophiceras Zones (Yin et al., 2001). Except for the inarticulate brachiopods Lingula and Orbiculoidea (Table 1), most of the survivors narrowly escaped the end-Permian disaster, but became extinct

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Table 2 Global distributions of the Mesozoic-type brachiopods Species

Family

Superfamily

Age

Locality

Authors

Lepismatina mansfieldi Lepismatina mansfieldi Lepismatina sp. Spiriferina sp. Abrekia sulcata Abrekia cf. procreatrix Abrekia procreatrix Akrekia chaqupuensis Meishanorhynchia meishanensis Nudirostralina subtrinodosi Paranorellina parisi P. mangyshlakensis P. dulongdeqingensis Piarorhynchella triassica Piarorhynchella sp. bP.Q gujiaoensis bRhynchonellaQ triassicus

Lepismatinidae Lepismatinidae Lepismatinidae Spiriferinidae Rhynchonellidae Rhynchonellidae Rhynchonellidae Rhynchonellidae Rhynchonellidae Rhynchonellidae Norellidae Norellidae Norellidae Norellidae Norellidae Norellidae Norellidae

Pennospiriferinoidea Pennospiriferinoidea Pennospiriferinoidea Spiriferinoidea Rhynchonelloidea Rhynchonelloidea Rhynchonelloidea Rhynchonelloidea Rhynchonelloidea Rhynchonelloidea Norelloidea Norelloidea Norelloidea Norelloidea Norelloidea Norelloidea Norelloidea

Induan Induan Olenekian Induan Induan Induan Olenekian Olenekian Induan Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian

Primorye, Russia Mangyslak, Kazakhstan Southwest Japan Daye, South China Primorye, Russia Caucasus, Armenia Himalayan region Lhasa, Tibet Meishan, South China Tulong, Tibet Primorye, Russia Mangyslak, Kazakhstan Lhasa, Tibet Primorye, Russia Selong, Tibet Guizhou, South China Idaho, western USA

Lissorhnychia wesca L. lielonggouensis Laevorhynchia tenuis Spirigerinella pygmaea Spirigerellina pygmaea Spirigerellina pygmaea Hustedtiella? spitzbergensis Hustedtiella planicosta Hustedtiella sp. Hustedtiella cf. planicosta Neoretzia fuchsi Neoretzia sp. Retzioidea gen. and sp. indet. Obnixia wittenburgi Obnixia thaynesiana Periallus woodsidensis P. aff. woodsidensis Fletcherithyris? margaritovi F.? margaritovi F.? margaritovi F.? aff. margaritovi Rhaetina incurvirostra Portneufla episulcata Protogusarella smithi

Pontisiidae Pontisiidae Wellerellidae Diplospirellidae Diplospirellidae Diplospirellidae Neoretziidae Neoretziidae Neoretziidae Neoretziidae Neoretziidae Neoretziidae Aulacothyroideidae Aulacothyroideidae Aulacothyroideidae Aulacothyroideidae Dielasmatidae Dielasmatidae Dielasmatidae Dielasmatidae Dielasmatidae Cryptacanthiidae Cryptacanthiidae

Wellerelloidea Wellerelloidea Wellerelloidea Athyridoidea Athyridoidea Athyridoidea Retzioidea Retzioidea Retzioidea Retzioidea Retzioidea Retzioidea Retzioidea Cryptonelloidea Cryptonelloidea Cryptonelloidea Cryptonelloidea Dielasmatoidea Dielasmatoidea Dielasmatoidea Dielasmatoidea Dielasmatoidea Dielasmatoidea Dielasmatoidea

Induan Olenekian Induan Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Induan Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian Olenekian

Caucasus, Armenia Lhasa, Tibet Guizhou, South China Primorye, Russia Mangyslak, Kazakhstan Dobrogea, Romania Spitzbergen Primorye, Russia Southwest Japan Dobrogea, Romania Tulong, Tibet Selong, Tibet Daye, South China Spitzbergen Idaho, western USA Idaho, western USA Idaho, western USA Primorye, Russia Mangyslak, Kazakhstan Dobrogea, Romania Idaho, western USA Southwest Japan Idaho, western USA Idaho, western USA

Vex semisimplex

Terebratulidae

Terebratuloidea

Olenekian

Idaho, western USA

Dagys, 1974 Dagys, 1974 Dagys, 1974 Xu and Grant, 1994 Dagys, 1974 Dagys, 1974 Bittner, 1899 Sun et al., 1981 Chen et al., 2002 Chen, 1983 Dagys, 1974 Dagys, 1974 Sun et al., 1981 Dagys, 1974 Chen’s unpubl. data Feng and Jiang, 1978 Perry and Chatterton, 1979 Dagys, 1974 Sun et al., 1981 Shen and He, 1994 Dagys, 1974 Dagys, 1974 Iordan, 1993 Dagys, 1974 Dagys, 1974 Dagys, 1974 Iordan, 1993 Chen, 1983 Chen’s unpubl. data Xu and Grant, 1994 Dagys, 1974 Hoover, 1979 Hoover, 1979 Hoover, 1979 Dagys, 1974 Dagys, 1974 Iordan, 1993 Hoover, 1979 Dagys, 1974 Hoover, 1979 Perry and Chatterton, 1979 Hoover, 1979

shortly thereafter (Fig. 2; see also Chen et al., 2005this volume). In addition, Retimarginifera is reported to persist into the Smithian of the early Olenekian (Waterhouse and Shi, 1991), whereas Orbicoelia is reported in the Spathian of the late Olenekian (Iordan, 1993), but their occurrences remain unconfirmed. In

summary, the Lower Triassic survivors may show ecologic adaptations to the harsh environments of the post-extinction oceans, but they do not represent the post-extinction recovery of brachiopod faunas because they extend from the pre-extinction Changhsingian (Permian).

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Stratigraphic ranges

MF 1 interval

Orbiculoidea

MF 2 interval

Paranorellina Piarorhynchella "Rhynchonella" Spirigerinella Hustedtiella Neoretzia Fletcherithyris Periallus Obnixia Rhaetina Portneufla Protogusarella Ves

Lissorhynchia Lepismatina Nudirostralina

Lingula

INDUAN L. PERMIAN

Changhsingian Griesbachian

Dienerian

Pustula

Smithian

Orbicoelia Paracrurithyris Paraspiriferina Araxathyris Prelissorhynchia Spirigerella Meishanorhynchia Laevorhynchia Spiriferina Retzioidea gen. and sp. indet. Abrekia

Retimarginifera

Spathian

Acosarina Tethyochonetes Paryohella Cathaysia Spinomarginifera

OLENEKIAN

Stages

MF 3 interval

275

end-Permian mass extinction

Fig. 2. Stratigraphic distribution of the global Early Triassic brachiopod genera. Note that these genera persisting from the pre-extinction strata are the survivors of the end-Permian event, while those originated in the Early Triassic represent the recovery brachiopods after the end-Permian event. For explanation of Stages MF 1 to MF 3 see Fig. 5.

True Mesozoic-type brachiopods have been reported mainly beginning from the Induan and Olenekian in the regions shown in Fig. 1. It is also possible that some Mesozoic-type brachiopods occur in the Lower Triassic of Oman, the Caucasus region, and New Zealand. 2.1. Spitzbergen Dagys (1974, p. 249) listed Hustedtiella? spitzbergensis and TTerebratular wittenburgi from the Olenekian (Table 2) of the Spitzbergen region. Later, Dagys (1993) re-assigned TT.r wittenburgi to Obnixia Hoover (1979). 2.2. South Primorye, Russia In the southern Primorye region of Russia, Dagys (1965, 1974) described seven monospecific genera. The Induan elements comprise Abrekia sulcata and Lepismatina (= Costispiriferina) mansfieldi. The Olenekian forms consist of Piarorhynchella triassica, Paranorellina parisi, Spirigerinella pygmaea, Hus-

tedtiella planicosta and Fletcherithyris margaritovi. Carter et al. (1994, p. 369) considered Costispiriferina (Dagys, 1974) as a junior synonym of Lepismatina (Wang, 1955) and assigned the genus to the family Lepismatinidae of the superfamily Pennospiriferinoidea. Although Dagys (1974) considered that his genus is a representative of the superfamily Spiriferinoidea, Carter et al.’s (1994) view is followed here. 2.3. Japan Dagys (1974, p. 250) reported Lepismatina sp., Fletcherithyris aff. margaritovi and Hustedtiella sp. from the Lower Triassic Hirobatake Formation of the Maizuru zone, southwestern Japan. This brachiopod fauna is associated with abundant bivalves and the ammonoid Paratirolites sp., which would indicate an Olenekian age (Nakazawa, 1961). In addition, Ichikawa (1951) listed Spiriferina cf. fragilis Schlotheim, S. cf. stracheyi, Spiriferina sp., and Terebratula sp. from the Fukkoshi Formation (lower Inai Group) of the Kitakami Mountain region of northeastern Japan. Besides the above brachiopods, the Fukkoshi Formation

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also contains the ammonoids Leiophyllites aff. pitamaha (Diener), L. aff. pradyumna Diener, L. sp., Danubites aff. ambika Diener, Balatonites cf. kitakamicus (Diener) (Murata, 1973). Bando (1968) assigned the Fukkoshi Formation to the latest Scythian (Early Triassic) or early Anisian (early Middle Triassic) based on correlations with the ammonoid fauna. However, a more recent study (Waterhouse, 2002) indicates that Leiophyllites is a zonal genus characteristic of the middle Anisian, and Balatonites indicates a Pelsonian age (early Middle Triassic) in the southern Primorye region of Russia (Waterhouse, 2002, p. 68). Thus, the Fukkoshi Formation and its faunas are Middle Triassic in age. In the same region, Murata (1973) described Lingula sp., Orbiculoidea cf. sibirica, and Orbiculoidea sp. from the Olenekian Osawa Formation. We should also point out that Orbiculoidea is one of the common survivors of the end-Permian extinction event in other regions (Table 1; see also Chen et al., in this issue), and Lingula is a disaster taxon of the event (Rodland and Bottjer, 2001). 2.4. Mangyslak, Kazakhstan Lepismatina mansfieldi is reported from the Induan succession of the Mangyslak Peninsula of the Caspian Sea, Kazakhstan (Dagys, 1974), whereas Piarorhynchella mangyshlakensis, Spirigerellina pygmaea, and Fletcherithyris margaritovi are described from the Olenekian of the same region (Table 2).

2.6. Central Oman Mountains, Oman In central Oman, Krystyn et al. (2003) reported unnamed ambocoelid spiriferids and rhynchonellids from the bioclastic limestone unit of late Griesbachian age in the Wadi Wasit area. Later, Twitchett et al. (2004) referred these small, smooth, spiriferid specimens to Crurithyris extima. Nonetheless, despite the fact that the specimens from Oman externally suggest either Crurithyris or Orbicoelia, they lack a median sulcus (or groove) in both valves (Twitchett, personal communication, 2004). As we specified in the case of the specimens from the Caucasus (Section 2.5), presence of the median sulcus (or groove) is one of the most important criteria distinguishing Crurithyris from Orbicoelia (see Waterhouse and Gupta, 1983; Chen et al., 2005-this volume). Thus, the Oman ambocoelid specimens should be assigned to Orbicoelia, which are known as survivors of the endPermian event (see above). Furthermore, we studied the specimens belonging to the rhynchonellids, and identified them to be identical to Prelissorhynchia pseudoutah Huang (1933), which is also one of the most common survivors of the end-Permian extinction event in South China (Xu and Grant, 1994; Chen and Shi, 1999; see also Table 1). Based on these data, we can infer that upper Griesbachian brachiopods of the central Oman Mountains are survivors of the endPermian mass extinction. 2.7. Romania–Bulgaria

2.5. Caucasus, Armenia Dagys (1974) reported Neowellerella wesca, Crurithyris? extima, and Abrekia cf. procreatrix from the Induan of the Caucasus areas, Armenia. Note that Neowellerella is a junior synonym of Lissorhynchia Yang and Xu (1966) (Xu and Grant, 1994; see below), a genus which is usually associated with the Induan. On the other hand, the species C.? extima, first reported from the P/T boundary beds of the Salt Range in Pakistan (Grant, 1970), is better assigned to Orbicoelia than Crurithyris because they lack a median sulcus in both valves and other morphological differences (Waterhouse and Gupta, 1983; Chen et al., 2005-this volume). In addition, the age determination of this fauna from the Caucasus areas of Armenia is also questionable, as will be discussed below.

Four brachiopod species, Spirigerellina pygmaea Dagys, Fletcherithyris margaritovi (Bittner), Orbicoelia extima (Grant), and Hustediella cf. planicosta Dagys, have been reported from the Lower Triassic Series of the northern Dobrogea area of Romania, and the Stara Planina area of Bulgaria (Mirauta et al., 1984; Iordan, 1993), in the western bank regions of the Black Sea. Among these taxa, Orbicoelia extima is a known survivor of the end-Permian event, as it has been demonstrated by its occurrence in other regions (e.g., Grant, 1970, Broglio Loriga et al., 1988). In contrast, in Romania, this brachiopod fauna is recorded in the Werfenian Platy Limestone Formation, which Iordan (1993) assigned to the Spathian (late Olenekian). Moreover, Iordan (1993) pointed out that the above brachiopods are character-

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istic of the Induan and the Olenekian as well. Because Orbicoelia extima is confined to the Griesbachian (early Induan) in the Salt Range of Pakistan (Grant, 1970; Wignall et al., 1996), and the Wadi Wasit area of Oman (Krystyn et al., 2003; Twitchett et al., 2004), hence, the Platy Limestone Formation may be considered to range from the Induan to the Olenekian in age. 2.8. Himalayan region In the Himalayan region Bittner (1899, p. 9, Plate 1, Fig. 12) first described Rhynchonella (Norella) procreatrix from the Otoceras beds (Lower Triassic) located in the northwestern Kiunglung area of the southwestern Niti Pass, and the Shalshal section of the Rimkin–Paiar area. Bittner’s species was subsequently reassigned to the new genus Abrekia (Dagys, 1974). In the Salt Range of Pakistan, Orbicoelia extima (Grant, 1970) is abundant in the middle and upper units of the Kathwai Member, which are constrained within the upper part of the Otoceras woodwardi Zone (or the conodont Hindeodue parvus Zone), and the Ophiceras Zone (Wignall et al., 1996). Thus, the brachiopod species Orbicoelia extima found at that locality is Griesbachian in age. This species, however, is a common survivor of the end-Permian event, as discussed above. Another pertinent study in the area revealed that some rhynchonellids are present in the upper Kathwai Member in the Chhidru section of the Salt Range (Wignall et al., 1996). These rhynchonellids may represent the Mesozoic-type elements, although they remain unnamed. In addition, Waterhouse and Shi (1991) found Spirigerella sp. in Nepal where it is associated with a fauna of the ammonoid Otoceras woodwardi Zone, which is correlated with the conodont H. parvus Zone (Waterhouse, 2002), whereas Retimarginifera was found in the Smithian (early Olenekian) of the Sabche member (Waterhouse and Shi, 1991; Waterhouse, 2002). Waterhouse (1994, p. 16) reported Spiriferella rajab and Neospirifer from the type section of the lowest Triassic Pangjang Formation, but he considered these brachiopods as being possibly derived from the underlying Permian strata. If the presence of these brachiopods is confirmed in these younger strata, they should be survivors of the end-Permian event.

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In the northern Himalayan region, Chen (1983) reported Nudirostralina subtrinodosi and Neoretzia fuchsi from the Lower Triassic of the Tulong section in Nyalam County, southern Tibet. The associated ammonoids Owenites, Pseudoceltites, Kashmirites, Eukashmirites, and Dieneroceras suggest an Olenekian age for these Tulong brachiopods (Chen, 1983). Sun et al. (1981) reported Lissorhynchia lielonggouensis (Sun), Akrekia chaqupuensis Sun, and Paranorellina dulongdeqingensis from the Spathian (late Olenekian) strata of the Lhasa area in southern Tibet. Wang et al. (1989) reported a brachiopod fauna from a horizon 20–30 cm below the P/T boundary of the Selong section and correlated this fauna with the post-extinction Changhsingian fauna of South China (see Chen et al., 2005-this volume). In addition to this surviving fauna, when one of us (ZQC, unpublished data) further re-examined the Selong brachiopod collection donated by Wang et al. (1989), he obtained several tiny specimens morphologically homologous to Piarorhynchella and Neoretzia, which are found in the Olenekian. 2.9. South China The stratigraphic record in South China shows the most diverse Paleozoic-type Lower Triassic brachiopod faunas in the world (Table 1). However, most of these Lower Triassic brachiopod faunas are survivors of the end-Permian event (Chen et al., 2005-this volume). Additional to these surviving brachiopods (Fig. 2), Feng and Jiang (1978) reported TPiarorhynchellar gujiaoensis from Lower Triassic strata of the Anshuan area of Guizhou Province, southwestern China. Xu and Grant (1994, p. 16) reported some specimens of Spiriferina sp. and Retzioidea from the Fourth Member (Dienerian) of the Daye Formation of the Xiushan section, Daye City, in the Hubei Province. Shen and He (1994) also described Laevorhynchia tenuis from the Guiding section in Guizhou (Table 1). Although its internal features remain unknown, this small and smooth rhynchonellide may represent a Mesozoictype element, because it is morphologically different from the brachiopods in the Changhsingian faunas (Shen and He, 1994). Indeed, this species from Guizhou was subsequently interpreted to be late Griesbachian in age (Rong and Shen, 2002), although we need to know more about its taxonomy and biostratigraphy.

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Chen et al. (2002) described a new form, Meishanorhynchia meishanensis, from the upper Griesbachian of the Meishan section in southeastern China. This taxon, which certainly occurs in the Mesozoic, may give some clues regarding the onset of brachiopod recovery in the Early Triassic as discussed below. In addition to these published data, we can also report that at least one distinct genus is recognizable from the Lower Triassic of the Three Gorges section, in the western Hubei Province. Zhan (1989) noted the appearance of the Mesozoic-type brachiopods in the Lower Triassic Feixianguan Formation at the Shangsi section of northern Sichuan Province, southwestern China. 2.10. New Zealand MacFarlan (1992) described a diverse fauna of Triassic–Jurassic age from New Zealand and New Caledonia. The following three species were assigned to the Lower Triassic: Aparimarhynchia dunrobinrensis MacFarlan from the Lower Triassic of the Aparima Valley; ?Maorirhynchia sp. A from the Lower Triassic of the Hokonui Hills; and Aorhynchia sp. from the Snowdon Formation in the Countess Range, and near Maraoa, northern Southland. Among these localities, age determination of the Snowland Formation to the Early Triassic seems to be corroborated by radiolarian correlations of Aitchison et al. (1988). If this age is correct, these three forms would represent genuine Mesozoic-type brachiopods that originated in the Early Triassic of that region. However, the brachiopod Aorhynchia in the Snowdon Formation is also associated with the conulariid Permophorus obovata and an ammonoid fauna (Waterhouse, 1978, 2002). These ammonoids have been determined to be of mid-Anisian age (Waterhouse, 2002), and regional stratigraphic correlations of the Snowdon Formation suggest a Middle Triassic age as well (Waterhouse, personal communication, 2004). Except for Aparimarhynchia, some well-preserved ammonoids of unequivocal Middle Triassic age are also present in the sequences exposed in the Aparaima Valley (Waterhouse, personal communication, 2004). Similarly, Hokonui Hills, where Maorirhynchia was described, includes abundant parapopanoceratids that are characteristic of the mid-Anisian stage of the Middle Triassic series (Waterhouse, 2002). The Early Triassic age assignment of the above localities

was further questioned when Campbell and GrantMackie (2000) reviewed Triassic stratigraphic correlations of New Zealand. In fact, these authors rejected the occurrence of the Lower Triassic in the above three areas suggested by MacFarlan (1992). Thus, it appears that the so-called Lower Triassic sequences inferred by MacFarlan (1992) in the above three fossil localities in New Zealand are most likely of Middle Triassic in age. Therefore, in our study these three New Zealand brachiopod genera are not considered among the Early Triassic taxa, although they were included as Early Triassic genera in the revised Treatise (see Savage et al., 2002, p. 1270; Owen and Mancengido, 2002, p. 1285; Mancengido et al., 2002, p. 1312). 2.11. Western USA Dagys (1974) listed five species in four genera from the Lower Triassic of North America. However, his generic identifications relied heavily on the literature and have been mostly rejected by Hoover (1979), who also described six species in five genera (including four new genera), Portneufia episulcata, Rhaetina incurvirostra, Vex semisimplex, Obnixia thaynesiana, Periallus woodsidensis, and Periallus. aff. woodsidensis from the western USA. Perry and Chatterton (1979) also described two species, TRhynchonellar triassica and Protogusarella smithi from Idaho. Dagys (1974) placed Pugnoides triassica Girty, originally described from the Olenekian of the western USA, in Piarorhynchella. However, this assignment was rejected by Perry and Chatterton (1979). This suggests that the Primorye materials may not be conspecific with the American species.

3. Taxonomic emendation and Lazarus taxa Among the brachiopods recorded in Lower Triassic strata, Orbicoelia (= Crurithyris of Dagys, 1974) is a representative of the relict Permian elements in the post-extinction Changhsingian faunas (e.g., Sheng et al., 1984). Its appearance in Induan strata of the Caucasus region is questionable because the so-called TInduanr strata of that region (e.g., Sarycheva and Sokolskaya, 1965) have been considered to be of Changhsingian age on the basis

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of revised ammonoid and conodont data (e.g., Tozer, 1969; Sheng et al., 1984; Kotlyar et al., 1983, 1999). In fact, although Dagys (1974, 1993) argued that Orbicoelia specimens came from the Induan of the Caucasus region, he did not document the detailed stratigraphy. Alternatively, a few similar brachiopods have been reported from the late Changhsingian successions of the same region by Kotlyar et al. (1983, 1999, 2004), but these authors never mentioned brachiopods in the Lower Triassic strata. Thus, despite the fact that the genus Orbicoelia occurs as a survivor taxon in the Induan stage of the Salt Range, in South China and Oman (Fig. 2; Table 1), it is excluded from the Induan in the Caucasus region because of the objections raised above. The following three taxa, Rhaetina, Neowerella, and Fletcherithyris, have been either misidentified or improperly placed among Paleozoic-type genera that reappeared in the Triassic. The genus Rhaetina was first proposed for a species from the Kossener schichten (Rhaetian or uppermost stage of the Late Triassic) in the Eastern Alps, but was later extended to Middle Permian specimens from Sicily, Italy (Gemmellaro, 1899). Subsequently, the Permian taxon originally assigned to TR.r lepton was illustrated and reassigned to the Permian genus Jusina, which is in a different family (Stehli, 1962; Cooper and Grant, 1976). Consequently, Rhaetina is confined to Triassic and younger strata. According to Hoover (1979), Rhaetina first occurred in the Olenekian and persisted until the Late Triassic. This genus, therefore, is a Mesozoic-type brachiopod that originated in the post-extinction oceans. Similarly, the Early Triassic genus Neowellerella, a junior synonym of Lissorhynchia Yang and Xu (1966), has been commonly reported in Upper Permian and the P/T boundary beds in South China (Liao, 1979, 1980, 1984; Zhan, 1989). However, most of the taxa reported at these levels have been reassigned to Prelissorhynchia (Xu and Grant, 1994). Furthermore, a recent study has now clarified that the Permian homologues and some earliest Triassic forms of the genus from South China are generically distinct from Lissorhynchia (Chen and Shi, 1999). Fletcherithyris was originally proposed to include an Early Permian brachiopod genus from New South Wales, eastern Australia (Stehli, 1961; Campbell,

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1965). Later, Dagys (1974) identified Fletcherithyris margaritovi as a cosmopolitan element (Table 2), and recognized this genus in collections from the Lower Triassic of Idaho, USA, and the Olenekian (Smithian and Spathian) of other localities in North America, as well as in the Primorye region of Russia, the Mangyslak Region of West Kazakhstan, and in Japan. However, Hoover (1979, p. 6) questioned the validity of Dagys’ (1974) identifications of the American specimens, which he removed from Fletcherithyris and reassigned to Rhaetina, or his new genus Periallus (Hoover, 1979). Moreover, F. margaritovi described by Dagys (1965) from the Primorye region of Russia is also very different from F. amygdala, the type species of the genus from eastern Australia (Stehli, 1961; Campbell, 1965). F. margaritovi has a conspicuous crural basis that separates the inner and outer hinge plates, and the outer hinge plates extending to the shell floor to form the V-shaped structure without the support of the median septum. By contrast, F. amygdala possesses a high median septum supporting a well-developed septalium in the dorsal valve. Hence, the Russian taxon F. margaritovi may represent a separate genus, and thus the generic assignment is questioned here. Apart from the inarticulate Lingula and the questionable Lazarus genus Fletcherithyris, no other Paleozoic-type genera occur in the Lower Triassic or younger strata. This phenomenon contrasts sharply with the notably abundant Lazarus genera in the post-extinction bivalve faunas (Sepkoski, 1984; Jablonski, 1986; Hallam, 1991), gastropods (Pan and Erwin, 1994; Erwin, 1996, 1998; Erwin and Pan, 1996), and echinoderms (Paul, 1988; Schubert and Bottjer, 1995; Foote, 1996). At the family level, Sepkoski (1982) and Harper et al. (1993) recognized 13 families that originated in the Paleozoic, and range through or occur in the Early Triassic. Some of these families may represent Lazarus taxa. However, classification of the Brachiopoda has been significantly revised recently (e.g., Carter et al., 1994; Brunton et al., 2000; Savage et al., 2002; Alvarez and Rong, 2002), and as a result some of the families listed by Sepkoski (1982) and Harper et al. (1993) have been abandoned and some families bear new stratigraphic ranges. For these reasons, the previous dataset is not followed here (Sepkoski, 1982; Harper et al., 1993). In addition, according to

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Jablonski (1986), Erwin and Droser (1993) and Erwin (1996), the Lazarus taxa bdisappearedQ in the mass extinctions and breappearedQ in the post-extinction recovery stage. The 10 families shown in Table 1 comprise surviving taxa. Of the families comprising the Mesozoic-type brachiopod genera (Table 2), both the Wellerellidae and Dielasmatidae also include post-extinction Changhsingian survivors (Chen et al., in this issue, Appendix). Hence, these 12 families are not Lazarus taxa, although they originated in the Paleozoic and occured until the Early Triassic. Of the remaining 10 families comprising the Mezozoic-type genera (Table 2), the Diplospirellidae, Neoretziidae and Cryptacanthiidae are the Lazarus families (33%), which comprise both Paleozoic and Early Triassic genera, not including the surviving elements, suggesting the considerable Lazarus effect during the event. As such, the Lazarus effect in the Brachiopoda during the end-Permian mass extinction is significant at the family level rather than the genus level.

4. Timing of initial recovery of post-extinction brachiopod faunas Dagys (1993) showed that at least five new genera are known from the Induan, but precise age constraints on his genera are lacking largely due to the paucity of detailed biostratigraphy in these brachiopod-bearing strata. Chen et al. (2002) documented an exceptional example from the Meishan section, South China, the Globe Stratotype Section and Point for the P/T boundary (Yin et al., 2001). In Meishan, the lowest horizon with the true Mesozoic-type brachiopod Meishanorhynchia meishanensis is about 9.5 m above the end-Permian mass extinction horizon (Jin et al., 2000). Although various radiometric ages for the clay beds in Meishan prevent a definite estimate of the absolute age of the lowest Meishanorhynchia horizon (Bowring et al., 1998; Mundil et al., 2001, 2004), the associated ammonoid, bivalve and conodont faunas support a late Griesbachian age for Meishanorhynchia (Chen et al., 2002). Therefore, the initial recovery of brachiopod faunas took place in the late Griesbachian, suggesting that the onset of the post-extinction recovery of brachiopod fauna is very short after the event.

5. Taxonomic selectivity of the recovery brachiopods Beside the relict Permian elements (Table 1), a total of 32 brachiopod species from 20 genera and 12 families have been reported from Lower Triassic rocks worldwide (Table 2). These Mesozoic forms belong respectively to two superfamilies of the Spiriferinida, three superfamilies of the Rhynchonellida, two superfamilies of the Athyridida, and three superfamilies of the Terebratulida. They represent the Early Triassic brachiopod recovery faunas. They are more diverse than the Early Triassic surviving brachiopods (27 species in 15 genera of 10 families, see Table 1). Compared to the pre-extinction Changhsingian and post-extinction surviving brachiopods, the number of brachiopod taxa originating in the Early Triassic is relatively smaller (Fig. 3A). Sample biases may have affected this figure to some extent, due to less attention paid to the recovery faunas after the end-Permian extinction in some regions (Erwin, 1998). Nonetheless, data from several relatively well-studied areas such as South China and the Himalayan region, where the Upper Permian and Lower Triassic successions are well exposed, demonstrate that Lower Triassic brachiopods are much less diverse than the pre-extinction and surviving Changhsingian assemblages (Ager and Sun, 1988; Tong and Yin, 2002; Chen et al., 2002; Rong and Shen, 2002). The Order Rhynchonellida comprises 15 species of the recovery fauna, and is the predominant group (46.8% of the total species), whereas the Terebratulida constitutes 25% with eight species, and Athyridida 18.7% with six species (Fig. 3A). The Spiriferinida is represented by three species (9.5%; see also Fig. 3A). At the superfamily level, Norelloidea is composed of seven species, Rhynchonelloidea and Retzioidea consists of five species each, Dielasmatoidea comprises four species, and Wellerelloidea and Cryptonelloidea include three species, respectively (Fig. 3B). Therefore, the recovery brachiopods are dominated by the Rhynchonellida. Within the Rhynchonellida, both the Norellidae and the Rhynchonellidae, which are dominated by Mesozoic forms, comprise seven and five species, respectively; whereas the remaining three species are allocated to two Paleozoic families, namely the Pontisiidae and the

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20

species

18

Number of species

16 14

genera

12 10

families

8

superfamilies

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A

8

Number of species

7 6 5 4 3 2 1

a

a

de oi at

ul

at

br re

la ie D

Te

sm

ne to yp

Cr

oi

id llo

io tz

de

ea

ea id

de oi

Re

id

A

th

yr

re le el

W

a

ea id llo

id lo el or

N

on ch yn

ea

ea id lo el

in er Rh

B

Pe

nn

os

Sp

pi

iri

rif

fe

rin

oi

oi

de

de

a

a

0

Fig. 3. Faunal compositions of the recovery brachiopods. (A) Number of species, genera, families and superfamilies of the Spiriferinida, Rhynchonellida, Athyridida and Terebratulida. (B) Number of species in the 10 superfamilies represented (data from Table 2).

Wellerellidae. The patterns of taxonomic selectivity indicate that the Rhynchonellida, Terebratulida, Athyridida, and Spiriferinida were capable of recovering from the end-Permian mass extinction (Fig. 3A). The Rhynchonellida rebounded worldwide, whereas the Terebratulida recovered to form a special biogeographic province (discussed below). Furthermore, it is an interesting question as to why the rhynchonellids were able to repopulate successfully in the aftermath of the end-Permian extinction (see also Ager, 1987). Carlson (1991) inferred that rhynchonellids benefited from a lack of calcareous lophophore supports that were responsible for the extinction of the impunctuate spiriferids.

6. Paleobiogeography of the Early Triassic brachiopod faunas Details of the Triassic, in particular the Early Triassic, provincialism of brachiopods remain obscure. Although several earlier studies focused on geographic differences between Triassic (largely Middle Triassic and younger) brachiopods (Ager, 1965, 1988; Dagys, 1974, 1993; Ager and Sun, 1988), they did not produce a detailed biogeographic framework for Lower Triassic brachiopods. Although the numbers of Lower Triassic brachiopod genera are too small for quantitative analysis, their endemism in geographic distribution demonstrates a very high provinciality

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(Fig. 1). Of these, western USA–Spitzbergen, South China and Paleo-Tethyan margin regions (Primorye, Japan, Mangyslak–Caucasus, Alpine Europe, and Himalaya) may represent independent biotic provinces due to their low faunal similarities and remote geographic positions (Table 2; Fig. 1). For example, apart from Obnixia, which is also found in the Spitzbergen region, other genera are endemic to western USA. Two Griesbachian genera and the Dienerian Spiriferina from South China are restricted to that region, although some Primorye genera migrated into South China in late Early Triassic (Olenekian) time. Chen et al. (2005-this volume) also conducted an analysis of brachiopod biogeographic selectivity across the end-Permian mass extinction. The pre-extinction Changhsingian brachiopods are grouped into the Austrazean, Cathaysian, Himalayan, Western Tethyan and Sino-Mongolian Provinces, with high provinciality. This generalized pattern is also consistent with previous biogeographic reconstructions of brachiopods (Nakamura et al., 1985; Shen et al., 2000). Generally, the pre-extinction Changhsingian brachiopod faunas were distributed in the warm Tethyan regions and rarely appeared in cooler Boreal and Gondwanan regions. Immediately after the end-Permian extinction the survival brachiopods populated in the Tropical Tethyan, southern Tibetan and Himalayan Provinces, all of which are confined to the PaleoTethys and its marginal seas. Through the phases represented by the Mixed Faunas (MF) 1–3 (terms of Sheng et al., 1984), the three provinces formed one large province. During the MF 3 phase, the survival brachiopods were dominated by the disaster Lingula fauna. The pattern of the cosmopolitan world closely reflects the brachiopod provinciality at the beginning of the Early Triassic (e.g., Hallam and Wignall, 1997; Erwin, 1998, 2001), coinciding with the maximum anoxic event stage (Wignall and Hallam, 1992; Wignall and Twitchett, 1996, 2002), or the time of maximum flooding event (Hallam and Wignall, 1999). In contrast, the postextinction brachiopods originated in the late Early Triassic appear highly varied, perhaps due to biogeographic adaptations (Fig. 1). These Early Triassic provinces formed in various stages. The earliest Mesozoic-type brachiopods are found in South China, where at least two new late

Griesbachian (early Induan) genera are present. By the late Induan, the Mesozoic-type brachiopods had recovered in the Primorye region of Russia and the Mangyslak–Caucasus region of Central Asia. The Mesozoic-type brachiopods in other regions (e.g. western USA, southwest Japan, Romania–Bulgaria of Alpine Europe, and Spitzbergen) appeared in the late Olenekian. The faunal composition of these five major regions is also very different (Fig. 4). Of these, the faunas in both Primorye–Japan region and western USA are highly diverse at the generic level. However, the American fauna is dominated by the terebratulids, whereas the Primorye–Japan fauna comprises elements representing all four orders of the recovery faunas. The Mangyslak–Caucasus–Alpine Europe fauna is fairly diverse and includes representatives of all four orders similar to the Primorye–Japan fauna, and the Himalayan fauna consists of rhynchonellids and athyridids, whereas the South China faunas are characterized by both rhynchonellids and spiriferinids (Fig. 4). Comparison between the biogeographic distribution of brachiopods in the pre-extinction Changhsingian, survival and recovery intervals, shows that, curiously, the recovery brachiopods are found in Spitzbergen, the Primorye region of Russia, the Mangyslak region of Central Asia, Romania, and western USA, although these regions were actually barren of brachiopods during either the Changhsingian or the post-extinction survival stages (Chen et al., 2005-this volume). Despite a few Changhsingian brachiopods occurring in some hotspots in the Primorye and Caucasus regions (e.g. Kotlyar et al., 1999, 2004), they are morphologically distinguishable from the Mesozoic-type brachiopods. Accordingly, this observation reveals that the recovery brachiopods cannot be readily related to the survival lineages (Fig. 2). In comparison, the Lower Triassic brachiopods tended not to rebound in the regions where the survival brachiopods were relatively abundant and diverse (except for South China), but recovered in the regions where the Changhsingian (latest Permian) faunas had been absent or very rare since the end-Guadalupian (Middle Permian) mass extinction. Some of these regions are remote, cold and high-latitude (i.e., Spitzbergen). As such, the re-population of the recovery brachiopods tended to occur in either the regions barren of the Changhsingian faunas or the regions

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N=10

N=7

N=8

Mangyslak-Caucasus-Alpine Europe

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N=8

283

Himalaya

N=4

South China

Pennospiriferinoidea

Spiriferinoidea

Norelloidea

Wellerelloidea

Athyridoidea

Rhynchonelloidea

Retzioidea

Cryptonelloidea

Dielasmatoidea

Terebratuloidea

Fig. 4. Superfamily compositions of the recovery brachiopod faunas in the Primorye–Japan, Mangyslak–Caucasus–Alpine Europe, Himalaya, western USA–Spitzbergen and South China regions.

where the pre-extinction brachiopods and post-extinction survivors were very rare. This unique biogeographic selectivity is probably partly responsible for the depauperate nature of brachiopods in the Triassic (even Mesozoic) oceans. Therefore, the biogeographic distribution of recovery brachiopods in Early Triassic time is not consistent with the pattern of a cosmopolitan world in the Early Triassic that has been commonly shown by other organisms [e.g., bivalves, ammonoids, see Hallam and Wignall (1997), and references therein]. Satges

Species

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15(1)

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Variation in brachiopod diversity across the endPermian extinction (Fig. 5) generally agrees with the Textinction–survival–recoveryr intervals of the biota proposed by Kauffman and Erwin (1995). Five stages are recognized. Brachiopods in Stage 1 were severely affected by an abrupt crisis (Fig. 6), which has been documented in most of the common benthos (e.g., Jin et al., 2000). Genera

Anisian SmithianSpathian

7. Recovery pattern and process of the post-extinction brachiopods

Fig. 5. Diversity of brachiopod faunas across the end-Permian mass extinction. MF 1 = the mixed fauna 1 (term of Sheng et al., 1984); MF 2 = the mixed fauna 2; MF 3 = the mixed fauna 3. The P/T boundary is placed at the middle part of the MF 2 interval. The horizontal axis represents number of species, genera and families, respectively. Numbers on the left and right sides of each diagram indicate the number of the survival and recovery taxa, respectively. Numbers of the pre-extinction Changhsingian taxa are indicated by arrows. Gray bars indicate recovery taxa; black bars indicate surviving brachiopod taxa; dark gray bars indicate pre-extinction Changhsingian taxa. Bar width indicates number of taxa in various stages. The bracketed numbers in the MF 3 interval indicate the number of taxa persisting into the late Griesbachian and associated with the earliest recovery brachiopods.

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Radiation stage

Recovery/ dispersal stage

Dien.

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Olenekian Anisian Smi. Spa.

284

MF 2 interval

survival stage

MF 1 interval

1 Dien. = Dienerian

Smi. = Smithian

Extinction stage

Spa. = Spathian

Fig. 6. Generalized extinction–survival–recovery pattern for global brachiopod faunas across the end-Permian extinction. (1) extinction stage; (2) survival stage including the mixed fauna (MF) 1–3 intervals; (3) survival–initial recovery stage in the late Griesbachian; (4) recovery– dispersal stage in the Dienerian–Spathian (Early Triassic); (5) radiation stage in the Anisian. Black arrows indicate stratigraphic distribution tendency of brachiopods, but each arrow does not represent numbers of taxa.

Four hundred and twenty species in 143 genera of 50 families of pre-extinction Changhsingian faunas were reduced to 78, 40, and 25, respectively (Fig. 5). Extinction rates of species, genera and families are 81.4%, 72% and 50%, respectively (Chen et al., 2005this volume). Stage 2 brachiopods survived the endPermian extinction. This stage is subdivided into three intervals corresponding to MF 1–3. Most of the survivors were extinct at the end of the MF 2 interval. Only a few articulate forms survived into the MF 3 interval and the late Early Triassic when the disaster taxon Lingula, however, flourished in many regions of the world (Rodland and Bottjer, 2001). Stage 3 is late Griesbachian and is here referred to as the Tsurvival–initial recovery intervalr. During this stage Lingula and several articulated forms survived in the Early Triassic shallow oceans around the world. The earliest Mesozoic-type brachiopods (Meishanorhynchia and Laevorhynchia) were sparsely distributed in South China, suggesting an initially limited recovery of the Mesozoic brachiopod faunas. Stage 4 extended from the Dienerian to the Spathian (late Induan to late Olenekian), more diverse faunas (30 species in 19 genera of 11 families) were present in a wide variety of ecological niches of four different biogeographic provinces. Stage 4 patterns suggest that re-

covery of the Mesozoic brachiopods was global. In addition, despite occurring in late Griesbachian, the earliest generalist fauna of South China neither dispersed over the South China block, nor spread to other regions so that the meaningful re-diversification of the post-extinction brachiopods in that region did not occur until the Anisian (early Middle Triassic). In the Paleo-Tethyan marginal Province, some rebounding genera (even species) such as Abrekia, Piarorhynchella, and Paranorella, for example, that first appeared in the Primorye and Mangyslak regions, northern margins of the Paleo-Tethys, have also been reported in late Olenekian successions of Alpine Europe, southwest Japan, and the Himalayan regions. This fact reveals that the recovery faunas experienced a rapid re-population in both Primorye and Mangyslak regions followed by a rapid dispersal in the late Early Triassic Paleo-Tethys oceans. Similarly, the western American element Obnixia migrated to the Spitzbergen region in late Olenekian. Thus, global migration accelerated the Early Triassic recovery of the brachiopod faunas and potentially also their radiation in the early Middle Triassic (Stage 5; Fig. 6), when at least 50 genera (Hallam and Wignall, 1997; Sepkoski, 2002) appear abruptly in the Anisian (early Middle Triassic) worldwide (Fig. 6).

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Abrupt mass extinction

Diversity

Compared with the extinction–survival–recovery pattern proposed by Kauffman and Erwin (1995), brachiopod response to the end-Permian extinction appears different in several aspects. The brachiopod faunas suffered an abrupt mass extinction at the end of the Permian, then their stepwise extinctions occurred in the post-extinction survival stages. Numerous new species arose during the survival interval, although there were no blooms of opportunistic species. In contrast, blooms of disaster species occurred in the late survival stage (MF 3) and might have persisted into the gap between the end of extinction of most survivors and initial recovery of progenitor taxa. Initial recovery of the progenitor taxa is followed by a global rebound of brachiopod faunas. It is particularly noteworthy that the recovery taxa did not originate from the survival evolutionary lineages, but share some morphological similarities with the Middle Permian forms. For example, the earliest Mesozoic genus Meishanorhynchia is readily differentiated from any survivors from the Late Permian forms, but is moderately related to the Middle Permian genus Paranorella (Chen et al., 2002). In addition, apart from a few hotspots where the pre-extinction Changhsingian brachiopods may be sparsely present (Kotlyar et al., 1999, 2004), the post-extinction Changhsingian brachiopods are generally missing in both the Caucasus and Primorye regions. No Late Permian brachiopod faunas have been reported from Kazakhstan and western USA where the Lower Triassic faunas obviously populated and re-diversified in the aftermath of the end-Permian extinction. The Early Triassic brachiopods also occur in both Japan and the Romania–Bulgaria regions where the late Changhsingian brachiopod faunas remain unknown. Survivors of the end-Permian extinction are highly diverse (Chen et al., 2005-this volume) in both the Himalayan and South China regions, but none of them participated in both post-extinction recovery and post-recovery diversification so that the recovery brachiopods in the Olenekian are dominated by elements, which migrated from other regions. These survivors therefore fall into a pattern that was termed TDead Clade Walkingr (DCW) by Jablonski (2002). Diversity data and recovery processes of the brachiopods during the Early Triassic can be gener-

285

Survival interval Recovery interval

Time Fig. 7. Generalized model showing variations in diversity of brachiopod faunas in the extinction, survival and recovery intervals.

alized as shown in Fig. 7. A dramatic reduction in brachiopod diversity at the end-Permian extinction is followed by several stepwise declines in diversity during the survival interval: the time when the survival brachiopods are dominated by geographically widespread, generalist genera adapted to a wide variety of ecological conditions. This interval is followed by a slow growth of diversity with predominance of progenitor taxa (Fig. 7). During post-extinction recovery, the re-population of brachiopod faunas appears to show marked regional differences, similar to the pattern suggested by Jablonski (1998, 2001) for the molluscan recovery after the end-Cretaceous extinction. In favorable regions the progenitor elements flourished, although brachiopod diversity among rebounding taxa remained notably low. During the late Early Triassic (Olenekian), the recovery of brachiopods is characterized by global spread of the generalist taxa. Of the three models proposed for the recovery of biodiversity after extinctions by Erwin (2000, 2001), our diversity pattern is closest to Model C (Fig. 7), although the diversity of the survival interval is slightly different.

8. Conclusions Taxonomic and stratigraphic review of the Early Triassic brachiopod faunas worldwide demonstrate that initial recovery of brachiopod faunas began in the late Griesbachian, and that the time interval between the end of extinction and the first occurrence of the recovery faunas is much shorter than previously proposed. The dispersal of rebounding

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taxa triggered full recovery of Triassic brachiopods in the late Early Triassic. A lack of Lazarus genera and multiprovincialism characterize the rebounding brachiopod faunas, although Lazarus effect may have affected the Brachiopoda at the familial level to some extent. Rhynchonellid brachiopods were most capable of recovery. The re-population of recovery brachiopods appears to vary geographically, although the recovery faunas show a preference for either regions barren of latest Permian forms or regions where the Changhsingian brachiopods are very rare. This unique biogeographic selectivity is probably partly responsible for the depauperate nature of brachiopod faunas in the Triassic (even Mesozoic) oceans. Five stages of extinction, survival, survival–recovery, recovery–dispersal, and radiation are recognized based on variations in brachiopod faunas across the end-Permian extinction. A dramatic reduction in brachiopod diversity at the end-Permian mass extinction is followed by several stepwise declines in diversity during the survival interval, when the surviving brachiopods were dominated by geographically widespread, generalist genera that adapted to a wide variety of ecological conditions. The survival interval, dominated by eurytopic lineages, was followed by slow re-diversification of new lineages dominated by progenitor taxa.

Acknowledgements The senior author is grateful to Associate Prof. G. R. Shi of Deakin University, Australia and Prof. Sandra Carlson of University of California (Davis) for their multiple discussions on the issues of the end-Permian mass extinction and consequences. Prof. A. Nicora of the Universita degli Studi di Milano, Italy and Dr. Jin-Nan Tong of the China University of Geosciences (Wuhan) are also thanked for guiding us to the Tesero section, Italy and assistance in the field in South China, respectively. Constructive suggestions by three journal reviewers have greatly improved this paper. This study was supported by a grant from the Japanese Society for Promotion of Sciences (JSPS P01103 to ZQC) and a large discovery grant from the Australian Research Council (DP0452296 to ZQC).

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