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Gondwana Research 21 (2012) 611–623

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Gondwana Research

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Constraining paleo-latitude of a biogeographic boundary in mid-Panthalassa: Fusuline province shift on the Late Guadalupian (Permian) migrating seamount

Akihisa Kasuya a, Yukio Isozaki a,⁎, Hisayoshi Igo b a Department of Earth Science and Astronomy, The University of Tokyo, Komaba, Meguro, Tokyo 153-8902, Japan b Institute of Natural History, Takada, Toshima, Tokyo 171-0033, Japan article info abstract

Article history: The Guadalupian paleo-atoll limestone (Iwato Formation) in SW Japan was primarily formed in low-latitude Received 24 June 2010 mid-Panthalassa and was later tectonically accreted to South China (Japan) margin during the Jurassic. The Received in revised form 4 June 2011 present biostratigraphic study clarified that the Iwato Formation consists of 5 biostratigraphical intervals; i.e. Accepted 6 June 2011 four fusuline assemblage zones (Assemblage zones 1 to 4) and a barren interval on the top. Assemblage zones Available online 12 June 2011 1 to 4 correspond to the Neoschwagerina craticulifera Zone, N. margaritae Zone, Yabeina globosa Zone, and Handling Editor: M. Santosh Lepidolina multiseptata Zone of the conventional Tethyan fusuline stratigraphy, respectively. The present study newly clarified the following significant aspects of paleobiogeography of the Permian fusulines as to the Keywords: extinction of large-tested taxa in the latest Guadalupian. 1) The long unknown stratigraphic relationship was Panthalassa documented for the first time between the Yabeina-dominant interval and the overlying Lepidolina-dominant Seamount one within a single limestone unit. 2) The occurrence of Lepidolina cf. kumaensis Kanmera, the unique last Fusuline runner of large-tested fusuine, was detected for the first time in mid-oceanic paleo-atoll limestones. 3) With Provincialism respect to the northbound migration history of the paleo-seamount capped by the Iwato Formation, the Paleo-latitude development of the two coeval fusuline biogeographic territories in the low-latitude Panthalassa, i.e., the Permian Yabeina territory on the south and the Lepidolina territory on the north, was confirmed. 4) The paleo-latitude Extinction of the biogeographic boundary between the Yabeina and Lepidolina territories is constrained around 12° in the southern hemisphere on the basis of the latest geomagnetic data from the same limestone. This new approach utilizing biostratigraphy on ancient migrating seamounts coupled with geomagnetic paleo-latitude data is applicable to other cases in different time-space co-ordinates and of other fossil groups for constraining position of ancient biogeographic boundaries within lost oceanic domains of deep past. © 2011 International Association for Gondwana Research. Published by Elsevier B.V. All rights reserved.

1. Introduction hosted symbiont photosynthetic algae to reach that size (Ross, 1972; Wilde, 2002; Vachard et al., 2004; Yang et al., 2004; Ota and Isozaki, Fusuline (foraminifera) represents one of the major Late Paleozoic 2006). marine fossil groups that has been conventionally utilized in bio- This long-term trend of fusuline gigantism, however, was punctu- stratigraphic zonation and international correlation of the Upper ated abruptly at the end of Middle Permian (Guadalupian), or more Carboniferous and Permian rocks. The overall evolutionary history of precisely in the late Capitanian (Late Guadalupian) around 260 Ma. fusulines recorded a clear trend of size increase and sophistication of The thorough extinction of large-tested fusulines (i.e. Schwagerinidae wall structures throughout the Late Carboniferous and Permian (e.g., and Verbeekinidae) left solely smaller and simpler-formed dwarf taxa Loeblich and Tappan, 1964; Ross, 1967; Ozawa, 1970; Rozovskaya, (Schubertidae, Ozawainellidae, and Staffellidae) in the Lopingian (Late 1975; Kanmera et al., 1976; Sheng, 1990; Rauser-Chernousova et al., Permian) considerably before the final extinction of all fusulines at the 1996). Like modern foraminifera, fusulines belong to protists, i.e. end of the Permian ca. 252 Ma (Wilde, 2002; Yang et al., 2004; Ota and unicellular eukaryotes. Nonetheless some of the Middle Permian taxa Isozaki, 2006). The selective termination of the large-tested fusulines become extremely large up to the centimeter-scale, e.g., Eopolydiex- at the end-Guadalupian is particularly noteworthy because this faunal odina up to 16 cm in length (Vachard and Bouyx, 2002). Judging from re-organization was the biggest bioevent in the nearly 100 million- the similarity to the modern large foraminifers (e.g. Hallock, 1999), year fusuline history besides their final extinction at the end-Permian. the Permian large-tested fusulines are likewise regarded to have Another interesting aspect of the Permian fusulines is the remark- able development of provincialism, as up to six realms were hitherto distinguished; e.g. western Tethys, eastern Tethys, Panthalassa, NE Asia, ⁎ Corresponding author. Tel.: +81 3 5454 6608; fax: +81 3 5465 8244. northern Gondwana, and western America (e.g., Ross, 1967, 1995; Ishii E-mail address: [email protected] (Y. Isozaki). et al., 1985; Ozawa, 1987; Kobayashi, 1997). Nevertheless the above-

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612 A. Kasuya et al. / Gondwana Research 21 (2012) 611–623 mentioned screening of large-tested fusulines occurred at the end- surrounded by the Jurassic sandstone and mudstone (Fig. 2). As the Guadalupian regardless of such clear provincialism, confirming that the Triassic part is highly limited in volume, the limestone is mostly extinction-relevant environmental change at the end-Guadalupian was composed of the Permian strata. The Permian limestone generally global in context. The selective extinction of warm-water adapted strikes in ENE–WSW with almost vertical bedding planes, facing faunas suggests the appearance of a cool climate; however, the direct kill consistently to the north. Later tectonic disturbance often dissected mechanism of marine animals, including fusulines, has not been fully the limestone by N–S running minor faults, but the internal stra- clarified yet. In this regard, the extinction pattern of temperature- tigraphy was kept coherent, thus is laterally traceable from section to sensitive photosymbiotic organisms, such as large-tested Capitanian section within the studied limestone block. fusulines, holds a key significance in the studies of the end-Guadalupian The Permian limestone is composed of the Guadalupian Iwato extinction and relevant global environmental changes. Formation (ca. 100 m thick) and the overlying Lopingian Mitai Among the Capitanian large-tested fusulines, two representative Formation (ca. 30 m thick) (Fig. 2). These formations consist of dark genera, Yabeina and Lepidolina, are of profound interest not only because to light gray, pure bioclastic limestone without coarse-grained both represent the pre-extinction last runners of the above-mentioned terrigenous clastics, and yield typical Tethyan shallow marine faunas gigantic fusulines but also because these two rarely co-occurred even that include various fusulines, smaller foraminifera, calcareous algae, within the same oceanic domains. The significance of this remarkable bivalves, gastropods, brachiopods, bryozoans, ostracods, crinoids, and dichotomy of the Capitanian fusulines was first pointed out by Toriyama rugose corals (Ota and Isozaki, 2006; Isozaki et al., 2007b; Kani et al., (1967) who tried to explain the phenomenon in terms of sedimentary 2008). The base of the Iwato Formation was not exposed in this area, but facies-control of fusulines by proposing two contrasting lithofacies in judging from the geology of neighboring areas, its basement is likely Japan; i.e., the limestone-dominated Kinshozan facies with Yabeina composed of basaltic lava of oceanic island basalt affinity. The Iwato versus the terrigenous clastics-dominated Kuma facies with Lepidolina. Formation is composed mainly of massive, dark gray wackestone with It was still not clear-cut, however, if this dichotomy simply reflected minor amount of packstone and lime mudstone. The dominant sedimentary facies control because Lepidolina in fact occurs also from occurrence of large-tested fusulines (Neoschwagerina, Yabeina, Lepido- non-terrigenous limestones as in South China, Indochina, British lina) and calcareous algae (Mizzia, Permocalculus, Gymnocodium), Columbia (e.g., Ishii et al., 1969; Goto et al., 1986). By analyzing fusuline together with rugose corals (Waagenophyllidae) and also with large- wall structure, Ozawa (1970) clarified two independent lineages for the shelled bivalves (Alatoconchidae) in part, indicates tropical–subtropical Guadalupian verbeekinids; i.e. the Neoschwagerina craticulifera– warm-water environments for the primary depositional site (Isozaki, N. margaritae–Yabeina lineage and the Cancellina–Colania–Lepidolina 2006; Isozaki and Aljinovic, 2009). In accordance, the latest paleomag- lineage, both branched off from late Cisuralian (Early Permian) Misselina netic measurement for the upper part of the Iwato Formation confirmed and evolved into the forms with larger test and complicated internal that the Capitanian limestone was deposited at ca. 12° in the southern wall structures. Later in the 1980s, two coeval but contrasting fusuline hemisphere (Kirschvink and Isozaki, 2007). (biogeographic) territories within the Tethys–Panthalassa realms were Previous studies provided fragmentary fusuline data; i.e. the proposed; i.e., the Neoschwagerina–Yabeina territory and Colania– sporadic occurrence of Middle Permian Neoschwagerina, Yabeina and Lepidolina territory (Ishii et al., 1985; Ishii, 1990; Hada et al., 2001). Lepidolina from the Iwato Formation, and that of Late Permian The strictly separated occurrence of Yabeina and Lepidolina, however, Codonofusiella, Reichelina, Palaeofusulina from the Mitai Formation has been the main obstacle to examine these hypotheses and to identify (Saito et al., 1958; Kambe, 1963; Kanmera and Nakazawa, 1973; the mutual province/territory boundary. Isozaki and Ota, 2001; Murata et al., 2003). These preliminary data We recently found out a stratigraphic relationship between the suggest that the Iwato Formation ranges by and large in the later half of Yabeina Zone and the Lepidolina Zone for the first time within a single the Guadalupian, and the Mitai Formation in the Lopingian (Wuchia- stratigraphic sequence of a paleo-atoll limestone in Japan. This article pingian+Changhsingian), respectively. Ota and Isozaki (2006) iden- reports our new finding that provides the first direct clue to solve the tified the top of the Iwato Formation, i.e. the boundary between the long-lasting conundrum of the Capitanian dichotomy of large-tested Guadalupian Iwato Formation and the Lopingian Mitai Formation; fusulines. In addition, we discuss some geological implications of the however, neither detailed fusuline zonation within the entire Iwato present results with particular emphasis on constraining paleo-latitude Formation nor the precise age of its base has been clarified yet. The of ancient biogeographic boundary within lost oceanic domains, by present study analyzed fusuline stratigraphy of the Guadalupian Iwato utilizing the biostratigraphical data from ancient migrating seamounts Formation at 12 stratigraphic sections (Sections 1–12 from east to and their paleo-geomagnetism data from the same limestone unit. west; Fig. 2) in the Kamura area; all are in the main limestone body except Section 10 in an isolated block. As all the efforts to extract 2. Geological Setting conodonts from this formation to date ended in vain likely owing to the strong facies control, fusulines provide a basis for subdivision of the The Permian and Triassic limestones in the Kamura area in central Iwato Formation. Although the relatively poor exposures with thick Kyushu (Takachiho town, Miyazaki prefecture) form a part of an vegetation and strong deformation/recrystallization hampered the ancient mid-oceanic atoll complex that primarily developed on a mid- high-resolution biostratigraphy, the general fusuline stratigraphy was Panthalassan paleo-seamount (Fig. 1; Sano and Nakashima, 1997; clarified as described below. Isozaki and Ota, 2001; Ota and Isozaki, 2006). After a long journey across the superocean, the limestone was secondarily incorporated as 3. Fusuline stratigraphy allochthonous (exotic) blocks into the Middle–Upper Jurassic accre- tionary complex in SW Japan (Isozaki, 1997b) together with deep-sea Nine stratigraphic sections (Sections 2–5and7–11) out of 12 yielded cherts and other rock types (e.g., Nakagawa et al., 2009). Within the informative fusulines for biostratigraphy of the Iwato Formation. The accreted limestone blocks in the Kamura area, the primary stratigra- stratigraphic columns of the studied sections are illustrated in Figs. 3 phy of mid-oceanic shallow marine carbonates is preserved; the and 4. Abundance of fusulines varies drastically from section to section limestone ranges in age from the Guadalupian to Norian (Late and/or from sample to sample. We conducted microscopic observation Triassic) with several sedimentary breaks in the Triassic part (Saito for over 400 thin sections for fusuline analysis. The preservation of et al., 1958; Kanmera and Nakazawa, 1973; Watanabe et al., 1979; fusuline tests is generally poor due to severe deformation, frequent Koike, 1996; Ota and Isozaki, 2006). calcite veining, and secondary recrystallization. Owing to the limited The Permo-Triassic limestone in the Kamura area occurs as an over number of available axial sections of fusuline individuals, their iden- 2 km long and 100–150 m wide body, forming an allochthonous block tification were mostly difficult on species level; however, identified Author's personal copy

A. Kasuya et al. / Gondwana Research 21 (2012) 611–623 613

A Akiyoshi

Mino-Tanba belt

Chichibu belt

Akasaka Kamura 300 km

Jurassic (latest Triassic to earliest Cretaceous) accretionary complex

accretionary complex paleo-atoll

paleo-seamount

oceanic plate Pangea BC

Fig. 1. Index map of the Kamura area in central Kyushu, Japan (A), a simplified cartoon showing a ridge-arc transect with a mid-oceanic seamount (B), and the paleogeography of the Permian world with the primary location of the Kamura seamount (filled circle) (C). genus names with some species names from the studied sections are Formation because we cannot identify precise horizons of the first and listed in Table 1 and partly illustrated in Fig. 5. A brief description of last occurrences of diagnostic taxa owing to the scarcity of well- identified fusulines is given in the caption to Fig. 5. preserved specimens. Under the circumstances, here we use assemblage The listed fusuline faunas are categorized roughly into the following zones for subdivision of the Iwato Formation. 4 assemblages; i.e. Assemblages 1 to 4. Assemblage 1 is dominated by As to the fusuline-bearing 9 stratigraphic sections, the following sets Neoschwagerina cf. craticulifera (Schwager), a primitive Neoschwagerina of assemblage zones are recognized; Section 2 with Assemblage zones 1 with relatively simple septula. Assemblage 2 is composed mainly of and 3, Section 3 with Assemblage zones 1 and 3, Section 4 with As- Neoschwagerina cf. margaritae Deprat, an advanced Neoschwagerina. semblage zones 1, 2, and 3; Section 5 with Assemblage zones 1, 2, 3 and Assemblage 3 is dominated by Yabeina and advanced Neoschwagerina, 4; Section 7 with Assemblage zone 3, Section 8 with Assemblage zone 4; and Assemblage 4 by Lepidolina and Gifuella, respectively. Defining strict Section 9 with Assemblage zone 3; Section 10 with Assemblage zones 3 biozonation in terms of taxon range zone appears difficult for the Iwato and 4; Section 11 with Assemblage zones 3 and 4. Sections 3–5, 8, 10,

Sec. 1 131°20 131°21 67

Saraito Kamura Formation (Triassic) Mitai Formation (Lopingian)

Iwato Fomation (Guadalupian) 87 Koseri studied section Sec. 3 Sec. 2

83 Shioinouso 60 Sec. 7 78 Sec. 4 73 Sec. 5 Sec. 6

32°45 N Sec. 8 80 Sec. 9

80

Sec. 10 N 0 500 m 74 59 79 72 77 71 79 Sec. 11 Sec. 12

Fig. 2. Geologic sketch map of the Kamura area, showing the distribution of the Permo-Triassic allochthonous paleo-atoll limestone within the Jurassic accretionary complex and the locations of analyzed sections (topographic map: 1/5000 basic forest map prepared by Miyazaki prefecture). The extensive blank areas represent the Middle Jurassic mudstone/sandstone that surrounds exotic limestone block. Author's personal copy

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150 Sec. 9 Kamura Fm (Triassic) Alatoconchidae coquina bed Mitai Fm (Lopingian) ammonoid Iwato Fm (Guadalupian) rugose coral

Sec. 8 Ota & Isozaki (2006) Sec. 2 Isozaki et al. Isozaki et al. (2007a) Sec. 6 (2007b) Sec. 12 100 Sec. 10 Sec. 7 Sec. 5 Murata et al. (2003) Sec. 1 Sec. 11 Isozaki et al. (2007b) Sec.4 Sec. 3 Y+N Y Y G Nm Y G Nm Y+L Y+G+N Y+G Nm N Y G Y L G N Y L L+G N Y+Nm L N+G L Nm Y Y Nm Y G N Nm Nc Nc Nc 50

Nc

fusuline L: Lepidolina G: Gifuella Nc Y: Yabeina N: Neoschwagerina Nm: N. cf. margaritae Nc: N. cf. craticulifera 0 m

Fig. 3. Stratigraphic column of the Permian and Triassic limestones at 12 analyzed sections in the Kamura area. Note Sections 2, 8, and 10 correspond to the previously studied ones by Murata et al. (2003), Ota and Isozaki (2006) and Isozaki et al. (2007a, 2007b). Note the close association of giant bivalve (Alatoconchiadae) and large fusulines (see text in detail).

and 11 provide critical pieces of information for fusuline stratigraphy of 4. Correlation the Iwato Formation. In particular, Section 5 demonstrates the suc- cessive occurrence of 4 distinct assemblage zones, plus the probable It is noteworthy that 4 fusuline assemblage zones and 1 barren barren interval on top, in a single stratigraphic sequence (Fig. 4). The interval are recognized in sequence within a single limestone body in results from the rest sections support the sequence of Section 5 without the Kamura area (Fig. 6) because previous studies randomly any discrepancy. Section 10 solely recorded the co-occurrence of reported sporadic occurrences of various fusulines. According to Yabeina and Lepidolina (Table 1; Fig. 3), suggesting that a tran- the conventional Middle Permian fusuline studies in Japan, the sitional interval develops between Assemblage zones 3 and 4. In above-described 4 fusuline assemblage zones roughly correspond to addition, Section 8 displays a direct contact between the Assemblage the following 4 Tethyan fusuline zones; i.e. N. craticulifera Zone, zone 4 and the overlying barren interval, and also that between the N. margaritae Zone, Yabeina Zone, and Lepidolina Zone, respectively. barren interval and the overlying Wuchiapingian Codonofusiella–Reich- The N. craticulifera Zone and N. margaritae Zone are correlated with elina Zone (Fig. 3; Ota and Isozaki, 2006). the Murgabian to lower Midian in Transcaucasia, and the latter two On the basis of these data, it is confirmed for the first time that the zones with the middle–upper Midian in Transcaucasia (Leven, 1996; Iwato Formation in the Kamura area comprises 4 distinct fusuline Ueno, 1996; Kobayashi et al., 2007). The former two zones are assemblage zones; i.e. Assemblage zones 1–4 in ascending order. When correlated also with the Wordian (Middle Guadalupian) at GSSP the barren interval over the Assemblage zone 4 is added, it makes 5 (Global Stratotype Point and Section) in Texas, while the Yabeina fusuline intervals in total (Fig. 6). It is noteworthy that Assemblage zone Zone and Lepidolina Zone are correlated with the Capitanian (Upper 4 replete with Lepidolina represents the topmost fusuline-bearing Guadalupian) at GSSP in Texas (Wilde et al., 1999). As the Lopingian interval in the Iwato Formation and that the top of this zone marks the fusuline fauna appeared from the base of the overlying Mitai end-Guadalupian extinction horizon of the large-tested fusulines (e.g., Formation, the barren interval is assigned to the upper Capitanian Stanley and Yang, 1994; Yang et al., 2004; Ota and Isozaki, 2006). In (Ota and Isozaki, 2006). Thus the Iwato Formation ranges in the addition, another interesting point to note is the occurrence of Lepidolina Wordian to Capitanian. cf. kumaensis Kanmera from Section 5 because L. kumaensis has never The Iwato Formation is by and large similar to the Akasaka been reported from mid-oceanic paleo-seamount limestone. limestone in central Japan (Fig. 1A) in terms of lithofacies and faunal According to the composite stratigraphic column based on the 9 content; however, a higher-resolution zoning of fusulines was sections, thickness of each assemblage zone is estimated as follows; performed for the same interval in the Akasaka limestone (Zaw Assemblage zone 1: over 20 m; Assemblage zone 2: ca. 25 m; Assem- Win, 1999). The present dataset from the Kamura area is not yet blage zone 3: ca. 20 m; Assemblage zone 4: ca. 10 m; and the barren sufficient enough to correlate in the same resolution. These two interval: 15 m (Fig. 6). Although the base of the Iwato Formation was limestones are separated from each other for ca. 500 km at present, not identified, the total accumulated thickness of the Guadalupian they were likely derived from independent paleo-seamounts; paleo-atoll carbonate buildup is estimated to be more than 100 m. The nonetheless their mutual similarity probably comes from the eastern half of the limestone body is dominated by the lower half of physiographical proximity in origin. Other Permian paleo-atoll the Iwato Formation, whereas the western half by the upper half limestone blocks in the Jurassic accretionary complex in SW Japan (Figs. 2 and 3). also have similar faunal composition to these two; e.g., the Tsukumi Author's personal copy

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D1

75 D2 Mitai Fm D2.5 D3

70 D4 D4.2

barren interval 65 D6 D7 D8

60 D9 Gifuella sp. D10 Gifuella gifuensis? D11 Lepidolina sp., L. cf. kumaensis, Gifuella gifuensis

55 Assemblage zone 4

50 D11.7 Yabeina sp. (advanced form) Assemblage zone 3

45 D13 D14 Neoschwagerina cf. margaritae

Assemblage zone 2 40 D15 Neoschwagerina margaritae D16 D17 Neoschwagerina cf. craticulifera Iwato Formation Iwato 35 Assemblage zone 1 D25

D26 30 D27

25 D28 fusuline

20 D29 Alatoconchidae (bivalve)

light gray dolomite

15 dark gray limestone D31 D31.1

10 D32 D32.5

5

Section 5 D33 0 m

Fig. 4. Stratigraphic column of the Iwato Formation at Section 5. Note that this section has 4 assemblage zones and a barren interval in sequence. limestone in eastern Kyushu, Torigatayama limestone in central nant assemblage and Yabeina-dominated assemblage for the first time Shikoku, Shirasaki limestone in western Kii Peninsula, and Kuzuu within a single limestone mass derived from a paleo-atoll complex in limestone in northern Kanto. These paleo-atoll complexes likely mid-Panthalassa. Sections 2, 5 and 9 display that the Neoschwagerina- developed on neighboring but different seamounts within the same dominated Assemblage zones 1 and/or 2 are directly overlain by the seamount chain or swarm (= hotspot track) developed in low- Assemblage zone 3 replete with Yabeina without any in-between latitude Panthalassa (Fig. 1C). occurrence of Lepidolina (Fig. 6). In particular, Section 5 demonstrates that the Assemblage zone 4 occurs at ca. 7 m above the horizon of 5. Discussion Assemblage zone 3 (with a 2 m-thick gap on the exposure between them) (Fig. 4). 5.1. The Lepidolina Zone above the Yabeina Zone On the other hand, Section 8 proves that the Assemblage zone 4 with Lepidolina is directly overlain by the Lopingian Mitai Formation The most significant result obtained in this study is the documen- via the barren interval without any in-between occurrence of Yabeina tation of the stratigraphic relationship between the Lepidolina-domi- (Ota and Isozaki, 2006). After all, at least within the Kamura area, the Author's personal copy

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Table 1 Occurrence of Middle Permian fusulines from the Iwato Formation in central Kyushu, Japan (data mostly from the present study and partly from Murata et al., 2003; Ota and Isozaki, 2006; Isozaki et al., 2007a, 2007b). Refer to Figs. 3 and 4.

Fusuline name Section number (stratigraphic level in each section in meter)

23 4 5 7 8 9 1011

Neoschwagerina cf. craticulifera 24 0 18.5–20.5 37.5 Neoschwagerina margaritae Deprat 39.5 N. cf. margaritae Deprat 25.5 49–53 22–28 N. larga Morikawa & Suzuki 5–5.5 N. cf. colaniae Ozawa 5–5.5 N. minoensis Deprat 5–5.5 Neoschwagerina sp. 40 41.5–43 43.5 Verbeekina sp. 28 Gifuella gifuensis Honjo 58 6 Gifuella sp. 39.5 59 12.5–17 33.5 6 Yabeina cf. globosa (Yabe) 5–5.5 Y. cf. katoi (Ozawa) 5–5.5 Yabeina sp. 55 53.1 50 10–20 28–38 2–4 Lepidolina cf. shiraiwensis (Ozawa) 1–6 L. cf. kumaensis Kanmera 58 Lepidolina sp. 58 0–65–5.5 13.3 Colania sp. 35–5.5 Parareichelina sp. 5 Kahlerina sp. 6

Fig. 5. Photomicrographs of large-tested fusulines from the Guadalupian Iwato Formation in the Kamura area (A–F: entire views of test; G–K: enlarged views of wall structure). A: Neoschwagerina cf. craticulifera (Schwager) from Sample D17 at Section 5, B, G: N. cf. margaritae Deprat from C5 at Section 4, C, H: Yabeina sp. from E1 at Section 7, D, I: Gifuella sp. from D10 at Section 5, E, J: Lepidolina sp. from D11 at Section 5, F, K: L. cf. kumaensis Kanmera from D11 at Section 5. Scale bar (upper left for A–F, lower right for G–K) is 1 mm for all specimens. Note the clear difference in preserved wall structures that are critical in identification for 4 genera discussed in this article despite of the severe secondary deformation, vein development, and recrystallization of calcite tests. G: N. cf. margaritae, wall rather thick; primary transverse septula short and broad, connecting with top of parachomata. H: Yabeina sp., thickness of wall moderate to thin; primary transverse septula elongate and triangular in shape; secondary transverse septula short and thin, well developed in outer volutions. I: Gifuella sp., wall thin; primary transverse septula slender, connecting with top of parachomata; secondary transverse septula almost lacking but one or two septula sporadically occur in outer volutions. J: Lepidolina sp., wall very thin; primary transverse septula thin and short; secondary transverse septula first appear in inner 3rd to 5th volution. K: Lepidolina cf. kumaensis, fragmented outer elongate volutions with very thin wall and many thin parachomata; primary transverse septula short and thin; secondary transverse septula well developed and pendant shape with parachomata-like deposits. Author's personal copy

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Sec. 9 Sec. 8 Sec. 2 fusuline Sec. 7 Alatoconchidae Codonofusiella- Sec. 5 Reichelina Zone Mitai Fm Sec. 11 100 barren interval

L G Assemblage Zone 4 Sec. 10 L L+G (Lepidolina Z.) L L Y+L transitional interval between Zone 3 and Zone 4 Sec. 3 Y Y Y Capitanian Wuchiaping. Y+G+N Y+N Nm Assemblage Zone 3 G G G G Y (Yabeina Z.) Y Y+G Sec. 4 Y+Nm Y Y Y Y Nm Nm Nm Nm N Nm Iwato Fm Iwato Assemblage Zone 2 Nm N 50 (Neoschwagerina margaritae Z.) N N+G

Nc

Wordian Nc Assemblage Zone 1 Nc Nc (Neoschwagerina craticulifera Z.) Fusulines Nc ? L : Lepidolina sp. G : Gifuella sp. Y : Yabeina sp. N : Neoschwagerina sp. Nm Neoschwagerina cf. margaritae Nc Neoschwagerina cf. craticulifera 0 m

Fig. 6. Fusuline zones of the Iwato Formation in the Kamura area. Five biostratigraphic intervals are recognized; i.e. Assemblage zones 1 to 4 and the barren interval in ascending order. Assemblage zones 1–4 correspond to the Tethyan standard fusuline zones, i.e. Neoschwagerina craticulifera Zone, N. margaritae Zone, Yabeina globosa Zone, and Lepidolina multiseptata Zone, respectively. The former two zones are correlated with the Wordian, whereas the latter two and the barren interval with the Capitanian.

Yabeina-dominated interval occurs always below the Lepidolina- Section 5 (Figs. 3–5) because this is the first record from mid-oceanic dominant part and not vice versa, suggesting the primary superiority paleoatoll-type limestone. Kanmera (1953, 1954) originally described of the Lepidolina-interval in stratigraphy. In addition, Section 10 has a L. kumaensis from the Kuma Formation that consists mainly of coarse- horizon that yields Yabeina cf. globosa together with Colania sp. and grained terrigenous clastics (conglomerate, sandstone, mudstone) Lepidolina sp. (Murata et al., 2003). This short section likely preserves with minor amount of fusuline-bearing impure (terrigenous grain- a transitional interval that spans across the two assemblage zones rich) limestone. As this unit likely represents fore-arc shelf/slope (Fig. 6). sediments covering older accretionary complexes (Isozaki, 1987) As mentioned before, the stratigraphic relationship between the along the South China margin, its sedimentary facies is remarkably Yabeina Zone (often defined as the Yabeina globosa Zone in Japan) and different from the mid-oceanic paleoatoll pure carbonates like the the Lepidolina Zone (the Lepidolina multiseptata Zone) has been an Iwato Formation. issue of long-lasting uncertainty because these two zones hardly co- The age of the Kuma Formation is no doubt sometime in the later half occur from any Permian limestone section in Japan. Also in other of the Permian but was not sufficiently constrained in detail owing to its Tethyan domains, their mutual stratigraphic relationship was not uniqueness in faunal composition and to the absence of stratigraphic clearly documented. Kanmera (1953, 1954) assigned the Lepidolina relationship with other dated units. L. kumaensis,aswellasaccompanying Zone above the Yabeina Zone, whereas Hanzawa and Murata (1963) L. toriyamai Kanmera as a synonym, represents one of the most advanced proposed the opposite relation (Fig. 7). Ever since the compromising forms of verbeekinidae fusulines. This species rarely occurs from the assignment by treating these two zones as being coeval by Yabe ordinary Lepidolina Zone dominated by L. multiseptata (Deprat) or (1966), both the Yabeina Zone and Lepidolina Zone were tentatively L. multiseptata shiraiwensis (Ozawa). Within a possible range from the regarded as facies-dependent variations for the same time interval Capitanian to the Wuchiapingian, nonetheless, the age of the Kuma (Toriyama, 1967), and both were correlated all together to the Formation and that of the L. kumaensis Zone per se remained Capitanian in Texas and the Midian in Transcaucasia (e.g., Ishii, 1990; unconstrained (e.g., Toriyama, 1967; Kanmera et al., 1976)despitethe Leven, 1996). Consequently, the present results proved that the added radiolarian information (Ishiga and Miyamoto, 1986). interpretation by Kanmera (1953, 1954) was correct at least for the The sporadic occurrences of L. kumaensis were reported addition- mid-Panthalassan paleo-atoll carbonates within the Jurassic accre- ally from the sub-Lopingian strata in East Asia; i.e., the South Kiatakami tionary complexes in Japan. The stratigraphical superiority of the belt in NE Japan (Choi, 1970), the Primorye region of Far East Russia Lepidolina Zone over the Yabeina Zone was confirmed to date solely in (e.g., Sosnia, 1960; Kotlyar et al., 2007), and South China (Rui, the Kamura area. Although no positive evidence for reversed super- 1983). The restricted occurrences in East Asia also suggest the facies- position has been hitherto reported, we definitely need to check the controlled endemic nature of the L. kumaensis fauna (Toriyama, 1967). mutual relationship in other sections in Japan and also in the rest of Under such uncertainty in age, the L. kumaensis-bearing interval was the world. In this regard, what is concerned is the extremely rare separated from the L. multiseptata Zone (or L. shiraiwensis Zone) as the occurrence of Lepidolina from the Yabeina-dominant limestone (Zaw L. kumaensis Zone that was tentatively placed stratigraphically above Win and Sakagami, 1996). the L. multiseptata Zone (e.g., Yabe, 1966; Kanmera et al., 1976; Ishii, 1990). Leven (1996) summarized all these data and concluded that the 5.2. Lepidolina kumaensis from mid-oceanic paleo-atoll carbonate L. kumaensis Zone is properly correlated with the Capitanian in Texas or with the Midian in Transcaucasia. Another noteworthy aspect on fusuline biostratigraphy found in The present find of L. cf. kumaensis from the upper part of the this study is the occurrence of Lepidolina cf. kumaensis Kanmera from Iwato Formation accords with Leven's interpretation because the L. cf. Author's personal copy

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Age This study Kanmera (1953, 1954) Hanzawa & Murata (1963) Yabe (1966) Ishii (1990)

Codonofusiella- Wuchiapingian Codonofusiella- Codonofusiella- Codonofusiella- Codonofusiella- Reichelina Reichelina Reichelina Reichelina Reichelina (Lopingian) L. kumaensis barren interval Lepidolina Lepidolina Yabeina Lepidolina (multiseptata multiseptata globosa kumaensis (kumaensis) + kumaensis) Yabeina Lepidolina Capitanian Yabeina Yabeina-Lepidolina globosa globosa multiseptata Lepidolina Lepidolina Yabeina Yabeina multiseptata Guadalupian globosa globosa + kumaensis multiseptata

Wordian Neoschwagerina Neoschwagerina Neoschwagerina Neoschwagerina Neoschwagerina

Fig. 7. Comparison of fusuline zoning scheme for the Middle Permian accreted paleo-atoll carbonates in Japan (compiled from Kanmera, 1953, 1954; Hanzawa and Murata, 1963; Yabe, 1966; Ishii, 1990; this study). For convenience's sake, some taxonomic names are modified from the original descriptions into the currently accepted synonymous names; e.g. L. toriyami Kanmera=L. kumaensis Kanmera, Y. shiraiwensis (Ozawa)=L. multiseptata (Deprat). kumaensis-bearing horizon is ca. 15 m lower than the stratigraphic effective in terminating large-tested fusulines (Isozaki et al., 2007a, contact between the Iwato Formation and the Wuchiapingian Mitai 2007b; Isozaki, 2009a, 2009b). In particular, the drop in sea-surface Formation at Section 5 (Figs. 3 and 4). Thus it appears likely that the temperature coupled with the ocean circulation-driven eutrophica- L. kumaensis fauna never made its way into the Lopingian but became tion may have been critical for the photosymbiotic organisms terminated by the end of the Capitanian (Fig. 7), and that the adapted to the pre-existing oligotrophic conditions, such as large- L. kumaensis Zone belongs to Capitanian. tested fusulines, rugose corals, and aberrant bivalves (Isozaki, 2006; In addition, it is also noteworthy to detect L. cf. kumaensis from mid- Aljinovic et al., 2008; Isozaki and Aljinovic, 2009; Isozaki et al., in oceanic paleo-atoll buildups for the first time in the viewpoint of press). The Phanerozoic sea-level minimum appeared around the end biogeography. The Kuma Formation was located at the fore-arc of the of the Capitanian (Haq and Schutter, 2008) and the coeval migration active margin of Permian South China, whereas the Iwato Formation of mid-latitude fauna into tropical regions (Shen and Shi, 2002) was migrating in somewhere within the low-latitude domain of mid- support the onset of cooling in the Capitanian. Although further proof Panthalassa, ca. 3000 km away from South China. This suggests for the kill mechanism is still needed, it is emphasized that the that L. kumaensis likely had a wider distribution than previously extinction of the Permian large-tested fusulines occurred extensively believed, i.e., not only in the peripheries of East Asia but also in the throughout Tethys–Panthalassa regardless of paleobiogeographic middle of the superocean. In other words, the former interpretation provincialism. of facies-dependent endemism of the L. kumaensis fauna appears Most of the Permian limestones in Japan occur as allochthonous highly unrealistic. Nevertheless we need to explain any possible reason blocks embedded within younger matrices, which were primarily for the rare occurrence of L. kumaensis from mid-oceanic paleo-atoll derived from paleo-atoll complexes in mid-Panthalassa (e.g., Kanmera limestones. and Nishi, 1983; Kanmera et al., 1990; Sano and Kanmera, 1991; Isozaki, 1997a). Among them, the occurrences of Lepidolina are 5.3. Two modes in extinction of large-tested fusulines restricted to the Permian accretionary complexes (of the Akiyoshi, Maizuru, and Kuroseagwa belts in Southwest Japan), whereas those of The disappearance of large-tested fusulines and the following Yabeina to the Jurassic accretionary complexes (of the Mino-Tanba and development of the barren interval essentially suggest the onset of a Chichibu belts) (Ishii et al., 1985; Ishii, 1990). The former group of the severe environmental stress that prohibited the survival of large- limestone with Lepidolina, including the well-known Akiyoshi lime- tested fusulines into the Lopingian (e.g., Jin et al., 1994; Stanley and stone (Fig. 1), was accreted to the Japan margin mostly during the Late Yang, 1994; Wilde, 2002; Yang et al., 2004; Ota and Isozaki, 2006). Permian, whereas the latter group (including the Akasaka limestone) Neither the main cause nor the precise timing of the extinction has yet with Yabeina during the Middle–Late Jurassic, nearly 100 Ma later than been identified; however, the common lines of evidence throughout the former (Maruyama et al., 1997; Isozaki et al., 2010; Fig. 8). These the Tethyan (e.g., Leven, 1996; Yang et al., 2004) and circum-Pacific two contrasting groups of the Guadalupian limestones recorded a regions (e.g., Ross, 1995; Zaw Win, 1999; Wilde, 2002; Ota and similar scenario for the large-tested fusuline extinction but in sharp Isozaki, 2006) suggest the appearance of a global-scale environ- contrast in major genera. mental stress during the Capitanian. Various possible kill mech- The Iwato Formation witnessed this extinction of large-tested anisms of fusulines, in particular, that of the large-tested ones have Guadalupian fusulines (Lepidolina, Yabeina) in the low-latitude mid- been proposed, e.g., drop of seawater temperature, change in salinity, Panthalassa (Kirschvink and Isozaki, 2007). A similar extinction pattern eutrophication etc. (e.g., Brasier, 1995; Wilde, 2002; Yang et al., 2004) of the Yabeina-dominant fauna and the following development of a through comparison with ecology of modern large foraminifers. barren interval were observed also in the Akasaka limestone (Zaw Win, Recent facies analysis, C-isotope chemostratigraphy and magnetos- 1999; Ota and Isozaki, 2006) and in other Permian allochthonous tratigraphy of fusuline-bearing Guadalupian sequences suggest that limestones in the Jurassic accretionary complex in Japan. In contrast, the the cooling in the tropics in Tethys and Panthalassa may have been highest fusuline zone in the Akiyoshi limestone is represented by the

Fig. 8. Paleogeographic maps of the Late Guadalupian (A, B: based on Ziegler et al., 1997 and modified according to Maruyama et al., 1997; Muttoni et al., 2009) and of the Mid- Jurassic (C: modified from Rees et al., 2000). A: the relative positions of the two groups (the Akiyoshi-type and Akasaka-type) of Permian seamounts within the superocean Panthalassa; B: the Middle Permian fusuline territories (compiled from Hada et al., 2001, Ueno, 2003, and the present study). Note the boundary between the Lepidolina territory and the Yabeina territory in mid-Panthalassa is marked by the paleo-seamount with the Iwato Formation (Akasaka-type paleo-atoll limestone) at 12° S and 3000 km to the east of South China. C: the relative position of the paleo-seamount immediately before accretion to the South China margin, together with the migration trajectory of the seamount and that of South China per se during the Late Permian, Triassic, and Early–Middle Jurassic times. Author's personal copy

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Late Guadalupian A (260 Ma) 60N

Panthalassa Bureya 30N N.China

Panthalassa Paleo-Tethys Akiyoshi-group S.China paleo-seamount equator

Pangea Indochina Sibumasu Akasaka-group paleo-seamount 30S

60S

Deep Ocean (< -200 m) Shelf (-200–0 m) Land (> 0 m)

B

60N

? 30N

Lepidolina territory equator Yabeina territory

? Yabeina territory 30S

60S

Lepidolina Yabeina Eopolydiexodina

Mid-Jurassic C 60N

accretion

S.China 30N

Panthalassa

equator Pangea

Iwato Fm Neo-Tethys on a paleo-seamount

30S

60S Author's personal copy

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Lepidolina Zone (Toriyama, 1967; Ueno, 1996). The fusuline stratigraphy latitudes higher than 12° S), whereas its upper part was formed within of the topmost Akiyoshi limestone is still not clear because this part is the Lepidolina territory on the north of the border. composed of limestone conglomerate with abundant L. multiseptata As to the paleo-seamount that accommodated the Iwato Forma- without any trace of the L. kumaensis fauna. A fairly short time interval is tion, the entire travel history from its departure in mid-superocean to estimated between the final deposition of limestone and the subduc- its arrival at subduction zone is summarized as follows; 1) originated tion–accretion at an ancient trench along the South China margin at somewhere in the mid-Panthalassa in the southern hemisphere during the latest Guadalupian and early Lopingian; i.e. the whole where Neoschwagerina (ancestor of Yabeina) dominated, 2) tres- limestone complex was tectonically destructed, partly accreted and/or passed northward through the Yabeina-dominant territory up to 12° S subducted at the active trench, immediately after the deposition (Sano and migrated into the domain replete with Lepidolina during the and Kanmera, 1991). Thus a full record of the extinction history of large- Capitanian (Fig. 8B), 3) experienced the end-Guadalupian extinction tested fusulines for the Lepidolina-dominant group was not preserved in near the equator, 4) kept moving northwestward during the detail in the Akiyoshi limestone and its equivalents contained in the Lopingian, Triassic, and Early Jurassic in the northern hemisphere, Permian accretionary complex in Japan. Nonetheless it is noteworthy and 5) finally docked to the South China margin at mid-latitude in the that the Akiyoshi-type limestone was completely free from Yabeina,and late Middle–early Late Jurassic (Fig. 8C; Maruyama et al., 1997; Ota that the end-Guadalupian extinction of large-tested fusulines in the and Isozaki, 2006). Akiyoshi-type limestone group was marked by the die-off of Lepidolina. The remarkable dichotomy in coeval fusuline assemblages sug- The development of the two contrasting faunas of large-tested fusulines gests the relatively sharp differentiation of the Yabeina and Lepidolina in Panthalassa and their coeval extinction provide an interesting aspect territories within Panthalassa, and also the relatively northerly and/or of the Middle Permian fusuline biogeography. westerly development of the Lepidolina territory, as suggested before (e.g., Ozawa, 1987; Kobayashi, 1999; Hada et al., 2001). Although merely a few examples were listed, Hada et al. (2001) in a preliminary 5.4. Migrating seamounts and fusuline territories in Panthalassa discussion proposed the chronological order of accretion of the Permian seamounts in Canadian Cordillera and in New Zealand. Between the two groups of Permian paleo-seamount limestones in Without practical paleo-latitudinal constraints, however, the actual Japan, there was a large time lag of ca. 100 Ma in accretion timing; i.e. distribution pattern of the two territories in mid-Panthalassa has the Lepidolina-bearing Akiyoshi-type group accreted around 260– remained no more than imaginary. In this regard, the Iwato Formation 255 Ma (Late Permian), whereas the Yabeina-bearing Akasaka-type is a sole recorder of such across-latitude migration of the Permian group around 165–160 Ma (mid-Jurassic) (Isozaki et al., 2010). seamounts with practical paleo-latitude data for the border. A similar During the Middle Permian immediately before their final subduc- across-latitude migration scenario was discussed for the Shan–Thai tion–accretion, the Akiyoshi-type paleo-seamounts were located (Sibumasu) block in Thailand (Hada et al., 2001) and the Ekonay adjacent to the South China margin (Fig. 8A) that stayed in the trop- Terrane in Koryak–Kamtchatka belt in NE Russia (Shi, 2006); ical domain across the equator (e.g., Maruyama et al., 1989; Ziegler however, these cases lack quantitative constraints for paleo-latitude et al., 1997). This is in accordance with the fact that Lepidolina occurs of any province boundary. Furthermore, continental blocks are often commonly from the shelf limestones both in South China (e.g., Sheng, involved in local rifting, collision, rotation, and/or strike-slip dis- 1990; his Yabeina corresponds to Lepidolina) and in Indochina (e.g., placement, thus some complexities are added to paleo-biogeograph- Deprat, 1912; Ishii et al., 1969; Fig. 8B). ical discussion. In addition, often in cases of poorly defined “terranes”, On the other hand, the Akasaka-type paleo-seamounts were basement-covering autochthonous strata and accreted allochthonous located far away from South China in the Middle–Late Permian ones are not clearly separated, therefore, loosely defined identity of when the Akiyoshi-type seamounts were under accretion along the fusuline-bearing limestone may lead erroneous biogeographical South China margin. Given a conservative plate consumption rate at interpretation. In this respect, paleogeographical analysis focusing trench around 3 cm/year, the minimum distance between the South solely on paleo-seamount appears more straightforward, as discussed China margin and the mid-oceanic Akasaka-type paleo-seamounts is above. estimated ca. 3000 km. The paleo-latitude of the depositional site of The present confirmation of the gradual transition from the the Capitanian Iwato Formation was recently determined at 12° S by Yabeina Zone to the Lepidolina Zone in the paleo-atoll limestone the geomagnetic measurement (Kirschvink and Isozaki, 2007). Thus (Fig. 6) demonstrated the first reference point for the actual location the Akasaka-type seamount chain (or swarm) was probably located of the territory border within the superocean Panthalassa (Fig. 8B). In in the low-latitude mid-superocean in the southern hemisphere, order to reconstruct the overall framework of the Middle Permian and was separated for more than 3000 km (ca. 40° in equatorial fusuline paleogeography in Panthalassa, we need more reference longitude) to the east from South China (Fig. 8). Thus the two groups points for confining positions of multiple territory/province bound- of accreted Permian limestones in Japan represent two distinct ocean aries. By performing similar analyses to other circum-Pacific orogens domains within Panthalassa characterized by distinct fusuline fauna that potentially preserve numerous remnants of far-traveled mid- for each (e.g., Ozawa, 1987; Ishii, 1990; Kobayashi, 1999; Hada et al., Panthalassan seamounts, more precise image of the provincialism will 2001). be available in much higher resolution. From the paleo-biogeographical viewpoint, therefore, it is significant As to the global fusuline biogeography, the province/territory that the Iwato Formation solely recorded the co-occurrence of Yabeina subdivision appears not simple. For example, the Sibumasu block of and Lepidolina in an intimate stratigraphic relationship. The Iwato the so-called “Cimmerian” affinity, originated from high-latitude Formation basically belongs to the Yabeina-bearing Akasaka-type Gondwanaland further to the south (Fig. 8B), is characterized by a limestones that accreted to South China (Japan) margin during the unique fauna with Eopolydiexodina (anti-tropical taxon) but without Jurassic; however, its top part exceptionally yields Lepidolina. This Yabeina and Lepidolina (Ueno, 2003). In the middle latitude of the stratigraphic change in faunal composition from the Yabeina-dominant northern hemisphere, on the other hand, the southeastern side of the to Lepidolina-dominant nature within a continuous sequence likely Bureya block in Primorye (Far East Russia) is characterized by another reflects the northbound migration history of the paleo-seamount across distinct fauna dominated by Monodiexodina (anti-tropical taxon) and the fusuline biogeographical territory/province boundary within Lepidolina (e.g., Kotlyar et al., 2007). The middle-latitude domains Panthalassa (Fig. 8B). The lower half of the Iwato Formation was both in the southern and northern hemispheres during the Permian deposited within the Yabeina territory (sensu Ishii et al., 1985)inmid- likely formed anti-tropical distinct fusuline provinces/territories, Panthalassa of the southern hemisphere (in low latitude domains with suggesting the latitude-controlled provincialism of Middle Permian Author's personal copy

A. Kasuya et al. / Gondwana Research 21 (2012) 611–623 621 biota (e.g., Shi, 2006). Nonetheless, along the eastern Panthalassan Guadalupian, and the extinction horizon was recorded ubiquitously margin in the tropical domain, the extension of the Yabeina territory in the Lepidolina Zone. Likewise, the accreted Permian limestones in was recognized in West Texas (Skinner and Wilde, 1953). Further- the western North America yield both Yabeina and Lepidolina more, the Middle Permian fusuline provincialism within Paleo-Tethys (Thompson et al., 1950; Skinner and Wilde, 1966; Goto et al., 1986; appears more complicated than in Panthalassa (e.g., Ueno, 2003). Ross, 1995; Stevens et al., 1997), suggesting the occurrence of the These indicate that the development of fusuline provincialism was similar examples to the Japanese ones mentioned above. In particular, dependent not only on paleo-latitude but also on other factors, such as the possible co-occurrence of Yabeina and Lepidolina in Marble ocean current, nutrient availability etc. Canyon in British Columbia (Skinner and Wilde, 1966; Goto et al., 1986; Kobayashi et al., 2007) appears interesting for further checking. 5.5. Paleolatitude of biogeographic province boundary Thus those records from migrating paleo-seamounts provide in- teresting data that can document the relationship between the fusuline The end-Guadalupian extinction horizon in mid-Panthalassa was territories and extinction. We can test this hypothesis by checking other marked either at the top of the Yabeina Zone in the Yabeina territory examples of Permian paleoatoll limestones in accretionary complexes in (e.g., Akasaka limestone) or at the top of Lepidolina Zone in the the Cordilleran orogen in western North America on the other side of Lepidolina territory (e.g., Akiyoshi limestone). Both Yabeina and Panthalassa, and also in the circum-Pacific subduction-related orogens Lepidolina were derived from the same lineage of late Early Permian in the southern hemisphere because southbound paleo-seamounts verbeekinidae, and radiated/adapted separately into different terri- might record an overturned stratigraphy with the Yabeina Zone above tories during the Guadalupian. When the end-Guadalupian extinction the Lepidolina Zone owing to the opposite direction of seamount occurred globally, involving both the Yabeina and Lepidolina terri- migration (Fig. 9). This approach utilizing biostratigraphy on migrating tories, the paleo-position of each Capitanian seamount decided the paleo-seamounts is applicable to other cases to constrain paleo- pre-extinction last runner of the large-tested fusulines according to its latitudes of biogeographical province boundaries, thus paleo-biogeo- relative position with respect to the territory border at 12° S (Fig. 8B). graphic frameworks, even though ancient seamounts and/or seafloors Even in the same seamount chain or swarm, therefore, the coeval already disappeared from the Earth's surface by the non-stop oceanic extinction was recorded in different faunas. For instance, except for subduction processes. the Iwato Formation, most of the Akasaka-type limestones witnessed the end-Guadalupian fusuline extinction solely in the Yabeina Zone. 6. Conclusions The paleo-seamount with the Iwato Formation likely represented a northernmost segment of the seamount chain that alone crossed the The present fusuline biostartigraphic study distinguished 5 biostrat- territory border before the extinction, whereas the rest still remained igraphic intervals in sequence within a Guadalupian paleo-atoll lime- on the south of the border within the Yabeina territory (Fig. 9). As to stone (Iwato Formation) in the Kamura area, SW Japan, which was the Akiyoshi-type limestones, on the other hand, the entire paleo- primarily formed in the low-latitude mid-Panthalassa and was later seamount chain/swarm was already positioned in the middle of the tectonically accreted to South China (Japan) margin during the Jurassic. Lepidolina territory adjacent to South China margin during the The Iwato Formation is composed of four assemblage zones (1 to 4 in

relative migration trajectory of mid-oceanic seamount with repsect to the terriotry boundary N Lepidolina territory equator territory boundary at 12o S S Yabeina territory

crossing territory boundary (~12o S) staying within staying within Yabeina territory northbound southbound Lepidolina territory position seamount Age Akasaka Kamura no known Akiyoshi seamount seamount example seamount Lop. Wuch. G-LB barren interval extinction Yabeina Lepidolina Capitanian Yabeina Guadalupian Lepidolina Yabeina Lepidolina

Fig. 9. Relative trajectories of paleo-seamount drifting within the mid-superocean (arrows) with respect to the biogeographic boundary between the Lepidolina territory and the Yabeina territory (sensu Hada et al., 2001) (above), and the resultant superposition of the Late Guadalupian fusuline assemblages recorded in paleo-atoll limestone; i.e., 4 possible alternatives with 3 confirmed examples (Akasaka, Kamura, and Akiyoshi seamounts in Japan) (below). Note the Kamura paleo-seamount with the Iwato Formation recorded the trespassing episode across the boundary between southerly Yabeina territory and the northerly Lepidolina territory. In contrast, the Akasaka paleo-seamount recorded its long stable residence within the Yabeina territory, and so did the Akiyoshi paleo-seamount within the Lepidolina territory. An imaginary southbound seamount might record the opposite superposition between the Lepidolina Zone and overlying Yabeina Zone. Author's personal copy

622 A. Kasuya et al. / Gondwana Research 21 (2012) 611–623 ascending order) and a barren interval on the top. Assemblage zones 1 to Ishii, K., 1990. Provinciality of some fusulinacean faunas in Japan. In: Ichikawa, K., Mizutani, S., Hara, I., Hada, S., Yao, A. (Eds.), Pre-Cretaceous Terranes of Japan. 4 correspond to the N. craticulifera Zone, N. margaritae Zone, Y. globosa Nihon-Insatsu, Osaka, pp. 297–305. Zone, and L. multiseptata Zone of the conventional Guadalupian Tethyan Ishii, K., Kato, M., Nakamura, K., 1969. Permian limestones of West Cambodia — fusuline stratigraphy, respectively. The former two are correlated with lithofacies and biofacies. Palaeontological Society of Japan Special Paper 14, 41–55. Ishii, K., Okimura, Y., Ichikawa, K., 1985. Notes on Tethys biogeography with reference the Wordian in Texas, whereas the latter two and the barren interval to to Middle Permian fuslinaceans. In: Nakazawa, K., Dickins, J.M. (Eds.), The Tethys: the Capitanian in Texas. As to the extinction of large-tested fusulines Her Paleogeography and Paleobiogeography from Paleozoic to Mesozoic. Tokai at the end of the Guadalupian, the following significant aspects in University Press, Tokyo, pp. 139–155. biostratigraphy and paleobiogeography of the Permian fusulines were Isozaki, Y., 1987. End-Permian convergent zone along the northern margin of the Kurosegawa landmass and its products in central Shikoku, Southwest Japan. newly clarified. Journal of Geosciences Osaka City University 30, 51–131. Isozaki, Y., 1997a. Contrasting two types of orogen in Permo-Triassic Japan: 1. The stratigraphic relationship of the Lepidolina-dominant interval accretionary versus collisional. The Island Arc 6, 2–24. above the Yabeina-dominant one was confirmed for the first time Isozaki, Y., 1997b. Jurassic accretion tectonics of Japan. The Island Arc 6, 25–51. Isozaki, Y., 2006. Guadalupian (Middle Permian) giant bivalve Alatoconchidae from a after the long-time controversy owing to the absence of their co- mid-Panthalassan paleo-atoll complex in Kyushu, Japan: a unique community occurrence. associated with Tethyan fusulines and corals. Proceedings of Japan Academy 82B, 2. The occurrence of L. cf. kumaensis was detected for the first time in 25–32. fi mid-oceanic paleo-atoll pure limestone. This species was hitherto Isozaki, Y., 2009a. The Illawarra Reversal: a ngerprint of the superplume triggering Pangean breakup and end-Guadalupian (Permian) extinction. Gondwana Research known to occur solely from continental shelf facies sequences 15, 421–432. enriched with terrigenous clastics. The age of the L. cf. kumaensis- Isozaki, Y., 2009b. Integrated plume winter scenario for the double-phased extinction – – – bearing horizon was also constrained to the Capitanian. during the Paleozoic Mesozoic transition: G LB and P TB events from a Panthalassan perspective. 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