Forty-four species assignable to 32 genera of late Middle Permian (Capitanian) foraminifers are distinguished from Lepidolina limestone and microbial limestone of the uppermost part of the Taishaku Limestone, northeastern part of Hiroshima Prefecture, Japan. The microbial limestone contains Kahlerina taishakuensis sp. nov. enclosing problematic microfossils within their tests and is taxonomically less diversified than the Lepidolina limestone. Faunal composition of nonfusuline foraminifers is clarified in the Lepidolina Zone of the Akiyoshi Terrane for the first time. In addition to the new species, seven species are systematically described. They are Codonofusiella sp. cf. C. ashioensis Kobayashi, Chusenella otai (Nogami), Parafusulina sp., Lepidolina multiseptata (Deprat), Hemigordiopsis renzi Reichel, Baisalina ovata Han, and Partisania typica Sosnina. Taxonomic comments are given on Kahlerina and Hemigordiopsis.
Introduction
The Taishaku Limestone in the northeastern part of Hiroshima Prefecture, Japan is one of the huge limestone blocks of middle Late Carboniferous (Visean) to late Middle Permian (Capitanian) age embedded in the Permian Akiyoshi Terrane. A fusuline fauna dominated by Lepidolina in its uppermost part is common in these huge blocks such as at Akiyoshi (Toriyama, 1958), Taishaku (Sada and Yokoyama, 1966), Atetsu (Nogami, 1961b), and Omi (Kobayashi, 1988). Paleobiogeographically, it is a striking contrast with the contemporaneous Yabeina globosa fauna found in blocks embedded in the Jurassic terranes of Japan (Kobayashi, 1997b).
Foraminifers herein described were collected from the uppermost part of the Taishaku Limestone. Among them, nine species of the Capitanian foraminifers were previously listed as the opportunity to summarize the morphology and stratigraphic distribution of the tetrataxid foraminifer Abadehella (Kobayashi, 1996, p. 90). Though faunal composition was not mentioned in detail, the Capitanian Taishaku fauna is taxonomically more variable, especially in nonfusuline foraminifers, than other contemporaneous faunas in the Akiyoshi Terrane, and is as variable as those of the Jurassic terranes of Japan. These foraminifers are contained in limestones lithologically and faunistically divisible into two, a Lepidolina limestone with many foraminifers and a microbial limestone with Kahlerina enclosing problematic microfossils in their tests.
This paper expresses the faunal characteristics of these foraminifers, classifying them into 44 species assignable to 32 genera. Among them, eight species, including the new species Kahlerina taishakuensis, are systematically described. Limestone thin sections of 228 slides in this study are stored in the collection of the Museum of Nature and Human Activities, Hyogo, Japan (Fumio Kobayashi Collection, MNHAH).
Material
The Lepidolina-bearing limestone and the surrounding mudstone in the uppermost part of the Taishaku Limestone were designated as the Yasumoto Formation of the Nishi-Uyama Group by Yokoyama (1959). Sada and Yokoyama (1966) described four species of Lepidolina and an unnamed species of Chusenella from the formation. The Yasumoto Formation was later reassigned to the uppermost part of the Uyamano Formation in the central facies of the Taishaku Limestone Group (Hase and Okimura, 1970; Hase et al., 1974).
Figure 1.
Index map showing the sample locality. Topographical map from 1:50,000 map “Shobara” of the Geospatial Information Authority of Japan.
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Seven samples were collected from the gray limestone of the Uyamano Formation exposed along a narrow road, 2.8 km east of the village of Taishaku (Figure 1). They are lithologically divided largely into two types.
One type is a microbial limestone (Sample A) showing undulated lamination and containing locally many individuals of Kahlerina and calcisponges coated with problematic algae (Archaeolithophyllum) (Figure 2.1). Lepidolina multiseptata is also rarely contained in this limestone. The outermost whorl of most Kahlerina is irregularly coiled or not coiled, and the outer test conjoins with that of other individuals in some specimens (Figure 2.2). Some tests enclose microfossils of obscure origin in the chambers of their outer whorls (Figure 3), suggesting a possibility of their being a host for these microorganisms.
The other type is a Lepidolina limestone (Sample B), which is stratigraphically a few meters higher than the microbial limestone (Sample A) and contains many fusulines, some of which have microbial encrustations. In addition, many nonfusuline foraminifers, microproblematica, dendroid corals (Waagenophyllum), phylloid algae, and red algae? (Parachaetetes?) are contained in this limestone. Larger fossil grains are packed with fine bioclasts of unknown affinities and lime mud.
Both samples completely lack limestone fragments reworked from the pre-Capitanian limestone and intercalations of siliciclastic seams. In these respects, they are distinguished from contemporaneous conglomeratic limestones and limestone conglomerates of the Uyamano and overlying Oshigatani Formations (Hase et al., 1974) of the Taishaku Limestone. No limestone like Sample A has been reported from other Capitanian outcrops in the Akiyoshi Terrane.
Faunal composition and comparison
Foraminifers in Sample A are rare except for Kahlerina taishakuensis sp. no v. and poorer in taxonomic diversity than those in Sample B. They amount to 12 species and 12 genera in Sample A, and 43 species and 31 genera in Sample B (Table 1).
The Capitanian fusuline faunas in the Akiyoshi Terrane are marked by dominant Lepidolina shiraiwensis (Ozawa, 1925) that is commonly associated with species of Sumatrina Volz, 1904, Kahlerina Kochansky-Devidé and Ramovš, 1955, Chusenella Hsu, 1942, Codonofusiella Dunbar and Skinner, 1937, Dunbarula Ciry, 1948, Nankinella Lee, 1934 and rarely with Verbeekina Staff, 1909, Pseudodoliolina Yabe and Hanzawa, 1932, Rauserella Dunbar, 1944, and Parafusulina Dunbar and Skinner, 1931 (Toriyama, 1958; Nogami, 1961a, 1961b; Kobayashi, 1988; Ozawa and Kobayashi, 1990). All the fusuline genera distinguished herein were reported by previous authors except for the addition of Sichotenella Tumanskaya, 1953.
The present Taishaku fauna is nearly coeval with or slightly younger than the known faunas from the Akiyoshi Terrane from the viewpoint of the gradualistic phyletic sequence established for Lepidolina (Ozawa, 1975). The taxonomic diversity of the fusulines is not so different and roughly equal between the Akiyoshi Terrane and Jurassic terranes of Japan. However, dominant Lepidolina and the almost complete absence of Yabeina Deprat, 1914 in the Akiyoshi Terrane versus the complete absence of Sumatrina and presence of Gifuelloides Kobayashi, Ross, and Ross, 2010 in the Jurassic terranes are paleobiogeographically significant and have special tectonic implications for the plate movement of Panthalassan seamounts (Kobayashi, 1997a, 1997b; Kobayashi et at., 2006, 2010).
Figure 2.
Photomicrographs of the microbial limestone (1 and 2) and the Lepidolina limestone (3). Many individuals of Kahlerina are arranged roughly parallel to the laminae (horizontal direction) in 2.1. Calcisponges, shown by S, coated with problematic algae (Archaeolithophyllum) are also present. Irregularly coiled Kahlerina conjoin with the outer test of other individuals in 2.2. The Lepidolina limestone (2.3) consists of many fossils [fusulines, microproblematica, and red algae? (Parachaetetes?) shown by P] packed with fine bioclasts of unknown affinities and lime mud.
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Table 1.
List of foraminifers in the two limestone samples studied herein.
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Figure 3.
Problematic microfossils, shown by arrow, preserved within chambers of the test of Kahlerina taishakuensis sp. nov. 1,4. K. taishakuensis contained in the same limestone thin section (D2-013875). 2, 3. enlarged photographs of Figure 7. 11. and Figure 7. 4. All × 30.
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The Capitanian nonfusuline foraminifers, as well as other Permian ones in the Akiyoshi Terrane, are poorly known. Though further investigation is needed, Kobayashi (1988) listed eleven such taxa from the Capitanian of the Omi Limestone: Tuberitina spp., Pachyphloia ovata Lange, 1925; Climacammina sp., Tetrataxis sp., Abadehella coniformis Okimura and Ishii in Okimura et al., 1975; Globivalvulina cyprica Reichel, 1946; Globivalvulina sp. cf. G. vonderschmitti Reichel, 1946; Agathammina sp. A, Agathammina sp. B, Baisalina hunanica Lin, 1978 and Calcitornella sp.
Figure 4.
1–6. Miliolina gen. and sp. indet., 1: D2-025874, 2: D2-025868, 3: D2-025902, 4: D2-013925, 5: D2-025862, 6: D2-025924; all from Sample B, all ×40. 7–9, 12–16. Neodiscus sp. B, 7: D2-025871; 8: D2-025934; 9: D2-013879; 12: D2-025871; 13: D2-013913; 14: D2-025893; 15: D2-025866; 16: D2-025861, all from Sample B, 9:×30; others: ×40. 10, 11. Neodiscus sp. A, 10: D2-013873, ×30; 11: D2-013876, ×40, both from Sample A. 17–21. Baisalina ovata Han, 1982, 17: D2-025868; 18: D2-013898; 19: D2-025868; 20: D2-013879; 21: D2-025874, all from Sample B, all × 15. 22. Baisalina sp., D2-025930, Sample B, ×40. 23. Brunsiella sp., D2-025930, Sample B, ×40. 24. Langella fragilis (Sellier de Civrieux and Dessauvagie, 1965), D2-013889, Sample B, ×50. 25, 26. Langella sp„ 25: D2-025887; ×50, 26: D2-025888; ×40, both from Sample B. 27. Hemigordius discoides Lin, Li, and Sun, 1990, D2-025882, Sample B, ×50. 28, 29. Nodosinelloides sp., 28: D2-025867; ×40, 29: D2-025857; ×50, both from Sample B. 30. Astacolus permicus (Miklukho-Maklay, 1954), D2-025817, Sample A, ×40. 31–33. Hemigordius sp., 31: D2-025919; Sample B, 32: D2-013874; Sample A, 33: D2-025828; Sample A, all ×40. 34, 35. Hemigordiopsis renzi Reichel, 1945, 34: D2-013900; 35: D2-025911, both from Sample B, both ×40. 36, 37. Pachyphloia sp., 36: D2-025889; 37: D2-025902, both from Sample B, both ×40. 38–44. Pachyphloia robusta Miklukho-Maklay, 1954, 38: 258893; 39: D2-025909; 40: D2-025877; 41: D2-025892; 42: D2-025897; 43: D2-025868; 44: D2-013890, all from Sample B, all ×40. 45, 46. Partisania typica Sosnina, 1978, 45: D2-025816; Sample A, 46: D2-013918; Sample B, both ×50.
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More taxa of nonfusuline foraminifers are distinguished in this paper (Table 1). Taxonomic composition, diversity, and abundance of the nonfusuline foraminiferal fauna in the Taishaku Limestone are similar to the contemporaneous Capitanian faunas of the Chichibu Terrane (e.g., Kobayashi, 2006b, 2007a), Maizuru Terrane (Kobayashi, 2007b), and Southern Kitakami-Kurosegawa Terrane (Kobayashi, 2001; Kobayashi et al., 2009). In striking contrast to the fusulines, they are also related to the Lopingian non-fusuline foraminifers in these terranes (e.g., Kobayashi, 1997c, 2002, 2004) and significant differences of the faunal composition are not discriminated between Guadalupian and Lopingian faunas except for newly emerging Colaniella Likharev, 1939 and Paraglobivalvulina Reitlinger, 1965 in the Wuchiapingian (Kobayashi, 1999).
These lines of evidence show that fusulines were severely affected by the abrupt changes of the shallow marine environment that caused the Guadalupian-Lopingian biotic event (Stanley and Yang, 1994; Jin et al., 2003), whereas nonfusuline foraminifers were less remarkably affected.
Systematic Paleontology
Order Foraminiferida Eichwald, 1830
Suborder Fusulinina Wedekind, 1937
Superfamily Fusulinoidea von Möller, 1878
Family Ozawainellidae Thompson and Foster, 1937
Genus
Kahlerina
Kochansky-Devidé and Ramovš, 1955
Type species.—Kahlerina pachytheca Kochansky-Devidé and Ramovš, 1955.
Ussuriella Sosnina in Kiparisova et al., 1956, p. 21 (type, U. ussurica).
Pseudokahlerina Sosnina, 1968, p. 103 (type, P. discoidalis).
Hubeiella Lin in Lin et al., 1977, p. 20 (type, H. simplex).
Discussion.—Kahlerina has been placed variously by authors under the families Verbeekinidae (e.g., Ross, 1967; Loeblich and Tappan, 1988), Ozawainellidae (e.g., Sosnina, 1968), Staffellidae (e.g., Sheng, 1963; Rozovskaya, 1975; Rauzer-Chernousova et al., 1996), or the nonfusulinoidean Endothyridae (Loeblich and Tappan, 1964). It is most probably assigned to the Ozawainellidae based on its lenticular to subnautiloid test with finely porous wall free from recrystallization and no deposits referable to true parachomata.
Pseudokahlerina was separated from Kahlerina by Sosnina (1968) based on some differences such as shape of the external test, test size, wall structure, and development of chomata. Pseudokahlerina cannot be distinguished from Kahlerina by these slight differences and test expansion, and is thought to be congeneric with Kahlerina. Ussuriella Sosnina in Kiparisova et al., 1956 and Hubeiella Lin in Lin et al., 1977 are closely related to Kahlerina in their occurrence from the late Guadalupian and in having similar characters such as size and shape of the test, thickness and structure of wall, and development of chomata. They are also thought to be junior synonyms of Kahlerina.
Kahlerina taishakuensis
sp. nov.
Figures 7.1–7.17
Pseudokahlerina discoidalis Sosnina. Ozawa and Kobayashi, 1990, pl. 11, figs. 5, 6.
Derivation of name.—Geographic name, Taishaku, northeastern part of Hiroshima Prefecture.
Type specimens.—Holotype D2-013873 (axial section, Figure 7.3). Paratypes D2-013875 (tangential section, Figure 7.1); D2-013873 (tangential section, Figure 7.2); D2-013869 (axial section, Figure 7.4); D2-025817 (axial section, Figure 7.5); D2-025816 (axial section, Figure 7.6); D2-013879 (axial section, Figure 7.7); D2-025823 (axial section, Figure 7.8); D2-013874 (sagittal section, Figure 7.9); D2-025806 (axial section, Figure 7.10); D2-013865 (sagittal section, Figure 7.11); D2-025817 (axial section, Figure 7.12); D2-025810 (axial section, Figure 7.13); D2-025805 (axial section, Figure 7.14); D2-037608 (sagittal section, Figure 7.15); D2-013869 (sagittal section, Figure 7.16); D2-025816 (sagittal section, Figure 7.17).
Type locality.—Limestone outcrop of the Uyamano Formation exposed at 2.8 km east of the village of Taishaku (Figure 1).
Diagnosis.—Large-sized Kahlerina, having thick lenticular to subnautiloid test with broadly rounded periphery, variably curved lateral slopes, and shallowly umbilicated poles. Outermost whorl is abruptly and irregularly expanding and uncoiled in specimens.
Description.—Test thick lenticular to subnautiloid with broadly rounded periphery, variably curved lateral slopes, and shallowly umbilicated poles. Axis of coiling straight to gently curved. Mature test with four to five whorls. Length, width, and form ratio highly variable among the specimens. The holotype has 4.5 whorls, and is 0.88 mm in length, 2.35 mm in width, and 0.37 in form ratio.
Proloculus small, spherical, 0.07 to 0.17 mm in outside diameter in the 15 specimens illustrated. Inner two whorls lenticular, involute, and tightly coiled. Succeeding ones rapidly and variably increasing in width. The terminal whorl further increases in width and is about one and a half to two times as high as the preceding one of the normally coiled specimens, and two times or more higher than in the abnormally coiled ones (e.g., Figure 7.4) or uncoiled ones (e.g., Figure 7.1). Length, width, and form ratio in the first to fourth whorls of the holotype 0.11, 0.23, 0.52, and 0.78 mm; 0.19, 0.37, 0.73, and 1.45 mm; and 0.58, 0.62, 0.71, and 0.54. Those of the paratype illustrated in Figure 7. 6 are 0.11, 0.21, 0.44, and 0.67 mm; 0.17, 0.32, 0.60, and 1.15 mm; and 0.65, 0.67, 0.73, and 0.58.
Septa planar and unfluted throughout whorls, and long and curved anteriorly. Septal counts from the first to the fourth whorls 6 or 7, 8 or 9, 8 to 10, and 10 in the five sagittal sections illustrated. Septa and wall very thin in inner two whorls and abruptly thicken in the succeeding ones. Wall exceedingly thin in the first whorl, and as thick as 0.04 to 0.10 mm in the outer whorls, consisting of an outer distinct tectum and an inner, much thicker, translucent porous layer. Chomata absent except for indistinct ones rudimentarily present only in inner whorls.
Material examined.—Seventeen illustrated types and more than 50 other specimens.
Discussion.—Highly variable morphologies in every character of the test are recognized, as represented by the height of chambers and mode of coiling in the outermost whorl. They are gradual in the population. These morphologic differences in thin sections, accordingly, are better considered to represent broad morphologic variation and irregular coiling of this species. Pseudokahlerina discoidalis illustrated by Ozawa and Kobayashi (1990) from Akiyoshi is reassigned to this new species because of its having more diagnostic characters common to those of this new species than of the original Primorye one.
Three specimens named as Hubeiella simplex were illustrated by Lin in Lin et al. (1977) from the top of the Maokouan (Guadalupian) of Hubei. Although morphologic variations of the Hubei specimens are unknown, the present new species differs from the Hubei one in having a larger and shorter subnautiloid test, broadly rounded periphery of not only the external test but also of the outer whorls, and exceedingly thin wall in the inner whorls. Five species of Pseudokahlerina, P. discoidalis, P. compressa, P. latispiralis, P. implexa, and P. crepida were proposed by Sosnina (1968) from the upper Middle Permian Metadoliolina lepida Zone of Primorye. The nine specimens illustrated by her that are classified into those five species are more or less similar to each other and are possibly conspecific, as discussed by Kobayashi et al. (2009). They are distinguished from Kahlerina taishakuensis in having much smaller tests.
Occurrence.—Common in Sample A and rare in Sample B.
Figure 5.
1–3, 7. Rauserella ellipsoidalis Sosnina, 1968, 1: D2-025928, 2: D2-025919, 3: D2-013871, 7: D2-013905, 3: Sample A; others: Sample B, all ×30. 4–6. Dunbarula schubertellaeformis Sheng, 1958, 4: D2-013885, 5: D2-025934, 6: D2-025869, all from Sample B, all ×30. 8–13. Codonofusiella sp. cf. C. ashioensis Kobayashi, 2006a, 8: D2-025874; 9: D2-013879; 10: D2-013920; 11: D2-013867; 12: D2-025889; 13: D2-013923, 11: Sample A; others: Sample B, all ×40. 14. Codonofusiella sp. cf. C. explicata Kawano, 1960, D2-025901, Sample B, ×40. 15, 16. Dunbarula cascadensis (Thompson, Wheeler, and Danner, 1950), 15: D2-02562, 16: D2-025915, both from Sample B, both ×40. 17–19. Spireitlina sp., 17: D2-025877; 18: D2-013920; 19: D2-025876, all from Sample B, all ×50. 20. Spireitlina? sp., D2-025919, Sample B, ×50. 21–23. Postendothyra tenuis Lin, Li, and Sun, 1990, 21: D2-025927; 22: D2-025919; 23: D2-025917, all from Sample B, all ×50. 24–27. Climacammina valvulinoides Lange, 1925, 24: D2-013905, ×15; 25: D2-025843, ×20; 26: D2-013922, ×30; 27: D2-013918, ×20; all from Sample B. 28. Lasiodiscus sp., D2-013911, Sample B, ×50. 29–35. Globivalvulina cyprica Reichel, 1946, 29: D2-025858; 30: D2-025927; 31: D2-025901; 32: D2-025923; 33: D2-025891; 34: D2-025900; 35: D2-025901, all from Sample B, all ×50. 36. Tetrataxis sp. A, D2-025924, Sample B, ×30. 37, 38. Tetrataxis sp. B, 37: D2-025867, Sample B; 38: D2-025842, Sample A; both ×30. 39–41. Dagmarita chanakchiensis Reitlinger, 1965, 39: D2-013898, 40: D2-013905; 41: D2-013881, all from Sample B, all ×50. 42–45. Abadehella coniformis Okimura and Ishii in Okimura et at., 1975; 45: D2-013886; 46: D2-013913; 46: D2-013905, all from Sample B, all ×30. 46. Hemigordius sp. B, D2-037633, Sample B, ×30. 47. Nodosaria sp., D2-013917, Sample B, ×40.
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Family Schubertellidae Skinner, 1931
Subfamily Boultoniinae Skinner, 1931
Genus
Codonofusiella
Dunbar and Skinner, 1937
Codonofusiella
sp. cf.
C. ashioensis
Kobayashi, 2006a
Figures 5.8–5.13
Cf. Codonofusiella ashioensis Kobayashi, 2006a, p. 73, 74, Figures 7.40–7.48.
Discussion.—Codonofusiella ashioensis was proposed by Kobayashi (2006a) from the Wordian Nabeyama Formation based on such morphologic characters as strongly fluted septa and rapidly expanding last whorl, the uncoiled terminal portion of which becomes rectilinear and is larger than the inner coiled whorls. The Taishaku specimens have a larger test than the original material and their uncoiled terminal portion is incompletely preserved. Although their exact identity is uncertain, they are closer to the types of this species than to those of other known species of Codonofusiella.
Figure 6.
1, 2. Parafusulina sp., 1: D2-013892, 2: D2-025921, both from Sample B, both × 10. 3–7, 12. Chusenella otai (Nogami, 1961a), 3: D2-025870; 4: D2-013915; 5: D2-013901; 6: D2-025908; 7: D2-025909; 12: D2-025934, all from Sample B, all ×10. 8, 9. Chusenella sp. A, 8: D2-025915; 9: D2-025882, both from Sample B, both ×10. 10, 11. Chusenella sp. B, 10: D2-025930; 11: D2-025841, both from Sample B, both ×10. 13, 17, 18. Chusenella sp. C, 13: D2-025822; 17: D2-025887; 18: D2-025809, 13, 18: Sample A; 17: Sample B, all ×15. 14–16, 19. Sichotenella ovoidea (Sosnina, 1981), 14: D2-025870; 15: D2-013912; 16: D2-025917; 19: D2-025894, all from Sample B, all ×50. 20, 21. Nankinella sp., 20: D2-025807, ×40; 21: D2-025813, ×20; both from Sample A.
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Family Schwagerinidae Dunbar and Henbest, 1930
Genus
Chusenella
Hsu, 1942
Chusenella otai
(Nogami, 1961a)
Figures 6.3–6.7, 6.12
Schwagerina otai Nogami, 1961a, p. 194, 195, pl. 6, figs. 1–5.
Chusenella sp. A Sada and Yokoyama, 1966, p. 313, 314, pl. 33, fig. 6, pl. 34, fig. 2.
Discussion.—This species, proposed from the “Yabeina” shiraiwensis- “Yabeina” sp. A Subzone of the Atetsu Limestone (Nogami, 1961a), is transferred to Chusenella on account of its tightly coiled inner whorls. The present specimens have a larger proloculus than the original material, but the two groups of specimens maintain a resemblance in other test characters. These specimens are probably identical with the types, though they are incomplete.
The specimen of Chusenella sp. A illustrated from Taishaku (Sada and Yokoyama, 1966; pl. 33, fig. 6) is more similar to this species than to the types of Chusenella conicocylindrica Chen, 1956 and Chusenella globularis (Gubler, 1935) and probably belongs with Chusenella otai. However, Sada and Yokoyama (1966) did not compare their Taishaku specimen with the types of C. otai from Atetsu in spite of the fact that it has some morphologic similarities such as shape and size of the test with nearly straight periphery and bluntly pointed poles, elongate fusiform inner whorls with sharply pointed poles, and septal folding. Some of the characters they have in common are obscure in the present specimens because of abrasion of the tests.
Another specimen of Chusenella sp. A (Sada and Yokoyama, 1966; pl. 34, fig. 2) seems to be considerably different from the larger specimen mentioned above in its more tightly coiled and more numerous inner whorls. On the other hand, its test expansion, sharply pointed poles in the inner and outer whorls, and presence of axial fillings suggest that it is possibly conspecific with Chusenella otai.
In this study, three unnamed species are tentatively distinguished from Chusenella otai. Chusenella sp. A (Figures 6.8, 6.9) has more tightly coiled inner whorls than C. otai and hexagonal middle and outer whorls in axial and tangential sections. Chusenella sp. B (Figures 6.10, 6.11), though the specimens are incomplete, has inflated fusiform outer whorls with sharply pointed poles. Chusenella sp. C (Figures 6.13, 6.17, 6.18) has a a smaller proloculus followed by four to five highly elongate whorls and a corrugated wall. It might only represent the inner whorls of Chusenella otai.
Genus Parafusulina
Dunbar and Skinner, 1931 Parafusulina? sp.
Figures 6.1, 6.2
Schwagerina sp. F Nogami, 1961a, p. 203, pl. 7, figs. 1, 2.
Discussion.—Two axial and other sections illustrated are similar to Chusenella otai, but have a larger proloculus and more loosely coiled inner whorls. Based on these characters, they are doubtfully assigned to Parafusulina. Among previously described schwagerinids, they are the closest to and possibly conspecific with Schwagerina sp. F of Nogami (1961a) from the “Yabeina” shiraiwensis-“Yabeina” sp. A Subzone of the Atetsu Limestone.
Family Neoschwagerinidae Dunbar and Condra, 1927
Subfamily Lepidolininae Miklukho-Maklay, 1958
Genus
Lepidolina
Lee, 1934 emend. Ozawa, 1970
Lepidolina multiseptata
(Deprat, 1912)
Figures 7.18–7.21, 8.1–8.13
Neoschwagerina (Sumatrina) multiseptata Deprat, 1912, p. 53–55, pl. 3, figs. 2–8.
Yabeina shiraiwensis (Ozawa, 1925). Nogami, 1961b, p. 186–190, pl. 6, figs. 1–8.
Yabeina sp. A Nogami, 1961b, p. 191–193, pl. 7, figs. 1–5.
Yabeina sp. B Nogami, 1961b, p. 193, 194, pl. 7, fig. 6.
Yabeina multiseptata multiseptata (Deprat). Sada and Yokoyama, 1966, p. 304–307, pl. 33, figs. 4, 5, 7, 8.
Yabeina multiseptata shiraiwensis (Ozawa, 1925). Sada and Yokoyama, 1966, p. 307–309, pl. 33, figs. 1–3.
Yabeina minuta Thompson and Wheeler, 1942. Sada and Yokoyama, 1966, p. 309–311, pl. 34, figs. 3, 4.
Yabeina elongata (Gubler, 1935). Sada and Yokoyama, 1966, p. 311, 312, pl. 34, figs. 1, 5.
Discussion.—Ozawa (1975) demonstrated an instance of phyletic gradualism in Lepidolina, from L. asiatica (Ishii, 1966) to L. multiseptata through L. shiraiwensis. Based on proloculus size and development of secondary transverse septula and axial septula, Yabeina shiraiwensis described by Nogami (1961b) from the Atetsu Limestone and Yabeina multiseptata shiraiwensis of Sada and Yokoyama (1966) from Taishaku are better thought of as belonging to L. multiseptata. Yabeina sp. A and Yabeina sp. B of Nogami (1961b) and Yabeina elongata of Sada and Yokoyama (1966) are referable to microspheric forms of Lepidolina multiseptata. Yabeina minuta of Sada and Yokoyama (1966) in association with Yabeina elongata is thought to be an incomplete specimen of L. multiseptata. Its smaller proloculus is considered to represent the wide morphologic variation of this species.
Figure 7.
1–17. Kahlerina taishakuensis sp. nov., 1: D2-013875; 2: D2-013873; 3: D2-013873 (holotype); 4: D2-013869; 5: D2-025817; 6: D2-025816; 7: D2-013879; 8: D2-025823; 9: D2-013874; 10: D2-025806; 11: D2-013865; 12: D2-025817; 13: D2-025810; 14: D2-025805; 15: D2-037608; 16: D2-013869; 17: D2-025816, all from Sample A except for 7 from Sample B, all ×20. 18–21. Lepidolina multiseptata (Deprat, 1912), 18, 19: megalospheric forms; 20, 21: microspheric forms, 18: D2-013885; 19: D2-013923; 20: D2-013919; 21: D2-025853, all from Sample B; 18, 19, 20a, 21a: ×10; 20b, 21b: ×20.
![f07_260.eps](ContentImages/Journals/jpal/14/4/1342-8144-14.4.260/graphic/WebImages/f07_260.jpg)
Suborder Miliolina Delage and Hérouard, 1896
Superfamily Cornuspiroidea Schultze, 1854
Family Hemigordiopsidae Nikitina, 1969
Genus
Hemigordiopsis
Reichel, 1945
Type species.—Hemigordiopsis renzi Reichel, 1945.
Gansudiscus Wang and Sun, 1973, p. 157 (type, G. luquensis).
Discussion.—This genus is easily distinguished from other hemigordiopsids in its large globular test with thick wall. Gansudiscus, proposed from the Middle Permian (Neomisellina Zone) of the Chinling Range in Gansu and Shaanxi, was discriminated by Wang and Sun (1973) from Hemigordius Schubert, 1908 by its planispiral rather than discoidal coiling, larger test, and thicker wall. As indicated by Loeblich and Tappan (1988), Gansudiscus is a junior synonym of Hemigordiopsis.
Some workers (e.g., Brönnimann et al. 1978; Zaninetti et al., 1979) considered Hemigordiopsis to be a junior synonym of Hemigordius. Others, such as Sheng and He (1983), treated this genus as a subgenus of Hemigordius. However, Hemigordiopsis is distinguished from Hemigordius by having a much thicker wall, a larger and more globular test, more whorls, and complete planispiral and involute coiling except for the inner few whorls, as represented by the type species. The former is thought to be a descendent of the latter (Gargouri and Vachard, 1988).
There are two reports of forms referable to Hemigordiopsis from Japan: Hemigordius? sp. C (=Hemigordiopsis harimaensis Kobayashi, 2007c) in association with Yabeina globosa from the Itsukaichi-Ome area, central Japan (Kobayashi, 2005); and Hemigordiopsis harimaensis and Hemigordiopsis renzi from Guadalupian limestone breccias of different ages contained in the Cretaceous Ise Formation near Tatsuno, west Japan (Kobayashi, 2007c).
Hemigordiopsis renzi
Reichel, 1945
Figures 4.34, 4.35
Hemigordiopsis renzi Reichel, 1945, figs. 1, 2; Kobayashi, 2007c, pl. 1, figs. 38, 39.
Gansudiscus irregularis Wang and Sun, 1973, p. 157, 158, pl. 4, figs. 13, 16.
Hemigordius renzi (Reichel). Brönnimann, Whittaker, and Zaninetti, 1978, pl. 10, figs. 1–4; pl. 11, figs. 4–7.
Discussion.—Hemigordiopsis renzi, a widespread species in the Middle Permian from the Mediterranean Sea region (Reichel, 1945; Zaninetti et al., 1979; Gargouri and Vachard, 1988) to Primorye (Nikitina, 1969), is characteristic in its almost planispirally coiled globular test with very thick, dark wall. It is also reported from the lower Upper Permian of eastern Myanmar (Brönnimann et al., 1978). At least one of three forms from the Chinling Range, Gansudiscus irregularis (Wang and Sun, 1973), is considered to be conspecific with this species based on its test size and wall thickness.
Some specimens referable to this species are strongly deformed, as is also the case with Shanita amosi Brönnimann, Whittaker, and Zaninetti, 1978, which is morphologically analogous to this species except for having internal pillars throughout its globular test (Zaninetti et al., 1979). Though noticeably deformed and not well oriented, the two specimens illustrated are interpreted to be an oblique section of inner whorls (Figure 4.34) and an off-center oblique transverse section of Hemigordiopsis renzi after comparison with this species from Japan (Kobayashi, 2007c) and with many forms, some of which are more or less deformed and worn, from Myanmar and Thailand (Brönnimann et al., 1978; Zaninetti et al., 1979). The very thick, dark wall of the present specimens is thought to be due to secondary replacement and to have been originally porcelaneous.
Family Baisalinidae Loeblich and Tappan, 1986
Genus
Baisalina
Reitlinger, 1965
Baisalina ovata
Han, 1982
Figures 4.17–4.22
Baisalina ovata Han, 1982, p. 104, pl. 2, figs. 20, 21.
Discussion.—Baisalina ovata described from the Neoschwagerina Zone of the eastern part of Jilin by Han (1982) was separated from Baisalina hunanica Lin, 1978 by its having a larger test and less developed septalike protrusions. The Taishaku specimens are identical with the types of this species in their large test, thick wall and poorly developed septal protrusions.
In Japan, Baisalina hunanica and B. sp. are reported from the Middle Permian limestones of the Omi Limestone (Kobayashi, 1988), Kaize (Kobayashi, 2006b), Osakama (Kobayashi, 2007a), and other locations. The present specimens have a larger test more than twice as large as these previously reported Japanese specimens. As far as the test size and thick wall, Baisalina ovata is similar to Baisalina rikuzensis Kobayashi, Shiino, and Suzuki, 2009 from the Middle Permian of the Kamiyasse Formation in northeast Japan (Kobayashi et al., 2009). However, the latter has much more septalike protrusions (“pseudosepta”) subdividing outer whorls into more than fifteen chamberlets and is easily distinguished.
Figure 8.
1–13. Lepidolina multiseptata (Deprat, 1912), ail megalospheric forms, 1: D2-025930; 2: D2-025878; 3: D2-013909; 4: D2-025919; 5: D2-025916; 6: D2-025924; 7: D2-025854; 8: D2-025933; 9: D2-025844; 10: D2-025837; 11: D2-025845; 12: D2-025858; 13: D2-025876, all from Sample B, all ×10.
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Suborder Lagenina Delage and Hérouard, 1896
Superfamily Robuloidoidea Reiss, 1963
Family Partisaniidae Loeblich and Tappan, 1984
Genus
Partisania
Sosnina, 1978
Partisania typica
Sosnina, 1978
Figures 4.45, 4.46
Partisania typica Sosnina, 1978, p. 41, 42, pl. 1, figs. 1, 2, 5–7; Kobayashi, 1997c, pl. 5, figs. 11–13.
Partisania flangensis Sosnina, 1978, p. 42, pl. 1, fig. 3.
Partisania sp. Kobayashi, 2004, p. 68, 69, Figures 6.52–6.55.
Discussion.—This species is based on six type specimens from the Metadoliolina lepida Zone of the Midian Chandalaz Horizon of southern Primorye (Sosnina, 1978). The first few whorls appear to be lenticular and coiled, the following chambers are biserially, sigmoidally and rectilinearly arranged, and the last chamber strongly overlaps laterally toward the initial chambers in the holotype (a nearly centered longitudinal section). However, the sigmoidal chamber arrangement and degree of overlapping of the outer chambers appear to be considerably variable according to two off-center longitudinal sections illustrated by Sosnina (1978). This variable appearance suggests that Partisania flangensis described from the same horizon might be conspecific with this species.
Kobayashi (1997c) thought that an oval lagenid from the Changhsingian of the Iwai-Kanyo area was identical with Partisania typica based on the last chamber strongly overlapping laterally and reaching back to the initial chambers, the close resemblance of transverse sections with those of the types, and the wall structure, although centered longitudinal sections could not be prepared.
The two specimens illustrated herein from Taishaku are more closely similar to the types of Partisania typica than those from the Iwai-Kanyo area in their chamber arrangements both in the initial and later stages in spite of their not being centered longitudinal sections. Partisania sp. from the Kamiyasse Formation (Kobayashi et al., 2009) is based on a single oblique transverse section. Although specific identification is impossible, it is certainly assigned to Partisania on account of the characteristic chamber arrangement of the later stage and a wall structure referable to that of the lagenids.
Partisania sp. described from a Lopingian limestone block of western Shikoku (Kobayashi, 2004) was distinguished from this species by the difference of the overlapping of the chambers in the later rectilinear stage. On the other hand, other characters well resemble each other, especially the chamber arrangement of the coiled initial stage and the immediately next stage. This unnamed species from western Shikoku is probably identical with Partisania typica, since the degree of overlapping of the outer chambers is variable from specimen to specimen among the Primorye type specimens and also depended upon the orientation of the longitudinal sections.
Acknowledgments
I am indebted to D. Vachard, K. Ueno and J. R. Groves for their review of the manuscript, H. Furutani for his cooperation in the fieldwork of the Taishaku area, and A. Ujimaru for her help drawing figures. This study was financially supported by a Grant-in Aid for Scientific Research (C) from the Japanese Ministry of Education, Science and Culture awarded in 1994 (Project No. 05640521) and a grant from the Japan Society of the Promotion of Science awarded in 2009 (Project No. 19540497).