New Permian Radiolarians from East Asia and The

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OPEN New Permian radiolarians from east Asia and the quantitative reconstruction of their evolutionary and ecological signifcances Yifan Xiao1*, Noritoshi Suzuki2, Tsuyoshi Ito3 & Weihong He1

The biostratigraphically important Permian radiolarian genera Pseudoalbaillella sensu stricto and Follicucullus (Follicucullidae, Polycystinea) are discriminated by morphological gaps in their wings and segmentation. Previous statistical analyses demonstrated that Longtanella flls morphological gaps between these two genera. Longtanella has long been regarded as a junior synonym of Parafollicucullus, and only a few species have been described. Herein several true Longtanella species are recognized from South China, and eight new species and fve indeterminate species are described and illustrated to prove the validity of the genus Longtanella. In addition, a new genus, Parafollicucullinoides gen. nov., is described. Their palaeogeographic distributions and living environments are explored by applying correspondence analysis (CA), with occurrence datasets of selected fusulinacean genera from the Japanese Islands, China and Sundaland. CA results indicate that Longtanella was present to a limited extent in warmer conditions in the fusulinacean Province B and C during Kungurian–Roadian time, and possibly lived above the thermocline and below the deepest limit of fusulinaceans. The Pseudoalbaillella and the Follicucullus group preferred open ocean conditions, living below the thermocline and distributed not only in the ‘Equatorial Warm Water Province’, but also the northern peri-Gondwana Cool Water Province and the southern North Cool Water Province.

Te radiolarian order Albaillellaria is of particular importance because it includes index fossils for the Upper Devonian to the end-Permian1. Albaillellarians are characterized by poreless coverage with an internal bony triangular frame made of three intersecting rods and they are precisely identifable at the species level thanks to rapid evolution. Teir utility in biostratigraphy means that almost all albaillellarian morphotypes from the late Palaeozoic worldwide have been illustrated and described. Because members of the Permian Albaillellaria are also reported worldwide (South China 2; Japan 3; Tailand4; North American midcontinent 5), biostratigraphic correlations in the uppermost Carboniferous and Permian can be conducted with high resolution across South China, Japan and Tailand, that roughly correspond to the fusulinacean Provinces B (Eastern Tethyan Province) and C (Panthalassan Province)6 established by Kobayashi 7. A well-studied albaillellarian group is the Cisuralian to Guadalupian family Follicucullidae Ormiston and Babcock, 1979 8, which consists of more than 70 species. Te Paleozoic Genera Working Group (PGW Group hereafer) decided to divide three genera, namely Parafollicucullus, Follicucullus and Ishigaconus, into 70 species 9,10. Soon afer this consensus was published, this three-genera scheme was undermined by data from the North America midcontinent5, China11 and Japan3. Xiao et al.12 re-evaluated the generic taxonomy of Follicucul- lidae and discussed their phylogenetic relationships using mathematical methods, such as Hayashi’s quantifcation theory II and parsimony theories. Tis study favoured these recent criticisms over the views of the PGW Group, and concluded on a ten-genera scheme, including the genus Longtanella12. Further, it not only confrmed the validity of Longtanella but also suggested that Longtanella is an important sister group in the evolution between the genera Pseudoalbaillella sensu stricto and Follicucullus. Just before this paper 12 was accepted, the validity of Longtanella was newly morphologically confrmed 11. Herein, we investigate the validity of Longtanella, a key genus for understanding the evolution of late Palaeo- zoic radiolarians, but also for the validity of stratigraphic and palaeogeographic analyses through the Devonian

1State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Wuhan 430074, People’s Republic of China. 2Department of Earth Science, Graduate School of Science, Tohoku University, Sendai 980‑8578, Japan. 3Research Institute of Geology and Geoinformation, Geological Survey of Japan, AIST, Tsukuba 305‑8567, Japan. *email: [email protected]

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Figure 1. (a) Tectonic map of South China (revised afer Zheng et al13) and location of the studied section. (b) Geological map of the studied area (revised afer Ke et al 14). Y.F.X. created this fgure using CorelDRAW X4 (https://www. corel draw. com/ en/ pages/ corel draw- x4/ ).

to Permian. In order to fulfl these objectives, we reviewed all papers with illustrations of representatives of the Follicucullidae and examined their identifcations with care. In exploring the recent suggestion12 that Longta‑ nella could be a phylogenetic bridging genus between Pseudoalbaillella sensu stricto and the Follicucullus group (including all the species of Follicucullus and Cariver), we reexamined all the Longtanella morphotypes from South China. Ten, we explored the palaeobioprovincial scheme of these three radiolarian genera together with fusulinacean genera by using correspondence analysis (CA). Comparisons of palaeogeographic distributions among Longtanella and its ancestor and descendent genera could be helpful in understanding the palaeobio- provincialism in radiolarians. Materials Geological and depositional setting. We examined Cisuralian and Guadalupian bedded cherts of the Bancheng Formation in the Shiti section, Guangxi Province, China, and found 86 well- and moderately preserved specimens of 15 morphotypes whose morphological characters ft the defnition of Longtanella. Tis section is located in the northern part of the Shiti Reservoir, about 4 km southeast of Bancheng Town, which is in the northern part of Qinbei District, Qinzhou City, Guangxi Province (Fig. 1a,b). Te Bancheng Forma- tion is dominated by thin-bedded chert, siliceous mudstone, carbonaceous mudstone and lithic greywacke. Te Shiti section is divided into 15 tectonic slices by a series of thrust faults from east to west (Fig. 1b, supplement 1 Fig. S1). Te total apparent thickness of the section is ca. 230 m, and the thickness of each slice ranges from 1 to 50 m except for poorly exposed parts. Folds with a short half wavelength are visible in some slices, but the continuity of each bed is traceable within a slice. Te Bancheng Formation was named by Zhong Keng and others in 1992 for the lithological body of greyish- yellow and greyish-red thin-bedded chert, radiolarian chert, muddy chert and mudstone near Bancheng Town15. Te formation conformably overlies the Lower Carboniferous Shijia Formation and is unconformably overlain by Upper Permian or Lower Triassic strata 15–17. Te age of the Bancheng Formation ranges from Late Carbon- iferous to early Lopingian, and it yields abundant radiolarians, siliceous sponge spicules, a few conodonts, and foraminifera 17–19. Most of the slices examined in this paper could be correlated to the Kungurian or Roadian on the basis of radiolarian index species (supplement 2), suggesting that the same horizon intervals are repeated along the section by thrust faults. Te depositional setting of these siliceous rocks is not a true ‘pelagic open-ocean’, but possibly a restricted ocean basin20 because true pelagic open-ocean chert as found in Japan have never yielded foraminifera. Tese rocks were deposited in the wedge-shaped Qinfang Basin21 which formed with the collision of the Yangtze block and the Cathaysia Block 20,22 (Fig. 1a). Te Qinfang Basin expanded as a deep-sea basinal setting in the Early Carboniferous and then it started to close as a bathyal sea during the Early to Middle Permian, and prior to the Late Permian the basin fnally disappeared 23,24.

Palaeobioprovincial scheme. Te most appropriate palaeobioprovincial scheme for these radiolarian genera (Longtanella, Pseudoalbaillella sensu stricto and the Follicucullus group) is the fusulinacean scheme. As radiolarians have never occurred with fusulinaceans in the same samples, this tendency is of interest. Palaeobioprovincial analyses combining diferent taxa have rarely been performed for Permian marine organisms. It is generally understood that tropical bioprovinces were widely developed around the equator and expanded to higher latitudes in Greenhouse times such as the Eocene and Cretaceous 25. Te Permian, however, regardless of age and Greenhouse mode, experienced palaeobioprovinces that were partitioned by

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Figure 2. Te global distributions of Longtanella, Pseudoalbaillella and the Follicucullus group. Palaeogeographical map revised afer Lucas35. Orange lines show the fusulinid bioprovinces from Kobayashi7; blue lines show the conodont bioprovinces from Mei & Henderson 27. Y.F.X. created this fgure using CorelDRAW X4.

both latitude and longitude 26; this phenomenon is shown by nektonic animals like conodonts, ammonoids and marine fshes 27–29, and planktonic organisms are predicted to follow the similar patterns. Fossilized planktonic organisms in the Palaeozoic are limited to groups such as radiolarians whose skeleton is siliceous. We know that Cisuralian (Early Permian) radiolarian faunas were signifcantly diferent among the South Urals 30, South China–Japan2, and North American midcontinent 31. Lopingian (Late Permian) radiolarian faunas also showed strong provincialism between the Delaware Basin 32 and the South China–Japan regions 33,34. According to previous correlations, Longtanella ranges from UAZ2 (middle Asselian) to the end of UAZ14 (the Changhsingian) with 6 relatively higher occurrence probability (p ≥ 0.20) in UAZ6 (Kungurian) and UAZ 7 (Roadian) . Te metadatabase from our studies demonstrates that locations yielding Longtanella overlap with those of Pseudoalbaillella and the Follicucullus group in diferent ways in diferent palaeobioprovinces. Te palaeogeographic distribution map shows that Longtanella are restricted to the fusulinacean Provinces B (Eastern Tethyan Province) and C (Panthalassan Province), whereas both Pseudoalbaillella and the Follicucullus group are reported not only in Provinces B and C, but also in Provinces A (Western Tethyan Province) and D (Cratonic North American Realm) (Fig. 2).

Selected fusulinacean genera. Te palaeogeography of fusulinaceans in east to southeast Asia including China and Japan has been well documented36–38. Te Permian fusulinoidean palaeogeographic provinces in China are divided into the Cathaysia Tethys Province (CTP), Angara Tethys Province (ATP) and Gondwana Tethys Province (GTP)39. All these provinces constitute the Eastern Tethyan Province (Province B)6,7. In consideration of this knowledge, locality maps of fusulinaceans (occurrences of fusulinaceans are listed in supplement 3) were prepared for the selected seven genera (Table 1). Tese genera were chosen based on the facts that their taxonomic concepts are very stable, marked by obvious characters, and they are reliable guides to geographic distribution 26,40,41. Te faunal afnity between ‘cold-water’ Monodiexodina and ‘warm-water’ Misellina is ofen reported 37,38,42, although these afnities seem to apply only for the Eastern Tethyan and Panthalassa provinces 43 in the sense of Kobayashi7. Chusenella is helpful to indicate fusulinacean-bearing strata because this genus is commonly found in east and southeast Asia. Additional localities of Permian fusulinacean-bearing strata are provided by the distribution maps of Biwaella and Rauserella. Difering from the major fusulinacean province of Japan, the Panthalassan Province (Province C) and Eastern Tethyan Province (Province B) are marked by the presence of Afghanella, Eopolydiexodina, Gallowaiina, Leella, Pisolina, Polydiexodina, Sumatrina, Wutuella and Zellia44,45. Of these, Afghanella roughly overlaps the range of Longtanella. Results Occurrence of Longtanella and its age. Faunal composition and sample ages in Shiti section. A total of 71 species belonging to 21 genera in the Shiti section have been identifed from 38 of the 46 samples from the Bancheng Formation. Tis assemblage includes many common species for age determination such as Albail‑ lella asymmetrica Ishiga & Imoto46, Albaillella xiaodongensis Wang 47, Parafollicucullus fusiformis Holdsworth & Jones48, Parafollicucullinoides globosus (Ishiga & Imoto)46, Parafollicucullinoides yanaharensis (Nishimura &

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Stratigraphic range (fusulinacean data from Zhang and Wang26)Province(fusulinacean data from Kobayashi7) Cisuralian GuadalupianLopingian Province A Province D Province B Kungurian Capitanian Cha. (Western Province C (Cratonic Genus (Eastern Tethys (Panthalassa North Ass. Sak. Art. Roa. Wor. Wuc. Tethys e. m. l. e. m. l. e. l. Province) Province) American Province) Realm)

Biwaella

Monodiexodina

Misellina

Chusenella

Pisolina

Rauserella

Afghanella

Longtanella

Pseudoalbaillella

Follicucullus

Table 1. Te stratigraphic range and province of the selected fusulinacean and radiolarian genera.

Ishiga)49. Besides, 15 morphotypes of Longtanella were recognized in the samples studied, including eight new species and fve indeterminate species (Fig. 3, supplement 1 Figs S2–S5; supplement 5). Te age of the samples was determined by their coexistent species using the statistical likelihood UAZ ranges 6 proposed recently . Most samples were correlated with UAZ6 or UAZ7 without contradiction in faunal association, except for samples ST9, ST8 and 13ST4-2. UAZ6 and UAZ 7 are also correlated with the Kungurian and Roadian, respectively (supplement 2). All the Longtanella specimens are found in UAZ6 or UAZ7.

Occurrences of related radiolarian genera. Japanese Islands (supplement 1 Fig. S6). Te basement rocks of the Japanese Islands comprise mainly Palaeozoic–Cenozoic accretionary complexes with island arc- related rocks 50,51. Permian radiolarians occur in several Palaeozoic–Mesozoic geological units of the Japanese Islands: North Kitakami Belt, Mino–Tamba–Ashio Belt, Ultra-Tamba Belt, Maizuru Belt, Akiyoshi Belt, Hida- gaien Belt, North and South Chichibu belts and Kurosegawa Belt. Pseudoalbaillella and the Follicucullus group have been reported from everywhere in these tectonic belts except in the North Kitakami Belt, so the occurrences of Longtanella only are mapped herein. Occurrences of Pseudoalbaillella and Longtanella have been reported 52 from the North Kitakami Belt of the Northeast Japan Zone. However, confrmed occurrences of the Follicucullus group have not been recognized yet, probably because of thermal metamorphism by Early Cretaceous granite53. Tere are no reliable Permian fusulinacean data, so we exclude the North Kitakami Belt from further discussion. In the Inner Southwest Japan Zone, occurrences of Longtanella are restricted compared to Pseudoalbaillella and the Follicucullus group. Pseudoalbaillella and Follicucullus occur abundantly in the Mino–Tamba–Ashio Belt, but only a few occurrences of Longtanella have been reported54. Longtanella and Pseudoalbaillella were reported 55 from the Hida-gaien Belt. No obvious Longtanella has been discovered from the Ultra-Tamba, Maizuru and Akiyoshi belts. However, Longtanella was noted 56 in tufaceous mudstone and chert of the Nagato Tectonic Zone. According to these authors, these rocks are possibly derived from the Akiyoshi Belt. Relatively abundant Longtanella occur in the Outer Southwest Japan Zone. Longtanella, Pseudoalbaillella and the Follicucullus group co-occur in some sites of the Northern and Southern Chichibu belts57,58. Te co- occurrence of Longtanella, Pseudoalbaillella and the Follicucullus group has been noted59 in the Kurosegawa Belt, which is located between the Northern and Southern Chichibu belts. Finally, Longtanella was discovered 60 from Permian chert pebbles in the Lower Cretaceous Choshi Group of the Kurosegawa Belt.

Mainland China (exclusive of the Changning‑Menglian Suture Zone) (supplement 1 Fig. S7). Te three radiolarian groups are quite common in the northern and western parts of South China, including Guangxi, Sichuan, Guizhou, Jiangsu, Hubei, and Anhui provinces, mainly from the Gufeng and Bancheng formations. Other occurrences are from accretionary complexes, such as the Xijinulan–Gangqiqu ophiolite complex and East Kunlun tectonic complex. Longtanella is found mostly from the ‘stable continental region’ in South China and less than Pseudoalbaillella and the Follicucullus group in numbers and distributions. Te Follicucullus group is distributed

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Figure 3. (a–h) Scanning electron microscopic image (a) and sketches showing the terminology of Longtanella species. (i) the diagram of the Cartesian coordinates. Te morphological terminology was explained in supplement 4 and the descriptions of the species were in supplement 5. Y.F.X. created this fgure using CorelDRAW X4. Photo credit: Y.F.X.

widely, even in eastern Nei Mongol. Generally, these three groups are not so signifcantly diferent in distribution and are ofen associated with each other.

Sundaland and the Changning‑Menglian Suture Zone (supplement 1 Fig. S8). Te tectonic continuity of the Changning-Menglian Suture Zone is closely related to tectonic zones in Sundaland (the continental core of SE Asia which is comprised of the Sunda shelf and parts of the Asian continental shelf), so the Changning- Menglian suture zone is considered here. Longtanella is not found in the Changning-Menglian Suture Zone where Pseudoalbaillella and the Follicucullus group exist61,62, but present in the Inthanon Suture Zone, the Sra Kaeo Suture and the Bentong-Raub Subzone. As discussed later, radiolarians are more limited in distribution than fusulinaceans. Tey almost all occur in suture zones except for Pseudoalbaillella, which is exclusively found from the Sukhothai Terrane and the Indochina Terrane. As with the distributions in South China, Longtanella, Pseudoalbaillella and the Follicucullus group ofen co-occurred, but there are more records of the latter two than Longtanella.

Occurrences of selected fusulinacean genera. Japanese Islands (supplement 1 Fig. S6). Occurrences of fusulinaceans in the Japanese Islands have been thoroughly plotted 63. Te fusulinacean zones that overlap those of the selected radiolarian genera are the Pseudoschwagerina, Parafusulina and Neoschwagerina zones in the sense of Toriyama 63. Te genus Pisolina was not reported in any of the Japanese Islands. Monodiexodina is exclusively found from the Hida-gaien and Kurosegawa belts. Rauserella is reported from the Akiyoshi, Mino- Tamba-Ashio and Northern and Southern Chichibu belts. Afghanella was only from the Akiyoshi Belt. Biwaella in the Japanese Islands is represented only by Biwaella omiensis Morikawa and Isomi, 1960 64 from the Akiyoshi, the Mino-Tamba-Ashio, the Northern and Southern Chichibu belts. Misellina is the most widespread fusulinacean genus in Japan, having been illustrated from the Hida-gaien, Kurosegawa, Akiyoshi, Mino-Tamba-Ashio and the Northern and Southern Chichibu belts. Like Misellina, Chusenella has been found in the Akiyoshi, Kuro- segawa, Mino-Tamba-Ashio, and Southern Chichibu belts, but not from the Hida-gaien Belt.

Mainland China (exclusive of the Changning‑Menglian Suture Zone) (supplement 1 Fig. S7). Carboniferous to Permian fusulinaceans are widely reported in China 45 from ‘stable continental regions’ such as the Yangtze Cra- ton and Cathaysia Fold Belt and ‘suture or accretionary complex zones’. Pisolina is mainly from ‘stable continen-

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Axis Eigenvalue Percentage of variance Cummulative percentage of variance Axis 1 0.309 33.5 33.5 Axis 2 0.178 19.3 52.8 Axis 3 0.151 16.4 69.2 Axis 4 0.105 11.4 80.6 Axis 5 0.076 8.3 88.9 Axis 6 0.039 4.3 93.1 Axis 7 0.029 3.1 96.2 Axis 8 0.026 2.8 99.0 Axis 9 0.009 1.0 100.0

Table 2. Eigen values and percentage variance of all axes.

tal regions’, namely Anhui, Fujian, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Qinghai, Shaanxi, Sichuan, and Zhejiang provinces. Difering from the Japanese Islands, Afghanella was reported not only from ‘stable continental regions’, namely Guangxi, Guizhou, Hubei, Hunan, Sichuan provinces, but also from ‘suture or accretionary complex zones’ such as the Songpan-Ganzi Zone and the sutures between the North and South Qiangtang Blocks, namely Qinghai, Tibet, and Xinjiang provinces. A very few Permian Biwaella specimens are reported from Xinjiang, Guangxi, Guizhou, and Shaanxi provinces. Rauserella is widely known from several tectonic zones in Japan, but from limited areas in China, namely Anhui, Guizhou, Jiangsu, Jilin, and Sichuan provinces. Monodiexodina is illustrated from Tibet, Hainan, Heilongjiang, Hubei, Jiangsu, Jilin, Nei Mongol, and Sichuan provinces. Almost all fusulinacean localities are in the North China block where no radiolarian-bearing strata are recognized. Misellina is found everywhere in the Permian fusulinacean-bearing strata in China, but is not reported from Hainan, Jiangxi, Jilin, Heilongjiang, and Xinjiang provinces. All the provinces except for Hainan province are located in the North China block or adjacent tectonic zones. Chusenella is more diverse at the species level in China than in the Japanese Islands, and it is known from Tibet, Xinjiang, Anhui, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Qinghai, Shaanxi, and Sichuan provinces. It occurs at high species diversity in the South China block (Guangxi, Guizhou, Hubei, Hunan, Sichuan provinces) and Tibet.

Sundaland and the Changning‑Menglian Suture Zone (supplement 1 Fig. S8). Te occurrence of fusulinaceans in Tailand and Malaysia was summarized by Toriyama 65. Pisolina is known only from the Indochina Terrane, and Biwaella only from one locality of Tailand in the Sundaland and Changning-Menglian suture zone. Rauserella is also a minor component in Tailand and Yunnan. Monodiexodina is found from the Ailaoshan Suture and relevant sutures as well as the Inthanon Suture Zone. Highly diverse Chusenella was reported from Yunnan province, Myanmar and Tailand. Afghanella and Misellina are reported from limited localities in Yunnan. In contrast to rare Afghanella in Yunnan, diverse Afghanella species are reported from Tailand.

Correspondence analysis (CA). Te combinations of radiolarian and fusulinacean occurrences at the scale of tectonic divisions is so complex that we used CA to analyse the data. Te analysis reduced the data to nine axes, of which the frst three explain 69.2% of the variance (33.5%, 19.3% and 16.4% in ascending order; Table 2). On Axis 1, all the fusulinacean genera except Monodiexodina receive positive scores, whereas all the radiolarian genera receive negative scores (Fig. 4, Table 3). Te highest absolute score on Axis 1 is negative, for Monodiexodina (− 1.77), and the highest positive scores are for Pisolina (0.90) and Biwaella (0.75). Values on Axis 2 are high (> 1.00) for Pisolina and low (< − 1.00) for Rauserella (Fig. 4). Te inertia scores (Table 3) show that three genera (44.77% for Monodiexodina, 11.99% for Biwaella and 11.74% for Pisolina) explain 68.5% of the contributions to Axis 1. A total of 81.65% of Axis 2 is explained by Rauserella (55.32%) and Pisolina (26.23%). On Axis 3, high absolute scores are detected for Afghanella (− 0.741) and Monodiexodina (0.717) (Table 3), but the contributions to Axis 3 of these two genera is not high (28.21% for Afghanella and 15.09% for Monodiexo‑ dina), compared with other relatively higher contributions (19.21% for Chusenella, 16.99% for Biwaella, 14.74% for Longtanella) (Table 3). Based on Cos 2 (Table 3), two radiolarian genera (0.262 for Pseudoalbaillella, 0.368 for Follicucullus) and two fusulinacean genera (0.380 for Biwaella and 0.750 for Monodiexodina) are largely explained by Axis 1. Two fusulinacean genera (0.432 for Pisolina and 0.805 for Rauserella) are explained by Axis 2, and one radiolarian genus (0.269 for Longtanella) and two fusulinacean genera (0.386 for Afghanella and 0.532 for Chusenella) are explained by Axis 3. Discussion 6 Longtanella occurrence in references. Xiao et al. noted that Longtanella spp. was found between UAZ 2 11 (middle Asselian) and UAZ 13 (Wuchiapingian). Once Ito documented detailed morphological characters for Longtanella, the source data in Xiao et al.6 must be rechecked for Longtanella. Referring to Supplementary data 6 5 in Xiao et al. , the most reliable range for Longtanella is revised as UAZ 6 (Kungurian) to UAZ8 (Wordian). Tis diference was caused from our earlier incomplete knowledge of the ‘broken wing’. We conclude that the genus Longtanella is a good marker for the Kungurian to the Wordian.

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Figure 4. Correspondence plot of CA applied to the occurrences of selected radiolarian and fusulinid genera. Abbreviations: B., Belt; S., Suture; SZ., Suture Zone; L, indicates the presence of Longtanella. N.S. created this fgure using R version 3.3.2 (https://www.r- proje ct. org/ ) and Y.F.X. revised this fgure using CorelDRAW X4.

CoordinatesContributions (%)Cos2 Taxon information Axis 1Axis2 Axis 3Axis1 Axis 2Axis3 Axis 1Axis2 Axis 3 Longtanella –0.343 0.333 –0.409 5.06 8.27 14.74 0.189 0.178 0.269 Radiolarian Pseudoalbaillella –0.255 –0.121 –0.050 3.74 1.47 0.30 0.262 0.059 0.010 genus Follicucullus group –0.308 –0.113 –0.102 5.46 1.27 1.22 0.368 0.049 0.040 Afghanella 0.491 0.010 –0.741 6.05 0.00 28.21 0.169 0.000 0.386 Biwaella 0.746 0.158 0.621 11.99 0.93 16.99 0.380 0.017 0.263 Chusenella 0.276 –0.205 0.488 3.01 2.89 19.21 0.170 0.094 0.532 Fusulinacean Misellina 0.384 0.195 –0.086 5.29 2.36 0.55 0.228 0.059 0.012 genus Monodiexodina –1.765 0.225 0.717 44.77 1.27 15.09 0.750 0.012 0.124 Pisolina 0.904 1.026 0.355 11.74 26.23 3.70 0.336 0.432 0.052 Rauserella 0.448 –1.490 0.003 2.88 55.32 0.00 0.073 0.805 0.000

Table 3. Coordinates, contributions and Cos2 of the columns (genus) in the frst three dimensions. Black box marks to bigger absolute values.

Interpretation of correspondence analysis. Axis 1. Te positive direction of Axis 1 is largely contributed by Pisolina whereas the negative direction is contributed by Monodiexodina (Fig. 4, Table 3). Te preferred palaeoenvironment of Pisolina is ‘semi-restricted platform facies’ which represents a low hydrodynamic environment66 or ‘inner gentle slope benthic biofacies’67. Te preferred palaeoenvironment of Monodiexodina has been subject to two interpretations. One is an anti-tropical distribution37 although the suggested preferred temperatures are diferent from cold water68, middle latitude between high latitudinal cool/cold-water and tropical warm-water realms37, and temperate cool-water zone between temperate- and warm-water zones43 to warm water69. Te other interpretation is high energy water conditions like clastic lithofacies44 and wave- and storm- reworked, transgressive lag deposits70. Considering the inner gentle slope benthic biofacies of Pisolina, the most likely interpretation of Axis 1 is a range from gentle water conditions (positive) to high-energy water conditions (negative). All three radiolarian genera are plotted in the negative area of Axis 1 (Table 3), probably refecting open ocean conditions. Pseudoalbaillella and the Follicucullus-group are signifcant in Cos2 score (Table 3) although their contributions are not signifcant (Table 3). Tis also implies that open ocean conditions are not the most important factor for the distribution of the Longtanella.

Axis 2. Te preferred palaeoenvironments of Rauserella have not been discussed; it occurs with abundant small foraminifers in intercoralite sediments 71, abundant bryozoans with rare corals 72, abundant non-colonial

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corals73, common calcareous algae74 and variable fragments of fossils75. Following this information, Axis 2 is defned by Pisolina-bearing limestone facies in the positive direction and Rauserella-bearing limestone facies in the negative direction. Te selected three radiolarian genera have never been found from such limestones, and thus coordinates, contributions and Cos2 (Table 3) all are small scores, and not related to Axis 2.

Axes 3 and 4. As discussed above, Monodiexodina may have had an anti-tropical distribution. Preferred palaeoenvironments for Afghanella are poorly described in previous papers, but Afghanella in Zagros, Iran, occurs in a warming event43. Te most likely interpretation of Axis 3 is an anti-tropical distribution in the positive direction and warmer conditions in the negative direction. Te distribution of Longtanella is mainly described by Axis 3 (14.74 in Contribution, 0.269 in Cos2, Table 3) but also Axis 4 (21.79 in Contribution and 0.277 in Cos2), but the interpretation of Axis 4 is difcult. Te coordinate score of Axis 3 for Longtanella is -0.409 (Table 3), suggesting warmer conditions on the line of anti-tropical to warmer conditions axis. Low values for Pseudoalbaillella and Follicucullus show that these two radiolarian genera were not related to this anti-tropical to warmer conditions axis.

Summary of interpretations for selected radiolarian genera. Te distributions of Pseudoalbaillella and Follicucullus correspond to open ocean conditions, but such conditions were not important for Longtanella. Instead, Longtanella preferred warmer condition along the anti-tropical to warmer conditions axis, but this condition did not impact on the distributions of Pseudoalbaillella and the Follicucullus group. Terefore, Longtanella appears to have been well adapted to warmer conditions, difering from the widespread ancestral Pseudoalbail‑ lella as well as its widespread descendant the Follicucullus group. Compared to Pseudoalbaillella, Longtanella shows atrophied pseudothoracic wings and reduced pseudothorax and increased test height, and it evolved into the Follicucullus group by complete reduction of undulated segmentation of pseudoabdomen.

Revisiting the palaeogeographic map. Te fusulinacean palaeogeographic map shows diferent palaeo-provinces among the western Palaeo-Tethys (Province A), eastern Meso-Tethys and Meso-Tethys (Province B), Panthalassa (Province C) and the eastern margin of Panthalassa (Province D)7 (Fig. 1). Our CA analysis shows the combination of Provinces B and C with a common distribution over this area for Pseudoalbaillella and the Follicucullus group. Tis result also supports the earlier recognition6 of the same low-latitudinal standard Permian radiolarian biostratigraphy between Provinces B and C. Te relatively restricted distribution of Longta‑ nella in Provinces B and C is explained by the preferred warmer conditions in our CA. Tis seems to contradict the higher latitudinal distribution in British Columbia and Far East Russia. However, the original depositional position of parts of the British Columbia blocks or terranes were at a northern middle latitude 200 Ma age (earliest Jurassic)76 and the original position in the Permian was probably at a lower latitude. Permian radiolarian and fusulinacean localities in Far East Russia are a northern extension of the tectonic divisions of the Japanese Islands 50 and thus the original depositional positions were also at low latitudes. Te Permian radiolarian faunas are the same between Japan and Far East Russia6. Te fusulinacean faunal similarity among British Columbia, Far East Russia (= Primorye, Sikhote-Alin) and the Japanese Islands is already known 77 and was identifed as the ‘Tethyan-Panthalassa fauna’ or ‘Yabeina territory’ in subsequent studies71,78,79. In consideration of these tectonic and faunal afnities, Longtanella may be present in a limited way in warmer conditions in the fusulinacean Prov- inces B and C. For conodonts, the Permian ‘Equatorial Warm Water Province’ along the zone between northern and southern mid-latitudes27 (Fig. 1) includes all of fusulinacean Provinces A, B and C. Both Pseudoalbaillella and the Follicucullus group are distributed not only in the ‘Equatorial Warm Water Province’ but also in the northern peri-Gondwana Cool Water Province and southern North Cool Water Province. By contrast, Longta‑ nella is distributed in a very limited way, as explained above. Te terrestrial and oceanic realms were mapped in the latest Permian 80. Because the Middle Permian was cooler than the latest Permian 81, the tropical ocean zone was narrower in the Middle Permian 80. Waters above the thermocline are variable whereas water below the thermocline is homogenously cold in low to middle latitudes82,83. Tis oceanic rule requires the warm-water-dependent Longtanella to live above the thermocline, which means that it cannot be associated with fusulinaceans because they lived in deep-water environments. In the maps of Middle Permian palaeoprovinces with brachiopods 84, the distributions of both Pseudoalbaillella and the Follicucullus group cover not only the Cathayasian realm but also the southern part of the Sino-Mongolian realm and part of the Cimmerian realm, just like the conodont palaeoprovinces27. Te distributions of brachiopods and conodonts are largely controlled by latitudinal surface water temperature (SST), but the diferences in distribution of Pseudoalbaillella and the Follicucullus group suggest that their distribution did not depend on SST. Tese two genera are found in the Middle Permian limestone in the Guadalupe and Apache mountains of West Texas. Noble et al. 85 explained that a stimulatory response to increased runof is related to abundance changes in Follicucullus ventricosus. Such runof conditions generate a similar oceanographic situation to an upwelling region in terms of oceanic physiology86 and deep-water radiolarians can then be found in the intermediate zone87. Xiao et al.88 defned water depths scheme for the Permian referred from the modern oceanography. Te thermocline is roughly situated in similar water depth to the deep chlorophyll maximum (DCM)” and we defned the very shallow and shallow zones by the DCM. As Longtanella lived in diferent water conditions from Pseudoalbaillella and the Follicucullus group, the latter would have lived below the thermocline.

Evolutionary perspectives. Xiao et al.6 showed mathematically that Longtanella evolved from Pseudoal‑ baillella, whereas the Follicucullus group was divergent from Longtanella (Fig. 5). Our study herein frst recognized that Longtanella was limited to Provinces B and C in the Kungurian to Wordian, difering from provinces where its ancestor (Pseudoalbaillella) and descendant (Follicucullus) lived. Te short range and limited distri-

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Figure 5. Phylogenetic tree of Follicucullidae obtained by the parsimony analysis (revised afer Xiao et al.11, the character data set is provided in supplement 6). Te species name is written as the original described name. Genus abbreviations: L., Longtanella; Pa., Parafollicucullus; Ha., Haplodiacanthus; Ho., Holdsworthella; Ps., Pseudoalbaillella; F., Follicucullus; Ca., Cariver. Y.F.X. created this fgure using TNT version 1.5 (http://www. lillo. org.ar/ phylo geny/ tnt/ ) and CorelDRAW X4.

bution of Longtanella might be related to some climatic changes. Although our data are insufcient to specify this prediction, the early Kungurian is known for the end of late Artinskian–early Kungurian warming and maximum marine fooding event in east Gondwana 89 and the early Capitanian is the time of diversifcation of Follicucullus species90. We hypothesized that Longtanella evolved near east Gondwana to adapt to warmer waters with less input of fresh water around the early Kungurian and lost the competition for survival against the newly diversifed Follicucullus species around the early-middle Capitanian. Conclusions Eight new species and fve indeterminate species of Longtanella are described, which greatly expands knowledge of this genus. Correspondence analysis was applied to occurrences of Longtanella, its sister taxa (Pseudoalbaillella and the Follicucullus group), and seven fusulinacean genera. Te most likely interpretation of the CA output Axis 1 is gentle (positive) to high-energy water conditions (negative). Axis 2 is defned by Rauserella-bearing (negative) to Pisolina-bearing limestone facies (positive). Axis 3 is warmer conditions (negative) to anti-tropical distribution (positive). Comparisons with fusulinacean palaeoenvironmental interpretations based on CA suggest that there are some diferences in the distributions of Longtanella, Pseudoalbaillella and the Follicucullus group. Pseudoalbail‑ lella and the Follicucullus group favour open ocean conditions, but this condition is not important for Longta‑ nella. Longtanella may be present in a limited way in warmer conditions in the fusulinacean Provinces B and C. Pseudoalbaillella and the Follicucullus group are distributed in not only the ‘Equatorial Warm Water Province’ but also in the northern peri-Gondwana Cool Water Province and southern North Cool Water Province in the conodont scheme. Longtanella lived above the thermocline and below the deepest limitation of fusulinaceans. Pseudoalbaillella and the Follicucullus group lived below the thermocline. Tis leads to the suggestion that Longtanella was well adapted to warmer conditions, difering from the widespread ancestral Pseudoalbaillella and the widespread descendant Follicucullus. Te evolutionary appearance of Longtanella contributed to the atrophy of pseudothoraxic wings and size of the pseudothorax and lengthening of the total height of the test from Pseudoalbaillella and evolved to the Follicucullus group by complete reduction of undulated segmentation of the pseudoabdomen. Te appearance of Longtanella may relate to a regional warmer event afer the early Kungurian glacial period in East Gondwana and its extinction is likely related with diversifcation of the Follicucullus group in the early-middle Capitanian.

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Methods Sampling and analysis procedures. Forty-six samples were collected from all slices of the Bancheng Formation in the Shiti section. Te fragmented samples were soaked in a 4% HF solution for 10 h at room temperature. Afer discarding the supernatant liquid, the acid residues were transferred to other containers, then water was added until the residues were greatly diluted. Afer this step, the same steps were repeated more than 40 times for each sample. Disaggregated particles were wet-sieved through a 54 µm mesh sieve and dried at the end of 40 time steps. By preliminary observation under a binocular microscope, 38 samples with rich identifable radiolarians were selected. Individuals with distinguishable morphological characters were picked for species identifcation under a binocular microscope, and then better-preserved specimens (over 800 specimens in total) were photographed under the scanning electron microscope (Hitachi SU8010 in State Key Laboratory of Bio- geology and Environmental Geology, China University of Geosciences) for further morphological examination and species identifcation.

Meta‑database. Detailed occurrences of selected fusulinacean and radiolarian genera were mapped in the Japanese Islands, China and Sundaland. Sundaland includes the southern to southwestern parts of China, Viet- nam, Tailand, eastern Myanmar, Indonesia and Malaysia. Global distribution maps for the selected radiolarian genera include all other regions in the world. Following the current concepts of radiolarian genera, Longtanella, Pseudoalbaillella sensu stricto and the Follicucullus-group (Follicucullus and Cariver)12 were re-identifed from illustrated specimens in publications that are archived in Tohoku University (ca. 5800 papers). Te world palaeogeographic map was drawn based on the Middle to Late Permian tectonic reconstruction map (Fig. 2). Both the Pseudoalbaillella and Follicucullus group are widely distributed in the western and eastern margins of Palaeo-Tethys (Sicily 91; Turkey92; South China2; Malaysia93), east of the Meso-Tethys (Tailand 94), western Panthalassan Ocean (Far East Russia 95; Japan58; New Zealand 96), eastern Panthalassan Ocean (Brit- ish Columbia 97; Alaska48; South America98). By contrast, known occurrences of Longtanella are limited to the western Panthalassan Ocean, eastern margin of Palaeo-Tethys, east of Meso-Tethys, and part of the eastern Panthalassan Ocean. Tis suggests a more limited distribution for Longtanella than its ancestral Pseudoalbail‑ lella or descendent Follicucullus group. Te middle to upper Permian marine deposits with radiolarians in the Japanese Islands, China and Sundaland have been well studied since the 1980s. In these areas, any identifable morphotypes without a taxonomic name are regularly illustrated, and study of these indicates the occurrence of Longtanella at the level of tectonic divisions.

Statistical analysis. Statistical analyses of radiolarian palaeobioprovinces were performed with the occurrence or absence dataset (binary data) of co-occurrences of radiolarian and fusulinacean genera at the level of tectonic belts. Questionable occurrences or questionable identifcations were not included in the occurrence list. Te tectonic divisional scheme comes from published work for the Japanese Islands51, China13, and Sundaland99. For the present studies, we used CA (the dataset for CA is in supplement 7). CA is one of the most useful multivariate statistical methods to explore occurrence and absence data 88. In consideration of taxonomic stability at the genus level, the occurrences of seven fusulinacean genera in the Early-Middle Permian from China, Japan and Sundaland were compiled to analyze possible factors in the distribution of Longtanella. Te CA suggests that Longtanella difers from the Pseudoalbaillella and the Follicucullus group not only in palaeogeographic distribution, but also in its preferred temperature living conditions. Te CA, in short, can output independent factors that describe distributions. Te CA was performed with the statistical sofware R (R ver. 4.0.2) and plugins (RStudio ver. 1.3, Rcmdr ver. 2.7–0, RcmdrPlugin.EZR ver. 1.52, RcmderPlugin.FactoMineR ver 1.7).

Received: 20 January 2021; Accepted: 12 March 2021

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Competing interests Te authors declare no competing interests. Additional information Supplementary Information Te online version contains supplementary material available at https://doi.org/ 10.1038/s41598-021-86262-7. Correspondence and requests for materials should be addressed to Y.X. Reprints and permissions information is available at www.nature.com/reprints. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional afliations. Open Access Tis article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. Te images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

Scientifc Reports | (2021) 11:6831 | https://doi.org/10.1038/s41598-021-86262-7 13 Vol.:(0123456789) Supplement material-

New Permian radiolarians from east Asia and the quantitative reconstruction of their evolutionary and ecological significances

Yifan Xiao1*, Noritoshi Suzuki2, Tsuyoshi Ito3, Weihong He1

1State Key Laboratory of Biogeology and Environmental Geology, China University of

Geosciences, Wuhan 430074, PR China;

2Department of Earth Science, Graduate School of Science, Tohoku University, Sendai 980-8578, Japan;

3Research Institute of Geology and Geoinformation, Geological Survey of Japan, AIST, Tsukuba 305-8567,

Japan.

*Corresponding author. Email: [email protected]

Content

Supplement 1. Supplement figures.

Fig. S1 Cross-section of the Bancheng Formation, Shiti section, Guangxi province, China.

Figs S2–S5 Longtanella species from the Shiti section, Bancheng Formation.

Fig. S6 Occurrences of the selected fusulinacean genera and Longtanella in Japanese Islands (Tectonic map revised after Wallis et al. 2020).

Fig. S7 Occurrences of the selected fusulinacean and radiolarian genera in Mainland China (Tectonic map revised after Zheng et al. 2013).

Fig. S8 Occurrences of the selected fusulinacean and radiolarian genera in Sundaland (Tectonic map revised after Metcalfe 2013). Supplement 2. UAZ range (age) of each sample in the Shiti section decided by radiolarian species.

Supplement 3. Occurrences of fusulinaceans and radiolarians.

Supplement 4. The morphological terminology for the albaillellids in Figure 3, Supplement 5 and 6.

Supplement 5. Systematic Palaeontology.

Supplement 6. The character data set used for phylogenetic tree of Follicucullidae.

Supplement 7. Meta dataset for correspondence analysis (CA)

Supplement 1. Supplement figures.

Figure S1. Cross-section of the Bancheng Formation, Shiti section, Guangxi province, China. W.H.H. created this figure using CorelDRAW X4.

Figure S2. Longtanella species from the Shiti section, Bancheng Formation. Scale bar equals 100 μm for all figures, except from N (50 μm). A–P, Longtanella lanceoliformis Xiao & Suzuki sp. nov. A, 13ST9- 1_012; B, 13ST9-1_014; C, 13ST9-1_086; D, paratype, ST1-2_i001; E, ST1-2_i015; F, 13ST9-1_i003; G, ST1-2_i028; H, ST1-2_i030; I, ST1-2_i029; J, 13ST9-1_013; K, ST1-2_i031; L, 13ST9-1_087; M, holotype, ST 1-2_i026; N, ST 1-2_i026; O, ST1-2_i003; P, ST1-2_i032. Q–X, Longtanella edamame Xiao & Suzuki sp. nov. Q, holotype, 13ST15-4_148; R, 13ST6-1_010; S, 13ST15-4_149; T, ST1-3_i060; U, ST6-1_i001; V, 13ST6-1_248; W, paratype, 13ST6-1_245; X, 13ST6-1_250. Y, Z, Longtanella sp. 1. Y, ST6-1_i008; Z, ST9-1_i020. Photo credit: Y.F.X.

Figure S3. Longtanella species from the Shiti section, Bancheng Formation. Scale bar equals 100 μm for all figures. A–C, Longtanella sp. cf. Longtanella edamame Xiao & Suzuki sp. nov. A, ST1-2R_006; B, ST1-3_054; C, ST1-3_055. D–G, Longtanella jingyi Xiao & Suzuki sp. nov. D, paratype, ST1-3_061; E, ST1-3_i025; F, holotype, ST1-3_i053; G, ST1-3_i039. H–O, Longtanella tokkuriformis Xiao & Suzuki sp. nov. H, 13ST1-4_004; I, 13ST1-4_005; J, paratype, 13ST1-4_072; K, 13ST1-4_i005; L, holotype, ST1- 3_i045; M, ST1-6_i011; N, ST1-3_i030; O, ST3-5R_036. P–U, Longtanella turrita Xiao & Suzuki sp. nov. P, ST1-3_049; Q, paratype, ST1-3_048; R, holotype, ST1-3_052; S, ST1-3_050; T, ST1-3_051; U, ST1- 3_053. V–W: Longtanella sp. 2. V, 13ST15-4_153; W, ST1-3_i047. Photo credit: Y.F.X.

Figure S4. Longtanella species from the Shiti section, Bancheng Formation. Scale bar equals 100 μm for all figures. A–N, Longtanella kushidango Xiao & Suzuki sp. nov. A, ST9-1_i001; B, ST9-1_i002; C, ST9- 1_i003; D, ST9-1_i004; E, holotype, ST9-1_i005; F, ST9-1_i012; G, ST9-1_i007; H, paratype, ST9- 1_i006; I, ST9-1_i009; J, ST9-1_i013; K, ST9-1_i013; L, ST9R_178; M, ST9-1_i021; N, ST9R_181. O, P, Longtanella? sp. 3. O, ST1-2_i006; P, ST1-2_i035. Q–U, Longtanella laxiflexus Xiao & Suzuki sp. nov. Q, paratype, 13ST9-1_083; R, ST1-1_i001; S, ST1-1_i002; T, 13ST9-1_i004; U, holotype, ST1-2_i011; V, W, Longtanella sp. cf. Longtanella laxiflexus Xiao & Suzuki sp. nov. V, 13ST9-1_i007; W, 13ST9-1_i002. Photo credit: Y.F.X.

Figure S5. Longtanella species from the Shiti section, Bancheng Formation. Scale bar equals 100 μm for all figures. A–G, Longtanella follicucullinoides Xiao & Suzuki sp. nov. A, ST1-2_i025; B, ST1-2_i013; C, ST1-2_i005; D, ST1-2_i016; E, ST1-2_i010; F, holotype, ST1-2_i007; G, paratype, ST1-2_i022. H–J, Longtanella sp. 4. H, ST1-6_i006; I, ST1-6_i004; J, ST6-1_i005. K–P, Longtanella sp. 5. K, ST1-2_i008; L, ST1-3_i026; M, ST1-3_i032; N, ST1-3_i033; O, ST1-2_i004; P, 13ST6-1_257. Photo credit: Y.F.X.

Figure S6. Occurrences of the selected fusulinacean genera and Longtanella in Japanese Islands (Tectonic map revised after Wallis et al. 2020). T.I. created this figure using Adobe Illustrator CC 2017

(https://helpx.adobe.com/au/illustrator/release-note/illustrator-cc-2017-21-0-release-notes.html).

Figure S7. Occurrences of the selected fusulinacean and radiolarian genera in Mainland China (Tectonic map revised after Zheng et al. 2013). Y.F.X. created this figure using CorelDRAW X4.

Figure S8. Occurrences of the selected fusulinacean and radiolarian genera in Sundaland (Tectonic map revised after Metcalfe 2013). Y.F.X. created this figure using CorelDRAW X4.

Supplement 2. UAZ range (age) of each sample in the Shiti section decided by radiolarian species.

Abbreviations: Ps., Pseudoalbaillella; Al., Albaillella; R., Ruzhencevispongus; La., Latentifistula; H., Hegleria.

Supplement 3. Occurrences of fusulinaceans and radiolarians Genus Region Tectonic division References Monodiexodina Japan Islands Hida-gaien Niwa et al., 2004 Ueno and Tazawa, 2004 Kurosegawa belts Kanmera, 1963 Tazawa and Hasegawa, 2007 Mainland China (exclusive of the Heilongjiang province Han, 1980 Changning-Menglian Suture Zone) Han, 1981 Jilin province Han, 1980 Nei Mongol Han, 1976 Tibet province Wang et al., 1982a Nie and Song, 1983a Geng et al., 2012 Zhang et al. 2014b Zhang et al. 2013 Hainan province Sheng, 1965 Hubei province Chen, 1984 Jiangsu province Zhou and Zhang, 1984 Sichuan province Chen and Yang, 1978 Sundaland and the Changning- Ailaoshan Suture Igo et al., 1979 Menglian Suture Zone Inthanon Suture Zone Basir Jasin, 1991 Rauserella Japan Islands Akiyoshi Belt Kawano, 1961 Kobayashi, 2010 Kobayashi, 2012a Kobayashi, 2012b Kobayashi, 2013 Mino-Tamba-Ashio Belt Kobayashi, 2006a Kobayashi, 2011a Ueno et al., 2006 Kobayashi and Furutani, 2018 Northern and Southern Morikawa, 1956 Chichibu belts Kobayashi, 1988a Kobayashi, 2006b Kobayashi, 2007 Kobayashi, 2016 Mainland China (exclusive of the Anhui province Wang et al., 1982b Changning-Menglian Suture Zone) Guizhou province Yang, 1985 Jiangsu province Wang et al., 1982b Jilin province Han, 1980 Sichuan province Zhang and Wang, 1974 Chen and Yang, 1978 Sundaland and the Changning- Thailand Ueno, 2003 Menglian Suture Zone Yunnan province Zhou et al., 1987 Zhou, 2001 Shi et al., 2008 Afghanella Japan Islands Akiyoshi Belt Hanzawa, 1954 Kanmera, 1957 Toriyama, 1958 Kawano, 1961 Hanzawa and Murata, 1963 Kobayashi, 1988b Kobayashi, 2012a Kobayashi, 2012b

Ishii, 1990 Ozawa and Kobayashi, 1990 Ueno, 1992 Mainland China (exclusive of the Guangxi province Lin et al., 1977 Changning-Menglian Suture Zone) Li, 1989 Guizhou province Sheng, 1963 Liu et al., 1978 Yang, 1985 Chang et al., 2012 Hubei province Chen, 1984 Hunan province Lin et al, 1977 Xie, 1982 Sichuan province Chen and Yang, 1978 Zhang,1982 Sheng, 1963 Qinghai province Sheng, 1958 Sheng and Sun, 1975 Niu and Wu, 2016 Tibet province Zhang, 1982 Cheng et al., 2005 Xinjiang province Da and Su, 1983 Da, 1985 Sun and Zhang, 1985 Sun, 1993 Sundaland and the Changning- Yunnan province Zhou et a., 1987 Menglian Suture Zone Zhou, 2001 Shi et al., 2008 Thailand Toriyama and Kanmera, 1979 Dawson, 1993 Biwaella Japan Islands Akiyoshi Belt Kobayashi 2012b Kobayashi 2017 Mino-Tamba-Ashio Morikawa and Isomi, 1960 Morikawa and Isomi, 1961 Kobayashi and Furutani, 2009 Kobayashi and Furutani, 2019 Northern and Southern Morikawa and Kobayashi, 1960 Chichibu belts Ishizaki, 1962b Suyari, 1962 Kobayashi, 1993 Kobayashi, 2005a Kobayashi, 2005b Mainland China (exclusive of the Guangxi province Huang and Zeng, 1984 Changning-Menglian Suture Zone) Guizhou province Liu et al., 1978 Xia, 1994 Shaanxi province Wang and Sun, 1973 Ding et al., 1987 Ding et al., 1991 Xinjiang province Da and Su, 1983 Sundaland and the Changning- Thailand Sakagami and Iwai, 1974 Menglian Suture Zone Changning-Menglian Sakagami and Iwai, 1974 Misellina Japan Islands Hida-gaien Belt Niwa et al., 2004 Kurosegawa Belt Toriyama, 1947 Kanmera, 1956

Kanmera, 1963 Akiyoshi Belt Toriyama, 1958 Kobayashi, 1977 Ozawa and Kobayashi, 1990 Ueno, 1991 Mino-Tamba-Ashio Belt Morikawa and Isomi, 1961 Kobayashi, 2008 Northern and Southern Fujimoto, 1936 Chichibu belts Ishizaki, 1962a Ishizaki, 1963 Suyari, 1962 Takaoka, 1966 Kobayashi, 1977 Kobayashi, 2005a Sundaland and the Changning- Yunnan province Zhou et a., 1987 Menglian Suture Zone Zhou, 2001 Shi et al., 2008 Chusenella Japan Islands Akiyoshi Belt Sada and Yokoyama, 1966 Kobayashi, 1988b Kobayashi, 2010 Kobayashi, 2012a Kobayashi, 2012b Ozawa and Kobayashi, 1990 Davydov and Schmitz, 2019 Kurosegawa Belt Kanmera, 1954 Kobayashi, 2001 Tazawa and Hasegawa, 2007 Mino-Tamba-Ashio Belt Kobayashi, 2006a Kobayashi, 2011a Kobayashi and Furutani, 2018 Southern Chichibu belts Kobayashi, 2005a Kobayashi, 2006b Kobayashi, 2007 Kobayashi, 2011b Mainland China (exclusive of the Anhui province Wang et al., 1982b Changning-Menglian Suture Zone) Guangxi province Hsu, 1942 Sheng, 1963 Lin et al., 1977 Li, 1989 Guizhou province Liu et al., 1978 Yang, 1985 Hubei province Chen, 1956 Lin et al., 1977 Lin, 1984 Hunan province Chen, 1956 Lin et al., 1977 Xie, 1982 Cao et al., 2013 Jiangsu province Wang et al., 1982b Jiangxi province Wang et al., 1982b Sichuan province Zhang and Wang, 1974 Chen and Yang, 1978 Zhang, 1982

Shaanxi province Sheng, 1956 Qinghai province Niu and Wu, 2016 Xinjiang province Da and Su, 1983 Sun and Zhang, 1988 Gaetani and Leven, 2014 Tibet province Chu, 1982 Wang et al., 1982a Zhang, 1982 Nie and Song, 1983b Wang and Zhou, 1986 Cheng et al., 2005 Zhang et al., 2009 Zhang et al., 2013 Zhang et al., 2014 Zhang et al., 2019 Jilin province Han, 1980 Sundaland and the Changning- Yunnan province Zhou, 2001 Menglian Suture Zone Shi et al., 2005 Shi et al., 2008 Shi et al., 2017 Huang et al., 2009 Huang et al., 2015 Huang et al., 2017 Myanmar Zhang et al., 2020 Thailand Pitakpaivan, 1966 Sakagami and Iwai, 1974 Toriyama, 1976 Toriyama and Kanmera, 1977 Toriyama and Kanmera, 1979 Dawson, 1993 Ueno, 2003 Pisolina Mainland China (exclusive of the Anhui province Regional Geological Survey Team of Changning-Menglian Suture Zone) Wang et al., 1982b Fujian province Hong et al., 1986 Guangxi province Lin et al., 1977 Li, 1989 Guizhou province Liu et al., 1978 Yang, 1985 Zhang et al., 2010 Hubei province Lin et al., 1977 Chen, 1984 Lin, 1984 Hunan province Lin et al., 1979 Zhang and Hong, 2000 Jiangsu province Zhou and Zhang, 1984 Qinghai province Sheng and Sun, 1975 Shaanxi province Sheng, 1956 Sichuan province Zhang and Wang, 1974 Chen and Yang, 1978 Zhang, 1982 Zhejiang province Wang and Tang, 1986 Sundaland and the Changning- Indochina Terrane Ueno and Sakagami, 1993 Hori, 2004

Longtanella Japan Islands Northern and Southern Ujiie and Oba, 1991 Chichibu belt Sashida, 1995 Suyari et al., 1982 Kurosegawa Belt Hada et al., 1992 Kashiwagi and Isaji, 2015 North Kitakami Belt Kametaka et al., 2005 Hida-gaien Belt Niko et al., 1987 Nagato Tectonic Zone Isozaki and Tamura, 1989 Mino Terrane Ito et al., 2016 Far east Russia Sikhote-Alin Rudenko and Panasenko, 1997 British Columbia Harms and Murchey, 1992 Mainland China (exclusive of the Qinghai province Li and Bian, 1993 Changning-Menglian Suture Zone) Guangxi province Yao et al., 2004 Shimakawa and Yao, 2006 Wu et al., 1994 Guizhou province Yu, 1996 Jiangsu province Sheng and Wang, 1985 Hubei province Shi et al., 2016 Anhui province Nagai et al., 1998 Kametaka et al., 2009 Ito et al., 2013 Wang, 1993a Sundaland and the Changning- Yunnan province Feng, 1992 Menglian Suture Zone Bentong-Raub suture Basir Jasin and Che Ali, 1997 zone Metacalfe et al., 1999 Sra Kaeo Suture Zone Saesaengseerung et al., 2009 Udchachon et al., 2018 Inthanon Suture Zone Sashida et al., 1993

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Supplement 4. The morphological terminology for the albaillellids in Figure 3, Supplement 5 and 6.

Supplement 5. Systematic Palaeontology.

The general morphological terminology for Permian albaillellids is summarized in supplement 4. Undulation patterns in the main species of Longtanella are shown in Figure 3. The specimens described herein are registered and deposited in the China University of Geosciences, Wuhan. We also include an updating of the genus concept since the paper of Xiao et al. (2020) was published.

Order Albaillellaria Deflandre, 1953, emend. Holdsworth, 1969 Family Follicucullidae Ormiston & Babcock, 1979 Follicucullidae. Ormiston & Babcock, 1979, p. 332; Holdsworth & Jones, 1980, p. 285; Kozur, 1981, p. 265; Nazarov, 1988, p. 83; Kozur & Mostler, 1989, p. 169-170; Cordey, 1998, p. 37; Afanasieva, 2000, p. 124-125; De Wever et al., 2001, p. 93, 95; Afanasieva, 2002, p. 28; Afanasieva et al., 2005, p. S270; Afanasieva & Amon, 2006, p. 105. Holdsworthellidae. Kozur, 1981, p. 268. Follicullidae [sic] (=Follicucullidae). Petrushevskaya, 1984, p. 125. Pseudoalbaillellidae. (nomen nudum) Cheng, 1986, p. 45; (nomen nudum) Wang & Qi, 1995, p. 380. Follicucullinae. Kozur & Mostler, 1989, p. 170-171 [in Follicucullidae]; Afanasieva, 2000, p. 125 [in Follicucullidae]. Holdsworthellinae. Kozur & Mostler, 1989, p. 182-183 [in Follicucullidae]. Remarks. The Paleozoic Genera Working Group organized by the International Association of Radiolarists (InterRad) decided to include only three genera, Follicucullus, Ishigaconus and Parafollicucullus, in the Follicucullidae (Caridroit et al. 2017; Noble et al. 2017). Soon after this decision, several claims arose from several non-member specialists about this excessive synonymization (Ito 2020; Nakagawa & Wakita 2020; Nestell & Nestsell 2020). Quantitative and parsimony methods as an objective test of the genus-level classification resulted in a scheme of ten genera (Cariver, Curvalbaillella, Follicucullus, Haplodiacanthus, Holdsworthella, Kitoconus, Longtanella, Parafollicucullus sensu stricto, Parafollicucullus in Lineage I (= Parafollicuculinoides gen. nov. in this paper), and Pseudoalbaillella sensu stricto) (Xiao et al. 2020, Fig. 5).

Genus Parafollicucullinoides Xiao, Ito & Suzuki, gen. nov.

Type species. Pseudoalbaillella globosa Ishiga & Imoto in Ishiga et al. (1982). Included species (written as an original described name). Pseudoalbaillella lomentaria Ishiga & Imoto, 1980; Pseudoalbaillella ornata Ishiga & Imoto, 1980; Pseudoalbaillella longtanensis Sheng & Wang, 1985; Pseudoalbaillella yanaharensis Nishimura & Ishiga, 1987. Diagnosis. Shell divided into apical cone, pseudothorax, pre-pseudoabdomen and segmented pseudoabdomen (culmina and crenellae). Apical cone straight or slightly curved, about 1/3 of the length of the test. Pseudothorax globular and usually inflated. Two blade-like wings long and wide. Pre- pseudoabdomen waist short. Pseudoabdomen with two or more culmina similar in height. Derivation of name. Named for its similarity to Parafollicucullus, but different. Range and distributions. Permian, Late Sakmarian to middle Capitanian. Remarks. The new genus contains the species of ‘Parafollicucullus’ in lineage I in Xiao et al. (2020). They differ from the true Parafollicucullus of lineage Ⅱ in having a long straight apical cone, inflated pseudothorax, and segmented pseudoabdomen.

Genus Curvalbaillella Kozur & Mostler, 1989, sensu emend. Xiao & Suzuki herein Curvalbaillella. Kozur & Mostler, 1989, p. 179–180; Noble et al., 2017, p. 427–428. Kitoconus. Kozur & Mostler, 1989, p. 178; Catalano et al., 1989, p. 95. Type species. Pseudoalbaillella u-forma Holdsworth & Jones, 1980. Revised diagnosis. Shell divided into apical cone, pseudothorax, and unsegmented pseudoabdomen. Apical cone short, straight or slightly curved. Pseudothorax globular and not inflated, less than 1/5 of the length of the test. Two hemicyclic-like wings short to medium in length which extend horizontally with straight shoulders. Pseudoabdomen unsegmented, almost invariant in width, straight or curved, sometimes the basal part of the test forms ‘U’-shape. Two flaps of the same size at the aperture, often extending in different directions (horizontal or upwards). Remarks. Statistical test made by Xiao et al. (2020) pointed to the conspecific relationship of Curvalbaillella and Kitoconus. The difference between these genera is in the curved (Curvalbaillella) or straight (Kitoconus) pseudoabdomen. The known Curvalbaillella species (Pseudoalbaillella bulbosa Ishiga, 1982; Pseudoalbaillella chilensis Ling & Forsythe, 1987; Pseudoalbaillella u-forma Holdsworth & Jones, 1980; and Pseudoalbaillella u-forma reflexa Ling & Forsythe, 1987, in the original description) do

25 not look like intermediate species for the Kitoconus genus (Pseudoalbaillella elegans Ishiga & Imoto, 1980 and Pseudoalbaillella elongata Ishiga & Imoto, 1980, in the original description) as was discussed by Kozur and Mostler (1989) . Some species identified as ‘Pseudoalbaillella elegans’ show a somewhat curved pseudoabdomen (eg., pl. 11, figs 10, 13, pl. 13, figs 11, 12 in Wang et al. 2012), supporting the statistically objective opinion of Xiao et al. (2020). Considering that the only obvious difference of Kitoconus and Curvalbaillella is the curvature of pseudoabdomen, we regard them as synonyms. Both Kitoconus and Curvalbaillella were established in the same paper (Kozur & Mostler 1989), but Curvalbaillella has objective priority over Kitoconus because the latter was established as a subgenus of the genus Pseudoalbaillella.

Genus Pseudoalbaillella Holdsworth & Jones, 1980 sensu emend. Xiao & Suzuki herein Pseudoalbaillella. Holdsworth & Jones, 1980, p. 285; Noble et al., 2017, p. 430. Type species. Pseudoalbaillella scalprata Holdsworth & Jones, 1980. Included species (written as an original described name). Pseudoalbaillella rhombothoracata Ishiga & Imoto, 1980; Pseudoalbaillella scalprata m. postscalprata Ishiga 1983; Pseudoalbaillella scalprata praescalprata Kozur 1989; Pseudoalbaillella triangularis Wang et al., 2012; Pseudoalbaillella desmoinesiensis Nestell et al., 2012. Revised diagnosis. Shell divided into apical cone, pseudothorax, and unsegmented pseudoabdomen. Apical cone strongly curved. Pseudothorax triangular or rhombohedral, usually inflated, wider than pseudoabdomen and more than 1/3 of height of the test. Two long and wide blade-like wings which are oriented downwards with arc-shaped shoulder. Pseudoabdomen unsegmented, almost invariant in width and similar to pseudothorax in height. Two flaps of the different size at the arc-shaped basal aperture, often extending downwards vertically or obliquely. Remarks. Holdsworth and Jones (1980) established Pseudoalbaillella and Parafollicucullus, and in the origin descriptions, Parafollicucullus differs from Pseudoalbaillella only for possessing a ring-like pre- pseudoabdominal segment between pseudothorax and pseudoabdomen. Later Kozur (1981) proposed Pseudoalbaillella is a synonym of Parafollicucullus because he thought these radiolarians usually having ring-like pre-pseudoabdominal segment. Most researchers agree with that these two genera are synonyms (Ishiga et al. 1982; Sheng & Wang 1985; De Wever et al. 2001; Nestell et al. 2012; Noble et al. 2017),

26 except for which name has the priority. However, Statistical test made by Xiao et al. (2020) implied that Pseudoalbaillella could be distinguished from Parafollicucullus in both morphology (eg., the size and shape of apical cone and pseudothorax of Pseudoalbaillella are much greater than Parafollicucullus) and phylogeny (they belong to different lineages). Their taxonomic and phylogenetic analysis supports that Follicucullus is more related to Pseudoalbaillella, compared to Parafollicucullus. Thus, the definition of Pseudoalbaillella is revised herein, and separated from Parafollicucullus.

Genus Parafollicucullus Holdsworth & Jones, 1980 sensu emend. Xiao & Suzuki herein Parafollicucullus. Holdsworth & Jones, 1980, p. 285; Noble et al., 2017, p. 430. Type species. Parafollicucullus fusiformis Holdsworth & Jones, 1980. Revised diagnosis. Shell divided into apical cone, pseudothorax, and segmented pseudoabdomen. Apical cone slightly curved. Pseudothorax globular, less than 1/5 of the height of the test. Two symmetric narrow blade-like wings which are oriented downwards with straight shoulder. Pseudoabdomen with variable number of segments or bands, widest in the middle part. Pre-pseudoabdomen between the pseudothorax and pseudoabdomen, often constricted. Two flaps at the arc-shaped basal aperture. Remarks. The concept of Parafollicucullus are expanded here, keeping its original important character “pre-pseudoabdominal segment”. The species of Parafollicucullus include the species of ‘Parafollicucullus’ in lineage II in Xiao et al. (2020).

Genus Follicucullus Ormiston & Babcock, 1979 Follicucullus. Ormiston & Babcock, 1979, p. 332; Noble et al., 2007, p. 428. Ishigaconus. Kozur & Mostler, 1989, p. 181; Noble et al., 2007, p. 429. Type species. Follicucullus ventricosus Ormiston & Babcock, 1979. Remarks. Noble et al. (2017) favoured the opinion of Kozur & Mostler (1989) who separated Ishigaconus (type species: Follicucullus scholasticus Ormiston & Babcock, 1979) from Follicucullus (type species: Follicucullus ventricosus) because of a lack of any inflation of the pseudothorax in Follicucullus scholasticus. The statistical results in Xiao et al. (2020, figs. 3, 4) showed a difference between F. ventricosus and F. scholasticus, confirming the opinion of Kozur & Mostler (1989). However, the difference scores are small in Xiao et al. (2020, fig. 3), and if we accept this score as a clustering line, the

Follicucullidae should be subdivided into 12 genera under the HQT-II statistical method, but this distinction cannot be supported by time-calibrated phylogenetic trees shown by Xiao et al. (2020, fig. 4). In consideration of the objective taxonomic scheme in Xiao et al. (2020), we apply the original concept of Follicucullus, with Ishigaconus as junior synonym.

Genus Longtanella Sheng & Wang, 1985 sensu emend. Ito, 2020, sensu emend. Xiao, Ito & Suzuki herein Longtanella Sheng & Wang, 1985, p. 175; Kozur & Mostler, 1989, p. 179; Ito et al., 2019, p. 270; Ito, 2020, p. 1–10. Type species. Longtanella zhengpanshanensis Sheng & Wang, 1985 Revised diagnosis. Shell smooth, straight, bilaterally symmetrical or slightly asymmetric. The shell divided into apical cone, pseudothorax, pre-pseudoabdomen and pseudoabdomen. Apical cone and pseudothorax small, wings retrograde to bulge-shaped or absent. Pre-pseudoabdomen long and slender. Pseudoabdomen with or without undulation. In some specimens, four flaps present. These four flaps can be divided into two groups: a pair of large ones at front and back sides, and a pair of small ones at left and right sides (see Fig. 3I), extending vertically or a little obliquely from apertural margin downward. Occurrences. The upper part of the Gufeng Formation in Chaohu City, Anhui province (Wang 1993; Nagai et al. 1998; Kametaka et al. 2009; Ito et al. 2013), Qinzhou city, Guangxi province province (Xia & Zhang 1998; Wang et al. 2012; this study), Hubei province province (Feng 1992) and Longtan region in Nanjing, South China (Sheng & Wang 1985); ophiolite complex in Qinghai province, western China (Li & Bian 1993); bedded chert of the Sra Kaeo Suture Zone in eastern Thailand (Saesaengseerung et al. 2009); Bentong-Raub Suture Zone in Malaysia (Metcalfe et al. 1999; Spiller 2002); Japan (Niko et al. 1987; Kashiwagi & Isaji 2015); Far East of Russia (Rudenko & Panasenko 1997). Ranges. Kungurian to Wordian. Remarks. The original definition of Longtanella is ‘Shell smooth, straight, bilaterally symmetrical turriformis. The shell wall divided into the spire, the turri-body, the turri-body and the turri-bottom composed of ring-like swollen segments, last segment constrictive with 4 flaps vertically extending downward’ (Sheng & Wang 1985, p. 179) and its diagnosis was revised as ‘Follicucullidae having a smooth, straight, bilaterally symmetrical, and turriform shell composed of a conical proximal spired part and a bladder, with or without a skirt’ (Ito 2020). Because our specimens also have similar overall

28 appearances with Longtanella, we redefined Longtanella and included our specimens in this genus. Ito (2020) applied different morphological terminology such as ‘spired part’, ‘bladder’ and ‘skirt’. Our paper, though Ito is a co-author, prioritized the more common general terms that are applicable to any follicucullids (see supplement 6 for the general characters of Follicucullidae). The fundamental difference of Longtanella from all other Follicucullidae genera is in its symmetrical morphology on the front-back view and the ‘dorsal-lateral view’. This character excludes confusion with poorly-preserved specimens of any other genera. Noble et al. (2017, p. 430) synonymized Longtanella with ‘Parafollicucullus’ by ambiguous reasoning, namely, ‘this genus fits within the parameters of Parafollicucullus and should be treated as a junior synonym.’ Ito (2020) proved the independence of Longtanella by observation of topotypes, and Xiao et al. (2020) confirmed this validity by statistical methods. Besides our standpoints and those of the original authors (Sheng & Wang 1985), the validity of Longtanella is accepted in many reports on Asian Permian radiolarians (Rudenko 1991; Nagai et al. 1998; Xia & Zhang 1998; Kametaka et al. 2009; Saesaengseerung et al. 2009; Muhanmad Ashahadi et al. 2016). By contrast, studies in North America are likely to reject its validity (Cordey 1998; Nestell & Nestell 2010; Nestell et al. 2012; Noble et al. 2017). This contradiction may be because North America, except for British Columbia, lies outside the Longtanella territory. The orientation of the follicucullid test is defined by Cartesian coordinates with the ventral-dorsal side, front-back side and top-bottom side axes (Nakagawa & Wakita 2020). This scheme needs to be slightly revised because Longtanella shows a symmetrical appearance with respect to the dorsal-ventral and the front-back axes. Taking into account of the realistic recognition of shell orientation, we simply apply the terms left-right side instead of dorsal-ventral side. Front-back side (Fig. 3A) as well as top-bottom side are simply applied for the external display. The genus Longtanella was originally included in the family Albaillellidae by Sheng & Wang (1985). The Albaillellidae has visible caveal ribs from the a-rod side (De Wever et al. 2001, p. 92) but Longtanella lacks these ribs. Feng (1992) depicted two Longtanella species, Longtanella mengshengensis and Longtanella turgida in the Parafollicucullus fusiformis assemblage, but he only described L. turgida but not L. mengshengensis. Subsequently L. mengshengensis is a nomen nudum.

Longtanella lanceoliformis Xiao & Suzuki sp. nov.

(Figs 3E, S2A–P) 1991 Follicucullus scholasticus Ormiston & Babcock; Ujiie & Oba: pl. 3, fig. 2. 1993 Pseudoalbaillella sp. H; Li & Bian: 416, 417, pl. 1, fig. 11. 1994 Pseudoalbaillella sp. B; Wu et al.: pl. 2, fig. 7. Diagnosis. Longtanella with ‘short spear’ lance-like shell. Derivation of name. Latin adjective, lanceola + formis (-is, -e), meaning lance-formed. Noun. Material. Holotype, Fig. S2M, specimen ST 1-2_i026; paratype, Fig. S2D, specimen ST1-2_i001. 21 specimens were examined by SEM and 16 of them are illustrated. Measurements (μm). Height of shell 223–357 (mean 279); height of AC 12–35 (mean 21); width of the basal AC 17–39 (mean 29); height of PPA 34–76 (mean 61); height of PX 26–63 (mean 42); width of PX 22–64 (mean 40); height of PA 130–214 (mean 162); based on 19 specimens. Occurrences. Middle Kungurian–Roadian; Ophiolite complex and the Bancheng Formation of South China, the Southern Chichibu Belt of Japan (Ujiie & Oba 1991). Description. Shell smooth, consisting of inflated oblong apical (AC) to pseudothoracic (PX) part, long thin pre-pseudoabdomen (PPA), and relatively wider pseudoabdomen (PA). AC very short (<1/2 of the PX height), straight, pyramidal with an apical spinulus. AC gradually joining with PX. PX slightly inflated without wings, forming ‘spearhead’ of a lance together with AC. No distinguishable pseudothoracic bulb in PX. The distal part of PX obviously narrowing to connect with PPA. PPA long and slender (1.3 to 1.5 times longer than the height of AC-PX part and 1/2 to 4/5 of the maximum width of PX). Pre-pseudoabdominal junction is easily distinguishable but does not form stricture. PA cylindrical and distally widening, straight. The crenelle between the distal end of PA and basal skirt (BS) is variable from smooth to necked connection. Some specimens with distinguishable BS is curved to the dorsal side. Four flaps vertically or little obliquely extending from apertural margin downward (Fig. S2M). Remarks. This new species is similar to Follicucullus porrectus Rudenko in Belyansky et al., 1984 in having a slender and smooth shell, but the new species is distinguished from the latter by the spearhead- like apical cone–pseudothoracic part and the presence of four flaps on the basal skirt. According to Nestell et al. (2019, p. 278), Follicucullus japonicus Ishiga, 1991 has a wider pseudothorax, a shorter and not cylindrical pseudoabdomen and two distinctive apertural flaps whereas F. porrectus lack apertural flaps. Besides the validity discussion on F. japonicus, no similar morphotypes with four flaps have been known in

Follicucullus.

Longtanella edamame Xiao & Suzuki sp. nov. (Figs 3D, S2Q–X) Diagnosis. Longtanella having edamame-shaped pseudoabdomen. Derivation of name. Edamame, noun, meaning a kind of soybean in Japanese. It literally means ‘stem beans’ with a few undulating zones. This species is named for the edamame-like outline of the pseudoabdomen. Material. Holotype, Fig. S2Q, specimen 13ST15-4_148; paratype, Fig. S2W, specimen 13ST6-1_245. 12 specimens were examined by SEM and eight of them are illustrated. Measurements (μm). Height of shell 216–340 (mean 270); height of AC 15–49 (mean 34); width of the basal AC 18–36 (mean 24); height of PPA 22–87 (mean 43); height of PX 22–58 (mean 38); width of PX 26–55 (mean 43); height of PA 109–270 (mean 183); based on 12 specimens. Occurrences. Middle Kungurian–Roadian; Bancheng Formation of South China. Description. Shell consisting of an acute apical cone, small rhomboidal to simple inflated pseudothorax with wing-like acute edges in its both sides, short and thin pre-pseudothorax, two to three undulated segments on pseudothorax with basal skirt. PX is generally rhombic in outline but it also appears as a laterally facing oblong pseudothoracic bulb. PX is separated from PPA in the case of smaller PX with a stricture whereas from PPA in the case of rhombic PX with a distinguishable connection. PPA cylindrical with or without one annular ridge (Fig. S2W, X for annular ridge). Height of PPA is nearly similar throughout shell and its height is variable from 22 μm in the case of no annular ridge to 87 μm in the case of the presence of distinctive annular ridge. PA joined with PPA by a significant stepped shoulder. PA is fusiform in shape with three crenellae to form an edamame-like outline. BS cylindrical, usually curving to ventral side. Remarks. This new species is unique in exhibiting morphological instability in the distinction between the pre-pseudothorax and upper part of the pseudothorax. Some specimens (e.g. Fig. S2U) look as if they have a short pre-pseudothorax with thinner upper pseudothorax or a stepped pre-pseudothorax. Others (e.g. Fig. S2W, X) look as if they have two crenellae and one culmen as a long and thin pre-pseudothorax. In consideration of these unstable morphological characters, the proximal end of the pseudothorax is

31 determined by the proximal end of the ‘spindle’ form (Fig. S2W, X). This new species is similar to Parafollicucullus longtanensis (Sheng & Wang, 1985) in having a similar shell, but the latter has an undulated pseudothorax with two long wings.

Longtanella jingyi Xiao & Suzuki sp. nov. (Fig. S3D–G) 2016 Longtanella? sp. A; Ito et al.: fig. 4.5. Diagnosis. Longtanella with a rhombic pseudothorax with tiny wings and elegant shaped smooth pseudoabdomen, and symmetric shell. Derivation of name. Chinese word ‘jing’ and ‘yi’, meaning ‘excellent’ and ‘beautiful’, respectively, for a lady. Material. Holotype, Fig. S3F, specimen ST1-3_i053; paratype, Fig. S3D, specimen ST1-3_061; Eight specimens were examined by SEM and four of them are illustrated. Measurements (μm). Height of shell 123–250 (mean 217); height of AC 23–58 (mean 36); width of the basal AC 21–40 (mean 27); height of PPA 19–64 (mean 40); height of PX 24–52 (mean 32); width of PX 31–63 (mean 50); height of PA 95–150 (mean 123); based on eight specimens. Occurrences. Middle Kungurian–Roadian; Bancheng Formation of South China and pebble of the Kamiaso Unit of the Mino-Tamba-Ashio Belt of Japan. Description. Shell consisting of a tiny apical cone, rhombic pseudothorax, pre-pseudoabdomen and smooth pseudoabdomen without undulation. AC with acute apex, similar in the height with PX; PX a little inflated with very short wings, forming an arrow shape together with AC. PPA cylindrical, growing wider downwards with no constriction with the PA, usually 1.2 times longer than the height of PX. PA fusoid and smooth without undulation. Posterior waist reduced in both dorsal and ventral sides; basal skirt short with flaps. Remarks. The taxonomic position of this new species at the genus level could be confused with a poorly preserved Parafollicucullus by loss of the wings if we refer to Parafollicucullus corniculatus (Rudenko & Panasenko, 1990) and Parafollicucullus internata (Wang in Wang et al., 2012). However, Longtanella jingyi sp. nov. differs from any Parafollicucullus species by its symmetrical shell. This new species is probably homeomorphic with some Parafollicucullus (eg. Pa. corniculatus and Pa. internata). However,

32 the small acute apical cone and rhombic pseudothorax without wings in L. jingyi sp. nov. form an arrow shape, and subsequently makes it easy to differentiate from the Parafollicucullus species with a distinct longer cone-shaped AC. This new species differs from Longtanella cf. edamame sp. nov. in having a smooth pseudoabdomen without undulation.

Longtanella tokkuriformis Xiao & Suzuki sp. nov. (Figs 3E, S3H–O) 2015 Pseudoalbaillella sp. cf. P. monacantha; Kashiwagi & Isaji: 42, figs 5.1–5.4. 2016 Follicucullus monacanthus Ishiga et al.; Shi et al.: fig. 4.16. 2016 Follicucullus dilatatus Belyansky; Shi et al.: fig. 4.17. 2020 Longtanella sp. B; Ito: figs 6F–6J. Diagnosis. Longtanella with strongly constricted pre-pseudoabdomen and strongly inflated oblong pseudoabdomen. Derivation of name. Noun. Tokkuri is a Japanese traditional small bottle with strong neck and large body for alcohol. Material. Holotype, Fig. S3L, specimen ST1-3_i045; paratype, Fig. S3J, specimen 13ST1-4_072. 19 specimens were examined by SEM and 8 of them are illustrated. Measurements (μm). Height of shell 136–247 (mean 200); height of AC 15–51 (mean 31); width of the basal AC 14–31 (mean 24); height of WA 19–55 (mean 33); height of PX 24–45 (mean 34); width of PX 29–58 (mean 41); height of PA 79–152 (mean 116); based on 17 specimens. Occurrences. Middle Kungurian–Roadian; Gufeng and Bancheng Formation of South China and Permian pebbles in the Lower Cretaceous Choshi Group of the Kurosegawa Belt in Japan. Description. Shell consisting of conical apical cone, small pseudothorax, strongly constricted pre- pseudoabdomen, and strongly inflated oblong and smooth pseudoabdomen without any undulation. AC is conical in shape and its shape varies from thin acute (Fig. S3H, K, L) to thick acute (Fig. S3M, N). PX slightly inflated with rudimentary wings. PPA is cylindrical and widens downwards. PA particularly wide, sharply increasing downwards, stocky in the bottom; no constriction existed between the pseudothorax and basal aperture. More than two flaps visible but the exact number is unknown. Remarks. This new species has a wide shell body. This new species was illustrated as Parafollicucullus

33 monacanthus (Ishiga and Imoto in Ishiga et al., 1982) or Follicucullus dilatatus Rudenko in Belyanskiy et al., 1984 in previous papers (Kashiwaki & Isaji, 2015; Shi et al. 2016). These two species are different from Longtanella tokkuriformis sp. nov., because Pa. monacanthus has one tiny wing on the dorsal side (e.g. Ito et al. 2015) and the pseudothorax of F. dilatatus is not obvious and inflated, compared to the new species. The new species is also similar to Parafollicucullus bella (Sheng & Wang, 1985) in the shell outline, but the latter has a distinguishable basal skirt with obvious constriction, wider pseudothorax and basal skirt in comparison with its pseudothorax, typical wing, and undulated apical cone.

Longtanella turrita Xiao & Suzuki sp. nov. (Figs 3F, S3P–U) 2016 Longtanella zhengpanshanensis Sheng & Wang; Shi et al.: fig. 4.13. 2020 Longtanella sp. A; Ito: fig. 6A. Diagnosis. Longtanella with turreted shell. Derivation of name. Latin adjective, turritus, -a, -um, meaning having a tower. Material. Holotype, Fig. S3R, specimen ST1-3_052; paratype, Fig. S3Q, specimen ST1-3_048. Six specimens were examined by SEM and six of them are illustrated. Measurements (μm). Height of shell 131–294 (mean 216); height of AC 17–33 (mean 21); width of the basal AC 17–26 (mean 22); height of PPA 33–66 (mean 44); height of PX 26–36 (mean 32); width of PX 35–50 (mean 41); height of PA 141–209 (mean 166); based on six specimens. Occurrences. Middle Kungurian–Roadian; Gufeng and Bancheng Formation of South China. Description. Shell tower-like, bilaterally symmetrical or slightly asymmetric, consisting of tiny apical cone, small pseudothorax, constricted pre-pseudoabdomen, and pseudoabdomen with culmina and crenellae. PX spherical, moderately inflated. PPA cylindrical and slim, 1.3–1.8 times longer than the height of PX. PA terraced, composed of 2–4 culmina which increase downwards, the first one shortest and sharply strictured. Flaps vertically or little bit obliquely extending from apertural margin downward. Remarks. This new species was once identified as Longtanella zhengpanshanensis Sheng & Wang, 1985 by Shi et al. (2016). Reinvestigating the topotypes, Ito (2020) confirmed no intermediate forms between L. zhengpanshanensis in the sense of Shi et al. (2016) and L. turrita sp. nov..

Longtanella kushidango Xiao & Suzuki sp. nov. (Figs 3A, S4A–N) 1987 Pseudoalbaillella lomentaria Ishiga & Imoto; Niko et al.: pl. 1, figs 3–6. 1992 Longtanella mengshengensis Feng [nomen nudum]; Feng: pl. 1, figs 9, 10. 1992 Albaillella sp.; Hada et al.: fig. 4F. 1992 Parafollicucullus lomentaria Ishiga & Imoto; Harms & Murchey: pl. 1, fig. B. 1993 Pseudoalbaillella lomentaria Ishiga & Imoto; Li & Bian: 416, pl. 1, fig. 12. 1993 Pseudoalbaillella lomentaria Ishiga & Imoto; Sashida et al.: fig. 6.5. 1994 Pseudoalbaillella lomentaria Ishiga & Imoto; Wang et al.: 181, pl. 1, figs 12, 13. 1995 Pseudoalbaillella lomentaria Ishiga & Imoto; Sashida: 39, fig. 5.18. 1997 Pseudoalbaillella lomentaria Ishiga & Imoto; Basir Jasin & Che Aziz: 331, pl. 1, fig. 1. 1997 Pseudoalbaillella annulata Ishiga; Rudenko & Panasenko: pl.1, fig. 1 1997 Pseudoalbaillella cf. lomentaria; Rudenko & Panasenko: pl.1, fig. 2. 1998 Pseudoalbaillella lomentaria Ishiga & Imoto; Sashida et al.: fig.11. 16, 17. 2004 Pseudoalbaillella aff. lomentaria; Hori: pl. 1, figs. 25, 32. 2006 Pseudoalbaillella lomentaria Ishiga & Imoto; Shimakawa & Yao: pl. 1, figs 12, 13. 2012 Pseudoalbaillella nodosa Ishiga; Wang et al.: 47, pl. 11, figs 5, 6, 8. 2018 Pseudoalbaillella sp. cf. P. lomentaria; Udchachon et al.: 1460, figs 7h, 7k. Diagnosis. Longtanella with cylindrical tube-like pseudothorax and pseudoabdomen with three to four culmina. Derivation of name. Latinized word from Japanese, meaning a bunch of vertically arrayed three pasted rice balls of Japanese traditional sweet. Material. Holotype, Fig. S4E, specimen ST9-1_i005; paratype, Fig. S4H, specimen ST9-1_i006. 17 specimens were examined by SEM and 14 of them are illustrated. Measurements (μm). Height of shell 233–344 (mean 295); height of AC 56–98 (mean 81); width of the basal AC 40–61 (mean 51); height of PPA 17–39 (mean 29); height of PX 44–69 (mean 54); width of PX 78–107 (mean 93); height of PA 104–167 (mean 136); based on 16 specimens. Occurrences. Middle Kungurian–Wordian; Gufeng and Bancheng Formation of South China; the Mizuyagadani Formation of the Hida-gaien Belt, Chichibu composite belts, Japan; Malaysia; Thailand;

Sylvester Allochthon in the Canadian Cordillera, Canada; British Columbia, the Samarka Terrane in the Primorye, Far East Russia. Description. Shell inflated and kushidango-like outline in shape. It consists of weakly undulated apical cone, inflated pseudothorax, short pre-pseudoabdomen, and pseudoabdomen with well-developed culmina and crenellae. AC usually with 3–5 segments but sometimes without segments (e.g. Fig. S4N). The total height of AC about 1/4 to 1/3 height of the shell. PX spherical, strongly inflated without wings, about 2/5 height of the PA. PPA short, about 1/2 height of the PX. PA composed of two inflated culmina,culmina of the same width of the pseudothorax; these culmina and crenellae oblique to the pseudothorax in some specimens, probably corresponding to the anatomical dorsal or ventral side (e.g. Fig. S4C–E). Posterior waist between the lower segment and basal skirt is narrower than segmental waists. BS similar to the segmental waist in width, but much narrower than the culmen. Remarks. This new species is commonly illustrated worldwide (see occurrence) but has been confused with Parafollicucullus lomentarius (Ishiga & Imoto, 1980). However, Pa. lomentarius is fundamentally different from L. kushidango sp. nov. by having wings. Longtanella mengshengensis Feng, 1992 (pl. 1, figs 9, 10 in Feng 1992) is a nomen nudum, as we mentioned earlier. Rudenko & Panasenko (1997) illustrated L. kushidango sp. nov. as ‘Pseudoalbaillella annulata’ on pl. 1, fig. 1, probably because of the many culmina in the pseudoabdomen in their specimen.

Longtanella laxiflexus Xiao & Suzuki sp. nov. (Figs 3G, S4Q–U) 2004 Pseudoalbaillella elegans Ishiga & Imoto; Hori: pl. 4, figs 43, 44. 2004 Pseudoalbaillella chilensis Ling & Forsythe; Yao et al.: pl. 1, fig. 5. 2006 Pseudoalbaillella chilensis Ling & Forsythe; Shimakawa & Yao: pl. 1, fig. 5. 2012 Pseudoalbaillella elegans Ishiga & Imoto; Wang et al.: 44, pl. 11, fig. 10. Diagnosis. Longtanella with weakly winding pseudoabdomen. Derivation of name. The species name ‘laxiflexus’, derived from the Latin adjective ‘laxus’(-a, -um) + masculine noun flexus (-us, m), meaning ‘weak winding’. Material. Holotype, Fig. S4U, specimen ST1-2_i011; paratype, Fig. S4Q, specimen 13ST9-1_083. Five specimens were examined by SEM and all of them are illustrated.

Measurements (μm). Height of shell 265–434 (mean 321); height of AC 50–102 (mean 75); width of the basal AC 33–60 (mean 47); height of PPA 27–69 (mean 40); height of PX 34–75 (mean 48); width of PX 60–82 (mean 69); height of PA 119–192 (mean 158); based on five specimens. Occurrences. Middle Kungurian–Roadian; Bancheng Formation of South China and the Chichibu composite belts of Japan. Description. Shell smooth, consisting of apical cone, rhombic pseudothorax, pre-pseudoabdomen and winding pseudoabdomen. AC conical and straight, about 1.5 times longer than PX. PX rhombic with no wings or wing-like rudiments. PPA has no obvious constriction, so PA is unable to be separated from PPA in the most cases with a few exceptions (Fig. S4U). PPA forms an annular ridge. PA long, constant in width without undulation, distinctly winding and forming an ‘S’ shape; the concave of posterior waist is on the ventral side. BS extends obliquely downwards. Remarks. This new species has been confused with Curvalbaillella chilensis and Curvalbaillella elegans (Hori 2004; Yao et al. 2004; Shimakawa & Yao 2006; Wang et al. 2012). The new species is similar to C. elegans in the long and smooth pseudoabdomen without undulation, but is distinguished from the former by the longer pseudothorax and winding pseudoabdomen. The winding pseudoabdomen in the new species also resembles that in C. chilensis. Referred to morphological variations in Curvalbaillella illustrated in Kuwahara (1992, pl. 1), any Curvalbaillella specimens curve once only and possess distinctive wings on the pseudothorax. It differs from Longtanella? sp. 3 (Fig. S4O, P) in the smooth pseudoabdomen.

Longtanella follicucullinoides Xiao & Suzuki sp. nov. (Fig. S5A–G) 1989 Pseudoalbaillella simplex Ishiga & Imoto; Isozaki & Tamura, figs. 4.11, 4.12. 2005 Pseudoalbaillella sp. cf. P. s i m p l e x Ishiga & Imoto; Kametaka et al.: figs 4.8–4.14. Diagnosis. Longtanella with long conical apical cone which smoothly continues from the apex to the slightly inflated pseudothorax and straight pseudoabdomen. Derivation of name. Similar to the genus Follicucullus. Material. Holotype, Fig. S5F, specimen ST1-2_i007; paratype, Fig. S5G, ST1-2_i022. Seven specimens were examined by SEM and all of them are illustrated. Measurements (μm). Height of shell 185–329 (mean 216); height of AC 81–102 (mean 91); width of the

37 basal AC 45–60 (mean 49); height of PPA 23–38 (mean 32); height of PX 43–55 (mean 50); width of PX 67–82 (mean 74); height of PA 18–149 (mean 51); based on seven specimens. Occurrences. Middle Kungurian–Roadian; Bancheng Formation of South China; Nagato Tectonic zone and North Kitakami Terrane in Japan. Description. Shell gracile and straight, consisting of a long apical cone, slightly inflated pseudothorax and cylindrical pseudoabdomen. Pre-pseudoabdomen generally absent. AC sturdy, conical without undulation, sometimes slightly curving to the ventral side. PX inflated with no wings, usually 1/2 of the height of AC. PPA, if present, cylindrical, similar to the width of PX. PA long, constant in width without undulation, about 1/2 of the height of shell. There develops a constriction between PPA and PA (see Fig. S5F), and the shell usually divided into two parts along this constriction. Remarks. The more complete specimen in Fig. S5F shows that this morphospecies is gracile and weak in the boundary of the PPA and PA. This species may sometimes be confused with poorly-preserved specimens of Curvalbaillella elongatus (Ishiga & Imoto, 1980), Parafollicucullus eurasiaticus (Kozur & Mostler, 1989), Parafollicucullus lanceolatus (Ishiga and Imoto in Ishiga et al., 1982), Parafollicucullus lomentarius, Parafollicucullus longicornis (Ishiga & Imoto, 1980) and Parafollicucullus simplex (Ishiga & Imoto, 1980). The new species is easily distinguishable from all these species in lacking wings even if specimens are poorly preserved. In the context of no wings in the new species, poorly-preserved Follicucullus is also similar to this species, but the new species is also easily distinguishable from the latter by its asymmetric shape which makes it hard to distinguish dorsal or ventral side.

Reference list:

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Supplement 6. The character data set used for phylogenetic tree of Follicucullidae.

Supplement 6-1. List of species and the codes of each character for each species. The codes and states of the 49 characters are explained in Supplement 6-2.

Abbreviations: Ho., Holdsworthella; Ha., Haplodiacanthus; Ps., Pseudoalbaillella; Pa., Parafollicucullus; L., Longtanella; K., Kitoconus; Ca., Cariver; Cu., Curvalbaillella; F., Follicucullus; I., Ishigaconus.

Supplement 6-2. The codes and states of the 49 characters.

Supplement 7. Meta dataset for CA. 1: occurrence, 0: absent.

Information on Zoobank records

Xiao, Yifan, Noritoshi Suzuki, Tsuyoshi Ito & Weihong He. 2021. New Permian radiolarians from ea st Asia and the quantitative reconstruction of their evolutionary and ecological significances. LSIDurn:lsid: zoobank.org:pub:8259B828-3B3B-4786-8A89-C31620AD8F4E

Parafollicucullinoides Xiao, Ito & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:99646A94-3503-4913-A04F-08A3D063DDD3

Longtanella edamame Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:4A402824-067A-4369-86DB-2AB658648B42

Longtanella follicucullinoides Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:0F51E98F-65E9-4AA4-B86A-41FEE8823B53

Longtanella jingyi Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:3F16D163-8180-459D-8EF9-996BC9850094

Longtanella kushidango Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:83CEAD3E-BCC5-4EAC-8775-B10BC37B6944

Longtanella lanceoliformis Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:20D2BBF5-4776-4B3D-9A95-4B40D586D502

Longtanella laxiflexus Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:62026843-0F8E-48D5-AD74-EB88EA804420

Longtanella tokkuriformis Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:A5B64F44-BF85-481E-8027-C024B51CC9F9

Longtanella turrita Xiao & Suzuki in Xiao, Suzuki, Ito & He, 2021 LSIDurn:lsid: zoobank.org:act:D23E56D5-F9AA-4971-81B5-28D6DB879831