Introduction
Permian crinoids are relatively rare compared with crinoids from earlier parts of the Paleozoic (Table 1). The most diverse and abundant articulated Permian crinoid crowns and cups were reported from Timor (Wanner, Reference Wanner1916, Reference Wanner1924, Reference Wanner1937, among others; Webster, Reference Webster2012a; Webster and Donovan, Reference Webster and Donovan2012). Although the stratigraphic position was uncertain, it was considered to be Artinskian by Charlton et al. (Reference Charlton2002). In addition, early Permian crinoid faunas are known from Russia (Yakovlev, Reference Yakovlev1926, Reference Yakovlev1927, Reference Yakovlev1930; Yakovlev and Ivanov, Reference Yakovlev and Ivanov1956), the United States (the midcontinent: Moore and Plummer, Reference Moore and Plummer1940; Pabian and Strimple, Reference Pabian and Strimple1974; southern Nevada: Lane and Webster, Reference Lane and Webster1966; Webster and Lane, Reference Webster and Lane1967, Reference Webster and Lane2007), Australia (eastern: Willink, Reference Willink1978, Reference Willink1979a, Reference Willinkb, 1980; Webster and Jell, Reference Webster and Jell1999b; Western Australia: Webster, Reference Webster1987, Reference Webster1990; Webster and Jell, Reference Webster and Jell1992; Teichert and Webster, Reference Teichert and Webster1993; Tasmania: Sieverts-Doreck, Reference Sieverts-Doreck1942), Oman (Webster and Sevastopulo, Reference Webster and Sevastopulo2007; Webster et al., Reference Webster, Angiolini and Tintori2009b), Thailand (Webster and Jell, Reference Webster and Jell1993), British Columbia (Webster et al., Reference Webster, Haggart, Saxifrage, Saxifrage, Gronau and Douglas2009a), Greece (Webster, Reference Webster2012b), Bolivia (Branisa, Reference Branisa1965; Strimple and Moore, Reference Strimple and Moore1971; Burke and Pabian, Reference Burke and Pabian1978), Mexico (Strimple, Reference Strimple1971), China (Tien, Reference Tien1926; Chen and Yao, Reference Chen and Yao1993), and Spitsbergen/Norway (Gorzelak et al., Reference Gorzelak, Błażejowski, Uchman and Hanken2013). Crinoid faunas are also known from the middle Permian of Tunisia (Valette, Reference Valette1934; Lane, Reference Lane1979), Europe (Gregorio, Reference Gregorio1930; Ramovš and Sieverts-Doreck, Reference Ramovš and Sieverts-Doreck1968; Strimple and Sevastopulo, Reference Strimple and Sevastopulo1982), and Pakistan (Waagen, Reference Waagen1887), as well as from upper Permian strata of Europe (e.g., Donovan et al., Reference Donovan, Hollingworth and Veltkamp1986; Reich, Reference Reich2007). Among them, the Timor, southern Nevada, Western Australia, and southern Ural Mountains faunas are the four main early Permian faunas representing more than 40 genera. The only Permian crinoids reported previously from China are Separocrinus discoides Chen and Yao, Reference Chen and Yao1993 (Webster et al., Reference Webster, Waters and Chen2009c) from the early Permian Sakmarian to Artinskian of Yunnan and one possible crinoid fauna from North China (southern Hebei Province), Taiyuan Series (exact age uncertain), including Delocrinus, Mathericrinus, and Sinocrinus (Tien, Reference Tien1926).
Table 1 Most significant previous localities, ages, and formations of Permian crinoid publications.
The fossils described herein were collected from an outcrop at Tianxi Village, Shanying Town, southern Anyang City, Henan Province (36.24730°N, 114.65929°E) (Fig. 1), approximately 120 km south of the Lincheng Coal Field Section of southern Hebei Province studied by Tien (Reference Tien1926). The present study adds to our knowledge of the diversity of crinoids during the earliest Permian in the North China Block.
Figure 1 Map showing the section locality at Tianxi Village, Shanying Town, southern Anyang City of Henan Province.
Geologic setting and stratigraphy
Lower Paleozoic strata of the eastern part of the North China Block (or Sino-Korean Platform) are composed dominantly of shallow marine carbonates (Feng et al., Reference Feng, Wang and Zhang1990). The block was largely uplifted during Ordovician time. A remarkable sedimentary gap (parallel unconformably) is present at the top of the Cambrian or Ordovician. The dividing line is from Xiaxian to Dengfeng. In the south of the dividing line, the gap is at the top of the Cambrian; in the north of the line, the sedimentary gap lies at the top of the Ordovician. No Upper Ordovician, Silurian, Devonian, or Mississippian deposits occur. Unconformably overlying the Cambrian or Ordovician limestones is the Pennsylvanian Benchi Formation, which is dominated by terrestrial deposits (e.g., Hou et al., Reference Hou, Xu, Yang, Wang, Liu, Zhao, Rui, Wang, Zang and He1987; Lü and Chen, Reference Lü and Chen2014). The Benchi Formation is composed dominantly of siltstone and contains abundant vascular plant fossils and bauxite. It is biostratigraphically dated by the conodont Idiognathodus delicates–I. podolskensis Assemblage Zone as the middle to upper Moscovian, Pennsylvanian. Strata above the Pennsylvanian Benchi Formation are the Permian Taiyuan, Shanxi, Shihezi, and Shiqianfeng formations in ascending order (Hou et al., Reference Hou, Xu, Yang, Wang, Liu, Zhao, Rui, Wang, Zang and He1987; Wu et al., Reference Wu, Zhang, Zhao, Jin and Liao1987). The base of the Taiyuan Formation is currently defined by the first occurrence of the limestones sharply differing from the Benchi Formation below. The Taiyuan Formation is composed of tens to hundreds of meters of strata and is characterized by limestones interbedded with siltstones and mudstones with rare coal beds indicating alternating of marine and terrestrial facies (Bureau of Geology and Mineral Resources of Henan Province, 1990). Conodont dating suggests that it is earliest Permian, Asselian, in age (Wang et al., Reference Wang, Yan, Jiang, Xi and Feng1987; Lang and Wang, Reference Lang and Wang2007).
Anyang is located in the central part of the North China Block. Coal, iron, limestone, and bauxite of Pennsylvanian and Permian age are important economic resources of this region. Upper Paleozoic outcrops along the Tianxi section are discontinuous. The lower part of the Taiyuan Formation is covered by roads and houses and partly measured as 15 m in thickness (Fig. 2). Brown siltstones yielding fossils of vascular plants, especially ferns, and insect fossils are indicators of a nonmarine environment. The marine fossil-bearing unit consists of thin-bedded calcareous siltstones and thick-bedded bioclastic limestones and is the Dajian Member, middle part of the Taiyuan Formation. Above the Dajian Member are sandstones of the Mojie Member (Wang et al., Reference Wang, Yan, Jiang, Xi and Feng1987). Thus, the Dajian Member biologically and lithologically indicates a dramatic transgressional interval yielding a shallow marine fauna.
Figure 2 Stratigraphic log of the section at Tianxi Village, Shanying Town, southern Anyang City of Henan Province.
Well-preserved crinoid fossils are concentrated within the calcareous siltstones in an interval approximately 1.5 m thick. Co-occurring macrofossils are the trilobite Ditomopyge sp.; brachiopods Choristites sp., Martinia spp., Eomarginifera sp., Punctospirifer sp., Neospirifer sp., Cleiothyridina sp., and Echinoconchus sp.; gastropod Pseudozygopleura sp.; rugose corals Lophocarinophyllum, Yakovleviella, and Tachylasma; and bryozoan Streblotrypa. These are common elements of the Taiyuan Formation (Wang et al., Reference Wang, Yan, Jiang, Xi and Feng1987; Zhao, Reference Zhao1987; Zhao and Liang, Reference Zhao and Liang1989). Sponge spicules are less common and preserved as fragments. The index conodont fossil, Streptognathodus cristellaris Chernykh and Reshetkova, Reference Chernykh and Reshetkova1987, is from the thick-bedded bioclastic limestones, approximately 20 cm above the argillaceous limestone. S. cristellaris is representative of the second conodont zone of the Permian Asselian in Russia (Chernykh, Reference Chernykh2006).
Faunal analysis
The Henan crinoid fauna is diverse and contains several new species. It contains representatives of each of the major groups of the crinoids known in the Paleozoic: Cladida (including Flexibilia), Disparida, and Camerata. Cladid crinoids (~59%) dominate the fauna as they do in nearly all post-Mississippian faunas. The fauna is divisible into two groups: cosmopolitans of the equatorial belt during the Permian and taxa known from the Americas. No endemic genera are present. Platycrinites, Synbathocrinus, and sagenocrinitids are all cosmopolitan in the equatorial belt in the Permian (Webster, Reference Webster2003). All genera in the Henan fauna are recognized in North America in the Carboniferous. The disparid Synbathocrinus and the eucladid Apographiocrinus are also known from early Permian (Artinskian) strata of North America as well as in Timor and Western Australia. Celonocrinus was reported from the middle Wolfcampian of North America by Lane and Webster (Reference Lane and Webster1966). In addition, Erisocrinus was reported from middle Permian strata of Russia by Yakovlev and Ivanov (Reference Yakovlev and Ivanov1956), and Artichthyocrinus was described from the early Permian (possibly within the Asselian to Artinskian) of Greece by Webster (Reference Webster2012b).
The Henan crinoid fauna is considered to have the greatest affinity with North American faunas. Ulocrinus and Neoprotencrinus have not previously been reported from Permian strata, and their ranges are extended upward into the Permian. The paleogeographic range of these genera is extended to include China. The Henan crinoid fauna is from the early Asselian; it may be helpful in understanding migration routes of fossil taxa and will be of fundamental value in helping paleontologists undertake paleogeographic reconstructions of the Permian globe.
Materials
Repository and institutional abbreviation
All specimens are deposited in the Nanjing Institute of Geology of Paleontology (NIGP), Chinese Academy of Sciences (NIGPAS).
Systematic paleontology
Crinoid morphologic terms follow Moore and Teichert (Reference Moore and Teichert1978) with anal and radial facet modifications by Webster and Maples (Reference Webster and Maples2006, Reference Webster and Maples2008). Measurement terminology follows Webster and Jell (Reference Webster and Jell1999a). The classification follows Simms and Sevastopulo (Reference Simms and Sevastopulo1993), Wright et al. (Reference Wright, Ausich, Cole, Peter and Rhenberg2017), and Wright (Reference Wright2017b).
Class Crinoidea Miller, Reference Miller1821
Subclass Pentacrinoidea Jaekel, Reference Jaekel1894
Infraclass Inadunata Wachsmuth and Springer, Reference Wachsmuth and Springer1885
Parvclass Cladida Moore and Laudon, Reference Moore and Laudon1943
Magnorder Eucladida Wright, Reference Wright2017a
Clade Articuliformes Wright, Reference Wright2017b
Cyathoformes incertae sedis
Family Stellarocrinidae, Strimple, Reference Strimple1961
Genus Celonocrinus Lane and Webster, Reference Lane and Webster1966
Type species
Celonocrinus expansus Lane and Webster, Reference Lane and Webster1966.
Celonocrinus cf. C. expansus Lane and Webster, Reference Lane and Webster1966
Figure 3 Crinoids from the Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block. (1–3) Celonocrinus cf. C. expansus Lane and Webster, Reference Lane and Webster1966, NIGP 166462 and NIGP 166463; (1, 2) basal views; (3) oral view. (4) Neoprotencrinus anyangensis n. sp., holotype, NIGP 166464, lateral view. Scale bar=1 cm.
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Crown spreading out and curling back in over the top; dorsal cup faint low and broad shape; cup plates smooth. Basals pentagonal in outline, curved longitudinally. Radials gently convex, short, much wider than long, with long basal-radial sutures that meet at a high angle; radial facet plenary, in contact with first primibrachial in the middle and first secundibrachials at lateral edges.
Arms branch isotomously twice, adjacent branches not in lateral contact proximally; first primibrachial small, short, triangular, and axillary in all rays, failing to extend fully across radials; first secundibrachials quadrangular, biserial, with angulated edges, in contact with the radial at lateral edges; higher branches on about secundibrachial eight; all brachials are broad, with flat outer surface. Column transversely round with crenulae.
Measurements for plates of the specimens are given in Table 2.
Table 2 Measurements for Celonocrinus cf. C. expansus specimens. Units=mm.
Materials
Figured specimens NIGP 166462 and NIGP 166463.
Remarks
The two cups are not well preserved; NIGP 166462 (Fig. 3.1) retains one relatively complete radial, one basal, and 12 arm fragments, but only two fragmentized arms and radials of NIGP 166463 are preserved (Fig. 3.2, 3.3). Although not complete, the pentagonal basals, wide and short radials, radial facet in contact with both first primibrachial and first secundibrachials, wide and flat arms and flat brachials, and branching mode are very suggestive of Celonocrinus expansus.
Superfamily Erisocrinacea Wachsmuth and Springer, Reference Wachsmuth and Springer1886
Family Erisocrinidae Wachsmuth and Springer, Reference Wachsmuth and Springer1886
Genus Neoprotencrinus Knapp, Reference Knapp1969
Type species
Paradelocrinus subplanus Moore and Plummer, Reference Moore and Plummer1940.
Neoprotencrinus anyangensis new species
Figure 3.4
Holotype
NIGP 166464.
Diagnosis
Neoprotencrinus with radials not in basal circlet, primibrachials and six uniserial secundibrachials, and slightly impressed sutures.
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Crown elongate cylindrical, length 85 mm; cup low bowl shape, length 6.5 mm, width 27 mm; basal concavity very shallow to flat; plates smooth, sutures depressed. Basals not visible in side view; radials large, length 6.5 mm, width 14 mm, width commonly about twice the length. Proximal tips of radials not in the basal circlet; radial facets plenary. No anal plates visible with arms attached. Arms 10, moderately long, closely appressed; branching isotomously on first primibrachials; first primibrachials slightly spinose; primibrachials and six uniserial secundibrachials.
Etymology
Named after Anyang city, Henan Province, which hosts the type locality.
Material
Holotype: NIGP 166464.
Remarks
One flattened specimen is assigned to N. anyangensis. Only one side of holotype NIGP 166464 (Fig. 3.4) is preserved and is slightly crushed without anal plates. Large first secundibrachials are followed by five quadrate uniserial secundibrachials with the sixth uniserial secundibrachials with an angular distal suture for the reception of the first wedge-shaped brachial. Thereafter, the brachials are biserial.
Webster and Kues (Reference Webster and Kues2006) moved Neoprotencrinus from the Protencrinidae, transferring it to the Erisocrinidae because of its biserial arms. Neoprotencrinus are reported from the Pennsylvanian of the United States, including one species, N. brachiatus Moore and Plummer, Reference Moore and Plummer1940, with the arms preserved. The differences in shape of the radials and primibrachials along with the difference in the number of nonbiserial secundibrachials are the justification for the species. N. anyangensis is distinguished from N. brachiatus by the nonbiserial secundibrachials, from N. gutschicki Webster and Kues, Reference Webster and Kues2006 by more impressed sutures, and from Neoprotencrinus cranei Strimple, Reference Strimple1949, N. regulates Strimple, Reference Strimple1949, N. disculus Strimple, Reference Strimple1949, and N. subplanus rockensis Moore and Plummer, Reference Moore and Plummer1940 by proximal tips of radials not in the basal circlet.
Family Protencrinidae Knapp, Reference Knapp1969
Genus Protencrinus Jaekel, Reference Jaekel1918
Type species
P. moscoviensis Jaekel, Reference Jaekel1918.
Protencrinus baliensis Webster, Reference Webster2012b
Figure 4.1–4.3
Figure 4 Crinoids from the Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block. (1–3) Protencrinus baliensis Webster, Reference Webster2012b, NIGP 166465, basal, oral, and lateral posterior views; (4–6) Apographiocrinus sp., NIGP 166466, basal, oral, and lateral posterior views; (7) gen. indet. sp. indet., NIGP 166467, lateral view; (8) family indet. gen. indet. sp. indet., NIGP 166468, lateral view; (9) Anal sac indeterminate, NIGP 166469, lateral view; (10–12) Sinocrinus sheareri Strimple and Watkins, Reference Strimple and Watkins1969, NIGP 166471, basal, oral, and lateral posterior views; (13–19) platycrinitid gen. indet. sp. indet., NIGP 166477–166483; (13, 14) facet views of columnals; (15–17) radial plates, basal views; (18, 19) basal circlets, basal views. Scale bar=1 cm.
Occurrences
Early Permian, series undesignated, Crete, Greece; Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Cup flat bowl shape (Fig. 4.3), medium-sized basal concavity, pentagonal in oral outline, smooth plate sculpturing, length 5 mm, width 23 mm. Infrabasals five, pentagonal, truncated distally in contact with radials, equal size, longer than wide, length 5 mm, width 4 mm; infrabasal circlet confined to basal concavity, down-flaring. Basals five, small quadrangular, not in lateral contact (Fig. 4.1), length 3 mm, width 3 mm, proximal tips in basal concavity, distal tips up-flaring with tips barely visible in lateral view. Radials five, hexagonal, much wider than long, length 5 mm, width 14 mm, gently convex transversely, moderately convex longitudinally, proximal tips in basal concavity, distally up-flaring form lateral walls; radial facet plenary (Fig. 4.2), deep ligament pit; narrow outer marginal area; deep central pit and intermuscular furrow; muscle fields shallowly concave; narrow lateral ridges. Anal notch not visible in lateral view with a narrow distinct V-shaped groove on the internal edge (Fig. 4.3). Column facet circular, 4 mm in diameter, impressed in infrabasal circlet.
Material
Figured specimen NIGP 166465.
Remarks
One well-preserved cup is assigned to P. baliensis Webster, 2012. It has a little deeper basal concavity than the specimens illustrated by Webster (Reference Webster2012b). The stratigraphic range of Protencrinus is Pennsylvanian to early Permian, and the paleogeographic range is North America, Russia, and Crete and, herein, is extended to North China.
Superfamily Apographiocrinoidea Moore and Laudon, Reference Moore and Laudon1943
Family Apographiocrinidae Moore and Laudon, Reference Moore and Laudon1943
Genus Apographiocrinus Moore and Plummer, Reference Moore and Plummer1940
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Cup low bowl shape (Fig. 4.6), length 6.6 mm, width 22.2 mm; basal cavity deeply concave in middle of cup; plates of cup slightly bulbous with fine nodose sculpture. Infrabasals not preserved, leaving small five-pointed star at bottom of basal concavity (Fig. 4.4). Proximal part of basals in basal plane, interbasal sutures faint. Radials five, length 5.5 mm, width 11.5 mm; transversely convex giving slightly scalloped appearance to summit of radials in dorsal view; radial articular facets thick, plenary, not well preserved in B, C, and D rays; ligament pits well defined, transverse ridge prominent with fine denticles; narrow, deep lateral furrows in E and A rays (Fig. 4.5). Single anal plate in cup between posterior radials, extending above radial summit (Fig. 4.6).
Material
Figured specimen NIGP 166466.
Remarks
The specimen is assigned to the Apographiocrinus on the basis of low bowl shape, small basal concavity, slightly bulbous plates, and single anal plate, but not preserved infrabasals, crushed plate surfaces, and faint sutures cannot be identified to a species. Judging from the phylogenetic position and the developmental biology of living crinoids, the single anal plate is almost certainly the radianal (Wright, Reference Wright2015).
Family Pachylocrinidae Kirk, Reference Kirk1942
Genus indeterminate species indeterminate
Figure 4.7
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Crown slender, cylindrical; cup slightly crushed, truncate bowl shape. Arms uniserial, faintly pinnulate; arm branching isotomously on primibrachial 1 and higher on secundibrachials 5 or 6; brachials narrow and long, rectangular to cuneate. Crushed column long.
Material
Figured specimen NIGP 166467.
Remarks
This specimen is assigned to Pachylocrinidae for truncate bowl shape, uniserial arms, and branching isotomously twice, but it is too incomplete to assign to a genus.
Family indeterminate
Genus indeterminate species indeterminate
Figure 4.8
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
One poorly preserved slender elongate cylindrical cladid crown with a low conical-shaped cup (Fig. 4.8). Four uniserial arms of two rays in enclosed lateral contact, branching on primibrachials 1; brachials very wide, rectangular to cuneate, ornament coarse nodes.
Material
Figured specimen NIGP 166468.
Remarks
This specimen is heavily weathered and cannot be identified to a genus.
Anal sac indeterminate
Figure 4.9
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Material
Figured specimen NIGP 166469.
Remarks
A partial anal sac consisting of three columns of plates moderately well preserved is not identifiable (Fig. 4.9). Plates are hexagonal and bulbous and interlock laterally with adjacent plates; distinctive one to three variable lengths along the lateral edges of some plates.
Superfamily Cromyocrinoidea Bather, Reference Bather1890
Family Cromyocrinidae Bather, Reference Bather1890
Genus Ulocrinus Miller and Gurley, Reference Miller and Gurley1890
Type species
U. buttsi Miller and Gurley, Reference Miller and Gurley1890.
Ulocrinus qiaoi new species
Figure 5 Crinoids from the Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block. (1–4) Ulocrinus qiaoi n. sp., holotype, NIGP 166470, lateral view, B ray view, anal view, and basal view; (5, 6) Erisocrinus cf. E. longwelli Lane and Webster, Reference Lane and Webster1966, NIGP 166472, lateral view; (7, 8) Artichthyocrinus limani n. sp., holotype and paratype, NIGP 166473 and NIGP 166474, basal view of holotype and paratype; (9–11) Synbathocrinus campanulatus Wanner, Reference Wanner1916, NIGP 166475, basal, oral, and lateral posterior views; (12–15) Synbathocrinus chenae n. sp., NIGP 166476, A ray view, anal view, basal view, and oral view. Scale bar=1 cm.
Holotype
NIGP 166470.
Diagnosis
Distinguished by the thick plates, pronounced irregular protrusions on plates, and three anals in cup.
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Cup high conical with convex base (Fig. 5.1, 5.2), length 20 mm, width 42.7 mm; plate sculpture irregular pronounced bulbous protrusions, impressed sutures; three cup circlets all visible in side view. Infrabasals five, up-flared strongly, length 6 mm, width 7 mm. Basals five, unequal size, hexagonal, large, length 12 mm, width 12.2 mm; basals larger than infrabasals. Radials five, pentagonal, unequal size, much wider than long, length 9.4 mm, width 15.5 mm; articular facets plenary, subhorizontal broad, transverse ridge marked by fine crenulations (Fig. 5.1). Three anal plates in cup, large primanal (equivalent to the radianal) obliquely between two basals, mostly to left of C radial, not in contact with D radial, supporting secundanal (equivalent to the anal X) and tertanal above (Fig. 5.3). Arms transversely well rounded, branching isotomously on first primibrachials; three thick first primibrachials (Fig. 5.2), length 6 mm, width 13 mm; secundibrachials cuneate in proximal portions, length 4 mm, width 6.8; distal part of arms not preserved. Column circular, 6 mm in diameter (Fig. 5.4).
Etymology
The specific name is in honor of amateur collector Mr. Qiao Liman, Anyang, for help with fieldwork.
Material
Holotype: NIGP 166470, only known specimen of this species.
Remarks
The partial crown of U. qiaoi n. sp. is crushed flat. It differs from the most similar U. neverovoensis Mirantsev and Rozhnov, Reference Mirantsev and Rozhnov2011 by having thicker plates and more impressed sutures. The new species also differs from U. grishini Mirantsev and Rozhnov, Reference Mirantsev and Rozhnov2011 in the less elongate cup and relatively long infrabasal circlet and differs from U. karchevskyi Mirantsev and Rozhnov, Reference Mirantsev and Rozhnov2011 in the strongly up-flared infrabasal circlet. It differs from all North American congeners in the presence of surface ornamentation.
Superfamily Erisocrinacea Wachsmuth and Springer, Reference Wachsmuth and Springer1886
Family Erisocrinidae Wachsmuth and Springer, Reference Wachsmuth and Springer1886
Genus Sinocrinus Tien, Reference Tien1926
Type species
Sinocrinus granulatus (Wanner, Reference Wanner1924).
Remarks
Note Sinocrinus was erected based on Sinocrinus microgranulosus Tien, Reference Tien1926, which is a junior subjective synonym of Sinocrinus granulatus Wanner, Reference Wanner1924 (cf. Sheffield, Reference Sheffield2015).
Sinocrinus sheareri Strimple and Watkins, Reference Strimple and Watkins1969
1969 Sinocrinus sheareri Reference Strimple and WatkinsStrimple and Watkins, p. 182, pl. 39, figs. 8–11.
1976 Sinocrinus asymmetricus; Reference StrimpleStrimple, p. 636, fig. 3A–B.
1977 Sinocrinus sheareri; Reference WebsterWebster, p. 156.
1986 Sinocrinus asymmetricus; Reference WebsterWebster, p. 284.
2015 Sinocrinus sheareri; Reference SheffieldSheffield, p. 468, figs. 7, 9A–C.
Occurrences
Pennsylvanian, Atokan, Big Saline Formation, Soldiers Hole Member, United States, Texas; Pennsylvanian, Moscovian, Bashkirian? Spain; Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Dorsal cup flat, asymmetrically bowl-shaped (Fig. 4.12), length 6 mm, width 21.4 mm; plate ornamentation dense nodes, impressed sutures. Infrabasals five, fuse into a single plate, not in contact with radials; infrabasal circlet subhorizontal, positioned slightly above basal plane, width 8.3 mm, not as high as radial circlet; only basal and radial plates visible in side view. Basals five, tumid, unequal size with the smallest one in the posterior, the proximal portion of basals in the basal plane and the distal portion sharply upward to form part of lateral walls, length 6 mm, width 6 mm; interbasal sutures short. Radials tumid, equal size, the proximal tips of radials in basal plane of cup but not entering basal concavity, length 5.2 mm, width 11.8 mm; radial facets plenary (Fig. 4.11), outer ligament furrow poorly developed, sharply defined ligament pit, wide transverse ridges bearing fine denticles, well-developed oblique furrows, broad muscle scars, and small V-shaped intermuscular notch. No anal plate in external view, small internal anal notch on distal extremities of adjacent C and D radials in Figure 4.12. Column large, circular, with strongly crenulated cicatrix.
Material
Figured specimen NIGP 166471.
Remarks
Sheffield (Reference Sheffield2015) refined Sinocrinus Tien, Reference Tien1926 using principal component analysis and neighborhood cluster analysis and extended the temporal span of the genus from the late Carboniferous (Moscovian) through the early Permian (Sakmarion). Sinocrinus asymmetricus Strimple, Reference Strimple1976 was rejected as the junior synonym of S. sheareri Strimple and Watkins, Reference Strimple and Watkins1969 because of their qualitative and quantitative similarities with one another.
Sinocrinus is mainly reported from the Houkou Limestone, Taiyuan Series of northern China. This specimen is distinguished from S. microgranulosus Tien, Reference Tien1926, S. microgranulosus pentalobus Tien, Reference Tien1926, and S. linchenensis Tien, Reference Tien1926 in having a proportionately lower, asymmetric dorsal cup, a proportionately larger column, lesser part of basals in cup walls, and short interbasal sutures. Three other forms are described by Tien (Reference Tien1926) as S. houkouensis, S. nodosus, and S. nodosus spinosus based on disarticulated plates.
Sinocrinus sheareri has been reported from the Pennsylvanian of United States and Spain. One well-preserved specimen is assigned to S. sheareri, extending the stratigraphic range of S. sheareri to early Permian and the paleogeographic range to include China.
Genus Erisocrinus Meek and Worthen, Reference Meek and Worthen1865.
Type species
E. typus Meek and Worthen, Reference Meek and Worthen1865.
Erisocrinus cf. E. longwelli Lane and Webster, Reference Lane and Webster1966
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Partial crown crushed flat, elongate cylinder (Fig. 5.5), length 52.7 mm; dorsal cup large, medium bowl-shaped, length 13 mm, width 33 mm; plate ornament dense nodes. Infrabasals not preserved. Basals large, length 7.8 mm, width 11 mm, two preserved in Figure 5.5 and two visible in Figure 5.6. Radials large, length 8 mm, width 14 mm, swollen slightly in middle; radial facets plenary; no discernible notch for an anal plate between any of the radials or above the cup view from the well-preserved side. Six total arms on one side, branching isotomously on large first primibrachials, biserial, slightly convex, closely apposed; first primibrachial large, quadrangular, more than twice as wide as long, length 6 mm, width 14 mm, strongly spinose, supporting a large quadrangular first secundibrachial on each shoulder; first secundibrachial quadrangular, length 3.8 mm, width 7 mm; secundibrachial 2 smaller than first secundibrachial, biserial just above the first secundibrachial, brachials smaller distally. Column circular.
Material
Figured specimen NIGP 166472.
Remarks
The specimen most closely resembles E. longwelli in size, but the infrabasals and the other side are crushed. Thus, it is assigned to Erisocrinus cf. E. longwelli.
Superorder Flexibilia Zittel, Reference Zittel1895
Order Sagenocrinida Springer, Reference Springer1913
Superfamily Sagenocrinoidea Roemer, Reference Roemer1854
Family Euryocrinidae Moore and Strimple, Reference Moore and Strimple1973
Genus Artichthyocrinus Wright, Reference Wright1923
Type species
Artichthyocrinus springeri Wright, Reference Wright1923.
Artichthyocrinus limani new species
Holotype
NIGP 166473.
Diagnosis
Artichthyocrinus with circular column impression covering infrabasals, basals, and most of radials. Circlet of plates beyond column impression includes interradials and anal plates.
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Crown bowl-shaped with a shallow depressed base, wider than long; cup discoid (Fig. 5.7, 5.8); broad shallow calyx formed by rigidly sutured plates; plates smooth, much wider than long, sutures flush. Infrabasal and basal circlets entirely concealed by proximal columnal. Radials five, much wider than long, widely out-flaring, lateral sutures fused in part; radial facets plenary, not exposed; one column of interradial plates in each interray, interradials and anal plates not easy differentiated, interradials in AB, BC, and AE relatively larger; smallest anal plate followed by additional small quadrangular or hexagonal plates in posterior interray. Arms branching on primibrachials 2; arms nearly flat outer surfaces; brachials fitted smoothly together laterally or interlocked. Column circular, large, impression surrounded by a flange. Measurements for plates of the specimens are given in Table 3.
Table 3 Measurements for Artichthyocrinus qiaoi n. sp. specimens. Units=mm.
Etymology
The specific name is after Mr. Qiao Liman, Anyang, the collector of the holotype.
Materials
Holotype: NIGP 166473; paratype: figured specimen NIGP 166474.
Remarks
Two specimens are assigned to A. limani n. sp. The crown of holotype, NIGP 166473 (Fig. 5.7), is slightly distorted, only two secundibrachials are preserved, and the distal parts of the arms are lost. Paratype NIGP 166474 (Fig. 5.8) is relatively smaller, only one secundibrachial in each ray, distal parts of the arms are lost, and the suture lines between these plates cannot be determined with certainty.
A. limani n. sp. differs from A. springeri Wright, Reference Wright1923 and A. koenigi Webster, Reference Webster2012b in having a column of interradials, more anal plates, and no basals extending beyond the circular column impression. The paleogeographic range of Artichthyocrinus extended to include China.
Parvclass Disparida Moore and Laudon, Reference Moore and Laudon1943
Superfamily Belemnocrinacea Miller, Reference Miller1883
Family Synbathocrinidae Miller, Reference Miller1889
Genus Synbathocrinus Phillips, Reference Phillips1836
Type species
Synbathocrinus conicus Phillips, Reference Phillips1836.
Synbathocrinus campanulatus Wanner, Reference Wanner1916
1916 Symbathocrinus campanulatus Reference WannerWanner, p. 16, pl. 96, figs 5, 6; text-figs 3, 4.
1924 Symbathocrinus campanulatus; Reference WannerWanner, p. 310, pl. 20, figs 23–25.
1942 Synbathocrinus campanulatus; Reference Moore and EwersMoore and Ewers, p. 105.
1943 Synbathocrinus campanulatus; Reference Bassler and MoodeyBassler and Moodey, p. 694.
1948 Synbathocrinus campanulatus; Reference BransonBranson, p. 215.
1973 Taidocrinus campanulatus (Wanner); Reference WebsterWebster, p. 252.
1973 Taidocrinus inflatus (Wanner); Webster, p. 252.
1987 Synbathocrinus campanulatus; Reference WebsterWebster, p. 114, Figs. 9A–L; 10A–D.
1993 Synbathocrinus campanulatus; Reference Teichert and WebsterTeichert and Webster, p. 4, pl. 65, figs. 7–10.
1993 Synbathocrinus campanulatus; Reference WebsterWebster, p. 114.
Occurrences
Early Permian, late Sakmarian, Callytharra Formation, Western Australia, Australia; Permian, Artinskian, Basleo, West Timor; Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
One well-preserved dorsal cup small, medium conical, truncate base with shallow basal concavity, length 4 mm, width 12.5 mm, perfect pentameral symmetry except very small anal plate articulated to mutual shoulders of C–D radial (Fig. 5.11); plate ornamentation nodose, flush sutures; basals and radials expand in width distally constant to slightly increasing rate of distal widening. Basals three, low, unequal, with small one in A–E interray, length 1.5 mm; the basal circlet visible in side view of cup, width 7 mm; ratio of basal circlet length/distal basal circlet width is 0.21, and basal plate circlet 37.5% of aboral cup length. Radials up-flaring, length 2.5 mm, distal-to-proximal radial plate width ratio about 1.79; radial articular facets plenary, subhorizontal, narrowing orally, triangular shape, with transverse ridge and ligament pit; anal notch obvious V-shape on distal extremities of adjacent C and D radials. Column circular.
Material
Figured specimen NIGP 166475.
Remarks
Webster (Reference Webster1987) pointed out that the degree of the anal notch of Synbathocrinus has varied throughout the geological range of the genus and is not a reliable specific or generic character and recommended the suppression of Taidocrinus as a subjective junior synonym of Synbathocrinus.
Synbathocrinus chenae new species
Holotype
NIGP 166476.
Diagnosis
Distinguished by low bowl-shaped cup, slender elongate anal plate, relatively long, wide transversely convex brachials.
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Crown elongate, slender (Fig. 5.12, 5.13), length 32 mm; cup small, broadly bowl-shaped, length 5.5 mm, width 19 mm; sutures between plates slightly impressed; plate sculpturing coarse nodes; basal circlet slightly impressed. Radials five, subquadrangular in outline, slightly wider than long, length 5.2 mm, width 6.8 mm, distal inclinate; narrow and deep concavity between radials and primibrachials; articular facets plenary. Anal plate slender, elongate, in notch on C and D radials, and nearly out of the cup (mostly above the cup). Arms five, long and slender, atomous, closely appressed when enclosed, only seven primibrachials preserved in each ray; first primibrachial approximately as wide as radial, quadrangular, gently convex, length 4.7 mm, width 6.8 mm; second primibrachial wider than long, length 3 mm, width 5.7 mm; brachials relatively long, wide, convex; longitudinal ridges present in middle of each brachial. Column circular.
Etymology
The specific name honors Mrs. Chen Guangjü, the holotype collector.
Material
Holotype: NIGP 166476, only known specimen of this species.
Remarks
The holotype NIGP 166476 of S. chenae n. sp. is elongate and slender, and the distal portions of the arms are not preserved (Fig. 5.12, 5.13). The basal of the cup is crushed and flattened, resulting in an elliptical column scar and small basal circlet (Fig. 5.14).
S. chenae n. sp. is assigned to the Synbathocrinidae on the basis of confinement of the elongate and slender crown, small bowl-shaped cup, wide and inclinate radial articular facets, five arms, and anal plate notching upper corners of C and D radials and nearly out of cup. It differs from most Synbathocrinus species in the low bowl-shaped cup and relatively long, wide, and convex brachials.
Subclass Camerata Wachsmuth and Springer, Reference Wachsmuth and Springer1885
Infraclass Eucamerata Cole, Reference Cole2017
Order Monobathrida Moore and Laudon, Reference Moore and Laudon1943
Suborder Glyptocrinina Moore, Reference Moore1952
Superfamily Platycrinitacea Austin and Austin, Reference Austin and Austin1842
Family Platycrinitidae Austin and Austin, Reference Austin and Austin1842
Genus indeterminate species indeterminate
Occurrence
Taiyuan Formation, Dajian Member, early Permian (Asselian), Anyang, Henan, North China Block.
Description
Several loose ossicles of unidentified platycrinitids, including two columnals (Fig. 4.13, 4.14), three radials (Fig. 4.15–4.17), and two basal circlets (Fig. 4.18, 4.19).
NIGP 166477–166478, two columnal nodals, much wider than long, bearing elliptical articula with latus smooth; angular divergence of upper and lower fulcra about 65° on opposite sides of the nodals in facet view.
NIGP 166479–166481, radial plates, subquadrangular in outline, being slightly wider than long; articulate facets rather narrow, less than half width; NIGP 166479 (length 10.2 mm, width 11.8 mm) and NIGP 166480 (length 10.2 mm, width 11.2 mm) quite smaller with a smoother surface than NIGP 166481 (length 18 mm, width 20.5 mm).
NIGP 166482 and 166483, two different basal circlets with perfect pentamerous symmetry; NIGP 166482, three smooth plates, width 30 mm; NIGP 166483, five plates with irregular nodes on the surface, width 21.2 mm.
Materials
Figured specimens NIGPA 166477–166483.
Remarks
Ausich and Kammer (Reference Ausich and Kammer2009) summarized the generic concepts of Platycrinitidae. The occurrence of thecal ossicles of Platycrinites in lower Permian rocks has been reported several times from Australia (Etheridge, Reference Etheridge1892; Marez Oyens, Reference Marez Oyens1940; Webster and Jell, Reference Webster and Jell1999b), Timor (Wanner, Reference Wanner1916), China (Tien, Reference Tien1926), Russia (Yakovlev and Ivanov, Reference Yakovlev and Ivanov1956), the United States (Webster and Lane, Reference Webster and Lane1967; Broadhead and Strimple, Reference Broadhead and Strimple1977), and Oman (Webster and Sevastopulo, Reference Webster and Sevastopulo2007).
The taxonomy of platycrinitids is, to a great extent, based on arm-branching patterns and tegmen plate arrangement. Such characters have no value in dealing with loose plates.
Acknowledgments
We thank the colleagues of the NIGPAS, Professors Z. Liao, W. Liao, J. Yuan, and F. Xia, respectively, for brachiopod, coral, trilobite, and bryozoan determinations and stratigraphic suggestions, as well as the postgraduate student S. Yu and the amateur collectors Mr. L. Qiao and Mrs. G. Chen for help with field collections. We are grateful to D.F. Wright and one anonymous reviewer, and the editors J. Jin and S. Zamora, for careful comments that greatly improved this manuscript. This research was supported by Strategic Priority Research Program (B) of the Chinese Academy of Sciences (No. XDB10010503), the National Natural Science Foundation of China (Nos. 41372022 and 41521061), and China Scholarship Council (File No. 201604910567).
