Records of the Western Australian Museum 19: 379-389 (1999). Some Late Cambrian molluscs from Liaoning Province, China Yu Wen' and Ellis L. Yochelson^ ‘ Nanjing Institute of Geology & Palaeontology, Academia Sinica, Nanjing, 210008, China Present address: Department of Earth & Planetary Sciences, Western Australian Museum, Francis Street, Perth, Western Australia, 6000, Australia ^Department of Paleobiology, National Museum of Natural History, Smithsonian institution, Washington, D.C. 20560 U.S.A. Abstract - Pilina liaoningensis sp. nov. is described from the Late Cambrian Wanwankou Member of the Fengshan Formation, Benxi, Liaoning, China. Muscle scars and other internal structures are preserved very well in the paratype; bifurcate bundle scars are described for the first time from the fossil Tergomya. This is the oldest known occurrence of Pilina. Three species of Late Cambrian gastropods, "Maclurites" ulrichi (Kobayashi), Matlwrella walcotti Kobayashi and Lytospira wanwankouensis sp. nov. are also described from the Late Cambrian of northeastern China. Lytospira has been previously reported in the Lower Ordovician to the Middle Silurian of North America and Europe. INTRODUCTION Ordovician in age. Kobayashi (1933) studied the The fossil Tergomya and Gastropoda described fauna, which included a great variety of molluscs and illustrated in this paper were collected in 1980 from the Wanwankou Dolomite of the Wanwankou by Xu Jun-tao and his colleagues of the Nanjing Series exposed in the Niuxintai Basin of Liaoning Institute of Geology and Palaeontology, Academia Province. In that work he described the following Sinica. They are from the Wanwankou Member of gastropods: Helicotoma wamoanensis Kobayashi, the Fengshan Formation exposed in two sections. Matherella xoalcotti Kobayashi, Clisospira The Chiushukou section is located at the Qiushugou niuhsintaiensis Kobayashi, Archinacella wanwanensis village of Niuxintai, northeastern Benxi County, Kobayashi, Proplina bridgei Kobayashi, P. ampla Kobayashi, P. (?) sp., Sceiu’lla sp. indet., Stenotheca (?) Liaoning Province, and the Doufangkou section is nianchurica Kobayashi, Scaevogyra ulrichi Kobayashi near the Yingzi village of Huolianzhai, in the northern part of Benxi. and S. naticaformis Kobayashi. Two decades later, The collection contains four species placed in four after restudying the strata of the Taizihe Valley, Wang and others revised the age of the Wanwankou genera: Pilina liaoningensis sp. nov.; "Maclurites" Limestone, Wanwankou Dolomite and Chiushukou ulrichi (Kobayashi); Matherella walcotti Kobayashi Shale, and assigned those three formations to the and Lytospira ivamoankouensis sp. nov. The first tw-o Late Cambrian Fengshan Formation (Wang et al., taxa are from the Doufangkou section; Matherella 1954; Lu, 1962; Lu et at., 1974; Chen et al., 1979, 1983; walcotti and Lytospira wanwankouensis are from the Chen and Teichert, 1983). Chiushukou section (Figure 1). Apparently among Tire Fengshan Formation in Liaoning Province is these fo,ssils, Pilina liaoningensis represents the oldest divided into three members, in ascending order: the known species of the genus; its excellently Yenzhou Member, the Wanwankou Member and preserved muscle scars and other internal structures the Chiushukou Member (Wang et al., 1954; Lu, are highly significant. 1962; Chen et al., 1979, 1983; Chen and Teichert, 1983). The Wanwankou Member is about 20 to 50 m GEOLOGICAL SETTING thick, and is noteworthy for its rich fauna of Cambrian sediments, especially Upper Cambrian, cephalopods, rostroconchs, gastropods, trilobites, are extensively distributed in Liaoning, Shandong brachiopods and conodonts. It is composed mainly and Shanxi Provinces and Inner Mongolia in North of stromatolitic limestone and stromatolitic China. The Upper Cambrian rocks are well dolomite, interbedded with intraformational developed and contain a diversity of fossils. limestone conglomerate (Chen and Teichert 1983). Kobayashi (1931) established the Wanwankou The molluscs described herein are generally well Series in Taizihe Valley, Liaoning Province, to preserved. Associated with them are the include the Wanwankou Limestone, Wanwankou cephalopods Sinoeretnoceras wanwanense Dolomite, Chiushukou Shale and Hsiapingkou (Kobayashi), S. taiziense Chen and Teichert and Dolomite. He considered the Series to be Early Wanwanoceras peculiare Kobayashi, and trilobites. 380 Yu Wen, E.L. Yochelson The fauna contains characteristic fossils of the Type locality Upper Cambrian which is widely distributed in the DoufangKou section exposed near Yingzi Upper Cambrian of Shandong, Anhui, and Zhejiang village, Huolinanzhai, Benxi Liaoning Province, Provinces (Lu, 1962; Chen et al., 1979; Chen and China (see Figure 1). Teichert, 1983; Chen et til., 1983). Illustrated specimens are deposited in the Nanjing Diagnosis Institute of Geology and Palaeontology (NIGP), Shell large, tryblidiform, elliptical in dorsal view. Academia Sinica. ProtcKonch symmetrically conical. Apex overhanging adapical margin. Surface ornamented with stout comarginal rugae, fine growth lines and radiating SYSTEMATIC PALAEONTOLOGY tlareads. Interior with eight metametric pairs of muscle scars, one pair of radular muscle scars, and other Class Tergomya Horny, 1965 internal structures on dorsal side of stcinkern. Order Tryblidiida Lemche, 1957 Description Superfamily Tryblidiacea Pilsbry, in Zittel- Shell large, elliptical in dorsal view. Protoconch Eastman, 1899 small, about 5 mm wide, symmetrically conical in Family Tr>’blidiidae Pilsbry, in Zittel-Eastman, 1899 dorsal view, protruding strongly forward, broadly rounded in the basal part, slowly and gradually Genus Piliiia Koken, in Koken and Perner, 1925 increasing in size away from the apex. Tip and external features of protoconch are not preserxed Pilina litiouiiitfcnsis sp. nov. (Figure 3c). Dorsum gently flattened, oblique Figures 2, 3a-i, 4a-e toward the abapical margin, with greatest shell Material Examined width just behind midlength. Abapical margin rounded; adapical margin narrowly rounded, Hololi/pe though not known in detail. Lateral margins more The holotype (NIGP 113849) shows the external broadly curved but also not known in detail. surface covered with ornament. Shell surface ornamented with comarginal rugae, growth lines and radiating threads. Comarginal Other material rugae stout and evenly distributed, more than 20 in The three paratypes, NIGP 113850-113852 are number, gradually becoming stronger from apex more or less complete to fragmentary, but all toward abapical margin, with the distance between contribute information on the musculature. rugae also gradually becoming wider. Growth lines Late Cambrian molluscs from China 381 Figure 2 Piliiia liaoitittgensis, sp. nov. a, b. Paratype 113850. a, dorsal view, showing the serially arranged muscle scars on the right side of the dorsum, x 5. b, diagrammatical drawing of NIGP 113850: A-H = pedal retractor muscle scars; 1 = medio-pedal muscle scar; 2 = latero-pedal muscle scar; 3 + 4 = musculus branchialis and pallial muscle; ra. m. s. = radular muscle scars; s.g.c. = scar of gill cavity; b.m.f. = bifurcated muscle fibres, x 2.25. Yu Wen, E.L. Yochelson :^82 Late Cambrian molluscs from China 383 rather fine, closely spaced. Radiating threads pair being larger and rope-like in appearance, crossing rugae and growth lines to form a concellate situated anterior to A to C pairs of muscle scars. sculpture. The second pair are located near the anteriorly In lateral view of steinkern (Figure 3g), apex lateral side and are stripe-like in appearance and protruding and overhanging adapical margin, extend posteriorly. Tine third pair are subcircular, dorsal side flattened and inclined gradually situated near the lateral margin and next to the downward to the abapical margin. Dorsum of second pair. Furthermore, between the stripe-like steinkern broadly rounded in the central part with a muscle scars and the subcircular muscle scars, there rather shallow dorsal furrow, bounded by two are five small grain-like muscle scars (Figures 2, obtuse ridges. On the outer side of these, muscle 3f,g). On both sides of A to C pairs there are at least scars are present raised above the general level of two pairs of muscle scars parallel to the radular the steinkern. Scars are interpreted as one pair of muscle scars. radular muscle scars and eight metameric pairs of A broken steinkern is shown on Figure 3i. composite muscle scars (i.e. Figure 2 A to H), other Preserved on the right of the inner side are five interior shell marking are present. preserved muscle scars, which are similar in shape and size to the D-FI pairs of muscle scars illustrated Details of inusciilations in Figure 2, 3f, g, h, 4a, b. Radular muscle scars are largest and most A more or less weathered specimen is figured m pronounced of internal features. These scars are Figure 4c, d, e. It has the traces of the radular subelliptical in plan view, complex, being composed muscle scars, and the A to FI muscle scar sets on the of more than thirty small scars, and situated one- right side are clearer than those on the left side third of distance between adapical margin and midline. Dimensions (mm) The A to C pairs of muscle scars are situated on Length Height Width both sides of the radular muscle scars. All three NIGP 113849, Holotype 45.50 12.00 29.50 pairs of muscle scars are linked together in the anterior portion of the shell. The D to H pairs of NIGP 113850, Paratype 43.50 - 23.00 NIGP 113851, Paratype 34.00 - 21.50 muscle scars are discrete and are located in the posterior of the shell. They are all similar and nearly horseshoe-shaped (Figure 2). Each muscle scar can Remarks be divided into four parts; two larger and two Species of Pilim have mainly been reported from smaller. Part (3) gradually becomes smaller and the Upper Ordovician of the Baltic Region (Koken weaker from D to H coordinate with Part (4) which and Perner, 1925) and Oklahoma (Peel, 1977), and becomes larger and stronger and develops two from the Middle Silurian of Gotland, Sweden small pits. Each muscle scar set but particularly in (Lindstrbm in Angelin and Lindstrom, 1880; D to H, is flanned by eight or more small scars is Lindstrom, 1884). The new species ol Pilina distributed. These muscles extend outward and described herein represents the earliest record of display a secondary, tertiary or more bifurcation. the genus. The species is peculiar in its prutoconch, In addition, on the outer sides of the muscle scars more complex muscle scars and thereby differs sets are smaller scars. These are on the outside of from the all known species. the main scars. Their placement is in the lateral gill Proplina Kobayashi, 1933 is known from the cavities, extending backward from C to G pairs. The Upper Cambrian to Lower Ordovician ot North anterior pairs are thinner and shallower, than those America and Northeast Asia. Musculature is more toward the posterior which wider and deeper; these prominent in species of Pilina. Proplina has a higher latter display a regular wave-like curvature. more rounded cross-section than Pilina. Thus we As the adapical part of this internal mould has felt justified in extending the downward range of been slightly damaged, the muscle scars of the Pilina. anterior body region are not completely known on Judging from the broadly flattened dorsum, the this steinkern, there are three pairs of muscle scars presence of eight pairs of pedal muscle scars, Pilina on both sides of the anterior body region. The first liaoningensis is similar to Proplina sibeliitsi Figure 3 Pilina liaoningensis, sp. nov., a-e, holotype NIGP 113849. a. dorsal view, showing comarginal rugae, radiating threads and growth lines; b. enlargement of ornament, c. anterior view, showing the symmetrical protoconch (black arrow); d. right-lateral view, showing profile and rugose comarginal ornament; e. oblique right-lateral view; f-h, paratype NIGP 113850. f. dorsal view, showing the eight metameric muscle scars and radular muscle scars' g right-lateral view, showing the scars of gill cavity and other internal structures on the right side of the shell; h. oblique right-lateral view; i. paratype NIGP 113832, interior view of a substantial specimen, showing five muscle scars on the right side of the shell. Scales = (a, e, h) x 1.5, (b) x 10, (c, d, f, g) x 2, (i) x 4. 384 Yu Wen, E.L. Yochelson Late Cambrian molluscs from China 385 Stinchcomb (1986, p. 613, figs. 2.6-2.8, 3.11-3.13) Diagnosis from the Lower Ordovician Gasconade F-onnation Spire depressed. Base flattened. Whorls of Missouri, U.S.A., but differs from the latter increasing slowly in early stages, more abruptly species in the shell being elliptical in dorsal view, in in body whorl, embracing about half of the flatter dorsal side, in the narrower A to C pairs penultimate whorl on basal surface. Width of of pedal muscle scars and in having bifurcated body whorl about two times that of inner whorls. muscle fibres. Approximately seven whorls at maturity. In some respects, particularly in the flattened Pseudoumbilicus wide and deep, about one-half dorsal side, this species resembles Biptilvina cwfisae diameter of shell. Aperture ovate. Growth linos Yochelson (1958, p. 10, figs. 5-9; Knight and closely spaced. Yochelson, 1960, p. 178, fig. 46-5; Wingstrand, 1985, p. 49, fig. 19) from the Lower Ordovician, Description Gasconade Formation of Missouri. That form is Shell of medium size, very low-spired, sinistral lower and flatter and its muscle scars have a more in orthostrophic orientation. Protoconch triangular shape. unknown. Whorls five to seven in number, increasing slowly and gradually in the early Horizon and locality ones, abruptly in the body whorl, especially near Wanwankou Member of Fengshan Formation, at the apertural region and embracing about one- Doufangkou. half of the penultimate whorl on its basal surface. Width of the body whorl about two times that of the inner whorls and the height Class Gastropoda Cuvier, 1797 about two-thirds of the width. Basal surface Order Archaeogastropoda Thiele, 1925 slightly convex in early stages, gently rounded at maturity. Basal suture distinctly impressed. Suborder Macluritina Cox and Knight, 1960 Outer whorl surface gently rounded, the Superfamily Macluritacea Fischer, 1885 juncture of basal and upper faces following the arc of a small circle. Upper surface inclined Family Macluritidae Fischer, 1885 inward, following the arc of a circle of smaller Genus Maclurites LeSueur, 1818 diameter than the basal one. Pseudoumbilicus wide and deep, approximately one half the Remarks diameter of the shell. Pseudoumbilical walls The type species of Maclurites is from the Middle inclined steeply downward. Aperture not well Ordovician and the limits of the genus are not clear known, but generally ovate in shape. Surface of at this time. Thus the generic name is used in a shell mainly smooth but faint lines of growth are quotational sense. This taxon does not seem to be present near aperture (Figure 4j, k). Operculum closely allied to Scaevogi/ra. unknown. "Maclurites" ulrichi (Kobayashi, 1933) Dimensions (mm) Figures 4g-4k Length Height Width Ap. height NIGP 113853 38.00 21.50 16.00 15.00 Scaevogyra ulrichi Kobayashi, 1933: 266, pi. V, figs. NIGP 113854 23.00 14.50 11.50 10.50 5-7;'Yu et al, 1963: 96, pi. 27, figs. 7-9. NIGP 113855 21.50 9.00 Scaevogi/ra naticaformis Kobayashi, 1933: 267, pi. V, fig. 8; Yu et al, 1963: 96, pi. 27, figs. 5-6. Remarks "Maclurites" ulrichi (Kobayashi): Yu, 1990: 145. The description is based on three specimens and one incomplete internal mould. The shell is Material Examined characterized by a distinct, but very low-spired and Four specimens, NIGP 113853-113856. rapidly expanded body whorl. Figure 4 a-e. Pilina liaoiiiiigeusis, sp. nov. a-b, paratype NIGP 113850. a,b, dorsal views, enlargement of muscle scars; c-e, paratype N’lGP 113851. c, dorsal view, showing the eight paired muscle scars on the right and left sides of the dorsum, d, e. right-lateral and left-lateral views, showing the muscle scars on the right and left sides of the shell, f. Matherella walcotli Kobayashi, abapertural view. NIGP 113857. g-k. "Maclurites" ulrichi (Kobayashi), g-h, apertural and top views, NIGP 113854; i, basal view, showing the depressed spire, NIGP 113855. j, k. basal and apertural views, showing the growth lines and the form of aperture, NIGP 113853.1-n, Lytospira wamvaiikouensis, sp. nov., holotype NIGP 113858. 1. apical view, m, n oblique apical views. Scales: (a) X 7.5; (b) x 4; (c, d,e) x 2; (f, i, j, k, 1) x 1.5; (g, h, m, n) x 1. 386 Yu Wen, E.L. Yochelson In the general outline of the shell, in the well Remarks rounded outer surface, especially the wide The specimen figured here agrees in general pseudoumbilicus, this species resembles character with those described by previous authors. "Macluriles" thomsoni Webers et al. (1992, p. 200, pi. However, our specimen is larger than any 6, figs. 1-8; pi. 7, figs. 1-7), from the Upper previously illustrated for the taxon. Cambrian Minaret Formation of Heritage Range, West Antarctica. It differs from the Antarctic Horizon and locality species in having a depressed spire, more slowly Wanwankou Member of Fengshan Formation, at increasing early whorls and in the proportion Qiushugou. between the height and the diameter of the shell and in the ovate aperture. In some respects, this species is also closely Superfamily Euomphalacea de Koninck, 1881 allied to Maclurites niuhsintaiensis Kobayashi Family Euomphalidae de Koninck, 1881 (1931, p. 97, pi. 10, figs. 5a-c; Yu, 1961, p. 379, pi. V, figs. 10-12; pi. VIIl, figs. 8-10; Yu et al, 1963, Genus Lytospira Koken, 1896 p. 86, pi. 25, figs. 4-6) from the Lower Ordovician Machiakou Formation of Niuxintai Basin, Remarks Liaoning. It is easily distinguished from M. This genus superficially resembles Ecculiomphalus niuhsintaiensis by its depressed spire, more Portlock, 1843, but differs in having a blunt, spiral rounded basal edge, more obtusely rounded angulation on or near the midline of the upper umbilical edge and by the ovate form of the whorl surface. aperture. To the best of our knowledge multiwhorled open- coiled gastropods have not been reported from beds as old as Upper Cambrian. Despite the limited Horizon and locality amount of material we deem it worthwhile to name Wanvvankou Member of Fengshan Formation, at a species so as to emphasize the early development Doufangkou. of this type of morphology. Family Scaevogyridae Wenz, 1938 Lytospira ivanivankotiensis, sp. nov. Genus Matherella Walcott, 1913 Figure 41-n Remarks Material Examined The genus Matherella Walcott, 1913 is Late A single internal mould, NIGP 113858 (holotype). Cambrian in age. It is widely distributed, having been originally described from the Upper Cambrian Type locality Hoyt Limestone of eastern New York and Chiushukou section, Niuxintai, Liaoning subsequently from the Upper Cambrian Minaret Province, China (see Figure 1). Formation of Heritage Range, West Antarctica (Webers t'f«/., 1992). Diagnosis Shell discoidal, with blunt spiral angulation near midline of upper whorl. Whorls four to five in Matherella walcotti Kobayashi, 1933 number, first two whorls in contact, but later ones Figure 4f openly coiled. Matherella walcotti Kobayashi, 1933, p. 261, pi. IV, fig. 7; pi. V, fig. 9. Description Matherellina walcotti (Kobayashi): Knight 1941: 189, Shell small and discoidal. Whorls four to five in pi. 90, figs a-c; Knight et al, 1960: 1187, Fig. 104- number, the first two in contact, with later whorls 5; Yu et al, 1963: 97, pi. 27, figs. 12-13; Yu, 1987: open-coiled and rapidly mcreasing in size. A blunt, 93, text-fig. 30a-b; 1990: 145. spiral angulation is near the midline of the upper whorl surface, sloping inward towards the inner Material Examined whorl surface. Outer whorl surface is gently convex. One internal mould, NIGP 113857. Suture in the first two whorls is distinctly impressed. Surface of the steinkern is smooth. Diagnosis Aperture and the characters of base unknown. High spired, trochiform, sinistral in orthostrophic Growth lines unknown. orientation. Outer face slightly convex, sutures distinctly impressed. Surface ornamented with Remarks rugae. According to previous records, the genus Late Cambrian molluscs from China 387 Li/tospira occurs from the Lower Ordovician to the laterally both sides of the radular muscle scars Middle Silurian in North America and Europe. The closer to the shell margin. These three pairs of present record of the genus from north-east China elongate muscle scars arc linked together. The D to thus extends the time-range of Lytospira back to the H pairs of muscle scars are located posterior to the Late Cambrian. midlength of the shell. They are all similar and In apical view, this species is somewhat similar to nearly horse-shoe-shaped (Figure 2). the type species Lytospira aiigelini (Lindstrbm) (1884: The recent Neopilina galatheae Lemche (1957) has 138, pi. XIII, figs. 36-38; Knight et al. 1960: 1191, fig. been studied in detail. (Lemche 1957; Lemche and 107-6) from the Middle Ordovician of Sweden. Wingstrand 1959; Wingstrand 1985). According to Lytospira ivanwankouensis differs in having the Lemche and Wingstrand (1959) each pedal retractor whorls discoidal in the same plane and in having a mu.scle can be divided into two parts: (1) and (2), more obtuse spiral angulation. It also shows some we interpret D-H as pedal retractors in part affinities to Lytospira valida Koken in Koken and because of their position and in part because of the Perner (1925, p.ll5, pi. XIII, fig. 3), but differs from similarity of the muscles to those of Neopilina. Even the latter in having tightly coiled early whorls and a though we can discern four parts, the overall shape more concave inner whorl surface. of the scars is basically bipartite. The same bipart division can be seen in the scar of P. liaoningensis. Horizon and locality Latero-posteriorly on the D to H pairs of muscle Wanwankou Member of Fengshan Formation, at scars there is a small distinct scar (part 3) is situated Qiushugou. in the left corner and another clear small scar (part 4) is located at the right corner of each main muscle scar. We present that the two main biparts of each DISCUSSION scar control: parts (1) and (2) control the foot and Tergomya are rare in the Fengshan Formation but provide for clamping and crawling movement; Filina liaoningensis, sp. nov. deserves special note parts (3) and (4) more likely control activity of gill because of its excellent preservation of the ornate lobes and pallial muscles. As noted part (3) tends to shell, muscle scars and other internal structures, gradually become smaller and weaker from D to H; which projects like a raised platfoim in outlook, while part (4) becomes larger and stronger and has including one pair of radular muscle scars and eight two small pits. In P. unguis, the A to C sets are metameric pairs of composite muscle scars (i.e. the bigger and wider than those in P. liaoningensis, and A-H pairs of muscle scars) (Lemche and the D to H sets are subdivided into more than four Wingstrand, 1959). The overall shape shows a smaller parts (Lemche and Wingstrand 1959, figs. configuration similar to that of the type species 133, 134). In P. cheyennica the A and B sets are Filina unguis Lindstrom (Lindstrbm, in Angelin and coalesced, the C to H sets are not subdivided into Lindstrbm, 1880; Lindstrbm, 1884) from the Middle the various minor scars. Silurian of Gotland, Sweden and that of P. The bundled impressions lateral to the pedal cheyennica Peel from the Upper Ordovician Keel muscle scars appear unique. They show secondary Member of the Chimneyhill Limestone, Oklahoma and tertiary bifurcation and extend to the lateral (Peel, 1977). margin. These do not stand in relief like the The paired radular muscle scars in P. liaoningensis presumed pedal scars. It is possible that these arc are well preserved and are the biggest in the the marks of blood vessels. We doubt that blood muscular field. In detail these scars are subelliptical vessels would be impressed into the shell. Our in plan view and very complex, composed of more interpretation is that this muscle extends to edge of than thirty small scars. The radular scars are the mantle which would shape the contours of the situated high on the dorsum just about one-third of lateral gill cavity. the distance along the length of the shell. In the The scars of the gill cavity are between the pedal type species P. unguis (Lindstrbm), the radular muscle scars and the shell margins, and are another muscle scars are nearly tadpole-shaped and smaller interesting structure in this species. They extend in size (Lindstrbm, in Angelin and Lindstrbm, 1880: backward from C pair to G pair, the anterior ones 16, pi. II, figs. 10-14, excl. fig. 15; Lindstrbm, 1884: being thinner and obscure, while those toward the 56, pi. I, figs. 33-37; pi. XIX, fig. 2; Knight, 1941: 246: posterior are wider and deeper, displaying a 4, figs. 1-3; Knight and Yochelson, 1960: 178, Fig. regular wave-like curvature. The concavely curved 48-6; Lemche and Wingstrand, 1959: 44, figs 133, surface is broader than the convex side, whereas in 134 162A-D; Wingstrand, 1985: 449, Fig. 19) while P. cheyennica the scars of the gill cavity are obscure. in Filina cheyennica Peel, the radular muscle scars In addition, there are at least three pairs of muscle arc subrectangular in shape (Peel, 1977, 117-121, scars on both sides of the anterior body region. The text-figs. 1-3; Wingstrand, 1985: 46, fig. 19; Peel first pair being larger and rope-like in appearance, 1991: 7, fig. 3c). situated anterior to A to C pairs of muscle scars. The A to C muscle scars pairs are situated The second pair are located near the anteriorly 388 Yu Wen, E.L. Yochelson lateral side and are stripe-like in appearance and in northern China and its international correlation. extend posteriorly. The third pair are subcircular, Gcologica et Palaeontologica 17: 1-15. situated near the lateral margin and next to the Clarke, A.H., Jr. and Menzies, R.J. (1959). Neopilina second pair. Furthermore, between the stripe-like (Venia) ewingi, a second living species of the Paleozoic muscle scars and the subcircular muscle scars, there Class Monoplacophora. Science 129: 1026-1027. are five small grain-like muscle scars (Figures 2, Knight, J.B. (1941). Paleozoic gastropod genotypes. 3f, g). On both sides of A to C pairs of muscle scar Geological Society of America, Special Paper 32: 1-510. sets, there are at least two pairs of muscle scars Knight, J.B. and Yochelson, E.L. (1960). Monoplacophora. In Moore R.C. (ed.). Treatise on invertebrate parallel to the radular muscle scars. paleontology. I, Mollusca 7: 177-183. Lawrence, The protoconch of Pilina liaoningensis sp. nov. is a University of Kansas Press. small, symmetrical cone, protruding strongly Knight, J.B., Batten, R. and Yochelson, E.L. (1960). forward (Figure 3c). Although external features and Descriptions of Paleozoic gastropods. In Moore, R.C. tip of the protoconch are not preserved, this (ed.). Treatise on invertebrate paleontology. 1. Mollusca 7: specimen further establishes that the Tergomya selected pagers. Lawrence, University of Kansas have a symmetrical protoconch (Clarke and Press. Menzies 1959; Menzies and Layton 1963; Peel 1977; Kobayashi, T. (1931). Studies on the stratigraphy and Wingstrand 1985). In P. citei/eitnica (Peel 1977), from palaeontology of the Cambro-Ordovician Formation the Upper Ordovician, the protoconch is a small of Hui-lien-Chai and Niu-hsin-tai, South Manchuria, mamillate protuberance. Japanese Journal of Geology and Geography 8: 131-186. Ornament in Pilim varies among .species. In P. Kobayashi, T. (1933). Faunal study of the Wanwanian liaoningenensis, the outer surface is covered with (Basal Ordovician) Series with special notes on the more than twenty stout and evenly distributed Ribeiridae and the Ellesmericeroids. Imperial comarginal rugae, the interspaces between the Universiti/ of Tokyo Faculty of Science Journal 3: 249- rugae being covered with fine lines of growth. This 328. ornament is crossed by radiating threads, resulting Koken, E., and Perner, J. (1925). Die Gastropoden des in a cancellate pattern. In P. unguis, the ornament Baltischen Untersilurs. Academic des Sciences de Russie Memores, Classe Pln/sico-Mathematique, 8th series, 38: comprises growth lines and faint, widely spaced 1-326. grooves, where as in P. cheymnica, the outer surface Lemche, H. (1957). A new deep-sea mollusc of the is covered with lamellar growth increments, which Cambro-Devonian Class Monoplacophora. Nature may be irregularly rugose. 179: 413-16, Lemche, FL, and Wingstrand, K.G. (1959). The anatomy of Neopilina galatheae Lemche, 1957. Galathea Report 3: ACKNOWLEDGEMENTS 9-71. We are grateful to Xu Jun-tao, Nanjing Institute of Lindstrdm, G. (1884). On the Silurian Gastropoda and Geology and Palaeontology, Academia Sinica, for Pteropoda of Gotland. Kungliga Si’cnska Vetenskaps- lending the specimens, to Song Zlii-yao also of the akademiens Fiandlingar 19: 250. Institute, for preparing photographs, and to Lu Yan-hao. (1962). The Cambrian System of China. Science Danielle West, Western Australian Museum for Press, Beijing, 177 pp. (in Chinese). drawing the text-figures. YW thanks to K.J. Lu Yan-hao, Zhu zhao-ling, Qian Yi-yuan, Lin Fluan- McNamara and G.K. Kendrick of the Western ling, Zhou Zhi-yi and Yuan Ke-xing. (1974). Bio- Australian Museum for reading the manuscript and environmental control hypothesis and its application appreciates the use of museum facilities. Prof. J.S. to the Cambrian biostratigraphy and Palaeo- zoography. Memoirs of Nanjing Institute and Peel significantly improved the work. Palaeontology, Academia Sinica 5: 27-116. Science Press, Beijing (in Chinese). Menzies, R.j. and Layton, W. Jr. (1963). A new species of REFERENCES monoplacophoran mollusc, Neopilina (Neopili)ia) Angelin, N.P. and Lindstrdm, G. (1880). Fragmcnla ivlero)iis from the slope of the Cedros Trench, Mexico. Siliiriai e lioiio Caroli Hcnrici Wegetin. 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