Yunannioidea—a new superfamily of Pteriomorphids (Bivalvia) from the Middle Ordovician, Yunkai Area, western Guangdong, South China
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Abstract:
The Ordovician in Yunkai area of western Guangdong Province mainly consists of sandstone, pelitic siltstone, silty mudstone and pebbly sandstone, and mudstone with intercalated carbonate rock lenses and conglomerates. The Ordovician includes the Lower Ordovician Luohong Formation and Luodong Formation, Middle Ordovician Dongchong Formation, and Upper Ordovician Lanweng Formation. Fossil bivalves, including the new taxa described in this paper, were collected from an about 2-meter-thick bed of gray to grayish yellow silty mudstone and pelitic siltstone of the upper part of the Dongchong Formation. The collection includes about one thousand bivalve specimens, which represent more than 22 species of 16 and several unnamed genera and may a possible new taxon. We report a group of unique bivalves including two new species of Yunannia gen. nov. of a new family Yunanniidae Zhang et Niu and a new superfamily Yunannioidea Zhang et Niu. This group demonstrate combined features normally attributed to different bivalve taxa. A small number of trilobites, including Nileus sp., Lonchobasilicus sp., Calymenesun tingi Sun, Calymenesun sp., Asaphopsis? sp., and brachiopods, including Paralenorthis sp., Aegira sp., Leptellina sp., Obolus? sp., Strophomena sp. and Nicolella sp., cooccur with the bivalves and indicate the Middle Ordovician age. SYSTEMATIC PALEONTOLOGYClass Bivalvia Linnaeus, 1758 in 1758–1759 Subclass Autobranchia Grobben, 1849 Infraclass Pteriomorphia Beurlen, 1944 Cohort Uncertain Superfamily Yunannioidea Zhang et Niu superfam. nov. Description Shell small, mytiliform, umbos anterior to terminal; strong inequilateral; buyssal sinus faint; the area beneath and anterior umbo edentulous; posterior to umbo a row of discrete chevron taxodont teeth; large anterior adductor muscle scar located in the umbonal angle and occupies the large and wide umbonal septum together with the anterior pedo-byssal retractors; posterior muscle scar unknown; external ligament, opisthodetic, submarginal; ligamental area narrow with 2-ranks of ligamental grooves and ridges, which are gently arched and extended continuously away from the dorsal margin; shell surface with faint comarginal growth lines. Remarks The Yunannioidea, a new superfamily of pteriomorphians is established based on the genus Yunannia proposed in the present paper. The Yunannia, the sole genus of the new superfamily Yunannioidea, is characterized by mytiliform outline, one posterior row of chevroned teeth, the large anterior adductor muscle scar located in the umbonal angle and inserted on the umbonal septum together with the anterior pedo-byssal retractor, and the arched two ranks of ligamental grooves and ridges which extend continuously away from dorsal margin. On the basis of features such as mytiliform, the musculature, and ligament, the Yunannioidea may be placed into the Cohort Mytilomorphi!. However, the genus Yunannia has only one row of posterior chevroned teeth. Together with its unusual hinge dentition, the new superfamily can be distinctly distinguished from other members of the same cohort. Meanwhile, features including the taxodont dentiotion and the two ranks of ligament indicate that Yunannia is likely in connection with Subcohort Ostreioni. Hence, the position of Yunannia at cohort level is undetermined. The genus Yunnania most possibly represents a new order of the infraclass Pteriomorphia. Family Yunannidae Zhang et Niu fam. nov. urn:lsid:zoobank.org:act:8A609836-4A82-4C6A-97B1-18168E761 891 Type genus Yunannia Zhang et Niu Yunannidae is the sole family of the new superfamily Yunannioidea. Distribution Middle Ordovician, Guangdong, China. Genus Yunannia Zhang et Niu gen. nov. urn:lsid:zoobank.org:act:B09DAFBA-A54D-4621-8FCD-07D8882 A6584 Type species Yunannia gankengensis Zhang et Niu gen. et sp. nov. Diagnosis Shell mytiliform, umbos anterior to terminal, with a row of posterior, discrete chevron taxodont teeth; anterior adductor muscle scar large and pedo-byssal retractor inserted on umbonal septum; ligament external and opisthodetic, extending continuously away from dorsal margin. Description The shell is small with height less than 21 mm. The shell shape is strongly inequilateral, mytiliform or narrow ovate with height longer than length. The umbos is anterior to terminal, projected and slightly incurved. Anterior lobe is absent. The dorsal-posterior margins are merged and widely arched. The anterior flange is thickened. The umbnal cavity is large and deep, separated by a lunate inner septum. The anterior adductor muscle scar is large and elliptical, located in the umbonalangle, and inserted on the umbanl septum together with the posteriorly extended pedo-byssal retractor. The growth line of anterior adductor muscle scar is present in one specimen. The anterior end of the anterior adductor scar is acuminate, likely marking the position of the pedal protractor but is obviously merged with the anterior adductor. The posterior adductor and posterior pedal retractor are unknown. The holotype, a left valve, bears about 20 teeth and sockets, while each half of a paratype bears about 8–15 teeth. The area anterior to and beneath umbo is edentulous. All teeth are discrete and chevroned with concavities towards distal end. The anterior two or three teeth are small and the rest become gradually larger posteriorly, achieving the maximum size about midway along the row of teeth. The teeth diminish gradually in size toward the posterior end of hinge plate. The ligament is external and opisthodetic, submarginal, sub-parallel to the gently arched dorsal-posterior shell margin. The ligamental area is narrow and extends posteriorly to about 3/4–4/5 height of shell, with one rather coarse ligamental ridge and two grooves above and below it (as the first-rank of ligament), while the weak and faint ligamental ridges and grooves (as the second-rank of ligament) within the first-rank ligamental grooves and ridge. Because of the number of first-rank ligamental groove-ridge couplets, it is likely the preduplivincular ligament. However, if the second-rank of ligament is also considered, the total number of ligamnental groove-rige couplets is more than two or three. Hence, it is likely the duplivincular ligament, rather than the preduplivincular ligament. In addition, the arched shape of ligamental area is similar to the simple arched ligament. The holotype specimen shows faint growth lines on the inner surface of shell wall. Etymology Yunan, a geographic name of a county in Guangdong Province. Remarks This genus is the sole representative of Yunannioidea superfam. nov. It is characterized by features such as the mytiliform shell having a posterior row of chevroned teeth, and the arched two ranks of ligamental groove-ridge couplets extending continuously away from the dorsal margin. The large anterior adductor located in the umbonal angle and inserted on the umbanl septum together with anterior pedo-byssal retractor. In addition, the large and wide umbonal septum is another distinguishing feature of Yunannia. The new genus combines some main features normally attributed to different bivalve taxa. Its dentition of taxodont teeth is similar to the Paleotaxodonta. The new genus and Ambonychioidea of Pteriomorphia share some features, including the mytiliform shape, terminal umbos and byssal sinus. Both the new genus Yunannia and cyrtodontoids have the same characteristic of sub-umbonaledentulous area. The present new genus and Dreissenoidea of Heteroconcha have common characters such as the anterior muscle scar inserted on umbonal septum, the submarginal, long, extended arched ligament. With the combination of features listed above, the new genus clearly distinguishes itself from all other genera in those taxa. Age and distribution Middle Ordovician; Guangdong, China. Yunannia gankengensis Zhang et Niu gen. et sp. nov. urn:lsid:zoobank.org:act:67E7B718-0211-4EBB-ADA9-898045A3 8637 (Figs. 3–5) Materials 8 specimens of internal moulds including 5 left valves and 3 right valves. Holotype: No. Ng4032, Paratypes: Ng4033, Ng4030, Ng4031, Ng1041. Diagnosis Height/length(width) ratio less than 1.5; ventral part of shell expanded and swollen. Description The shell is small, mytiliform, strongly inequilateral with height less than 20 mm. The height is longer than the length (width) with height/length(width) ratio less than 1.5. The umbos is terminal, projected and slightly incurved. The anterior lobe is absent. A faint byssal sinus is present at the upper part of anterior margin. The dorsal-posterior margins are merged and widely arched. A long and thickened anterior flange is present behind the anterior margin. A large umbnal cavity is separated by a lunate umbonal septum. The elliptical and large anterior adductor muscle scar is located in the umbonal angle, together with anterior pedo-byssal retractor, occupying the umbonal septum. The anterior adductor muscle scar being covered by growth line is showed in one paratype specimen (No. Ng1041). The anterior end of the anterior adductor scar is acuminate, which likely marks the position of the pedal protractor, but is obviously merged with the anterior adductor (as to otherwise indistinguishable). The posterior adductor and posterior pedal retractor are unknown. The area anterior to and beneath the umbo is edentulous. Posterior to the umbo there is a row of teeth consisting of 13–20 discrete and chevroned teeth with concavities towards the distal end. The anterior and posterior two or three teeth are small with the maximum size at the midway along the tooth row. The ligament is external and opisthodetic, submarginal, sub-parallel to the gently arched dorsal-posterior shell margin. The ligamenal area is narrow, extending posteriorly away from dorsal margin to about 4/5 height of the shell with one rather coarse ligamental ridge and two grooves laid above and below it (as the first rank of ligament), which are subdivided by the weak and faint (as the second rank) ligamental ridges and grooves. Because of the number of the first rank ligamental groove and ridge couplets, it is likely the preduplivincular ligament. However, if the second-rankof ligament is also considered, the total number of ligamnental groove-rige couplets is more than 2–3. Hence, it is likely the duplivincular ligament, rather than the preduplivincular ligament. Faint growth lines are present on the shell surface. Measurement Holotype: No. Ng4032: height 17.5 mm, length 13, H/L = 1.35. Etymology Gankeng, a geographic name of Guangdong. Comparison The present new species evidently differs from Yunannia yunkaiensis gen. et sp. nov. in having mytiliform shape, shell becoming gradually wider towards ventral side, with the maximum width near the ventral part of the shell, and a height/length ration less than 1.5. The latter species has a narrow and narrow ovate shape and a height/length ration about 2. Occurrence Gankeng Village, Yunan County, Guangdong Province; Dongchong Formation, Middle Ordovician. Yunannia yunkaiensis Zhang et Niu gen. et sp. nov. urn:lsid:zoobank.org:act:BE7BF3A3-3F07-4049-88F3-2EE989D4 E914 (Fig. 6) Materials 3 internal moulds of left valve. Holotype: No. Ng4035, Paratypes: Ng074, Ng4034. Diagnosis Shape narrow ovate and H/L ratio about 2.0. Description The shell is small, narrow ovate, strongly in-equilateral, 16.5–21 mm high. The height is much longer than the length with a H/L ratio of 2.0. The umbos is projected and slightly incurved. The anterior lobe is absent. The dorsal-posterior margins are merged and gently arched. The thickened anterior flange is relatively narrow. The umbonal cavity is separated by lunate, relatively narrow umbonal septum. The large and elliptical anterior adductor muscle scar is located in the umbonal angle, and together with the pedo-byssal retractor, inserted on umbanl septum. The anterior end of the anterior adductor scar is deeply inserted and becomes shallow poster-ventrally. The posterior adductor and posterior pedal retractor are unknown. The area anterior to and beneath umbo is edentulous with a posterior row of hinge tooth consisting of more than 8 to 13 discrete and chevroned teeth with concavities towards the distal end. The ligament is external and epithetic, submarginal. The ligamental area is narrow, sub-parallel to the gently arched dorsal-posterior shell margin, extending posteriorly away from dorsal margin to about 3/4 height of shell, and with one rather coarse ligamental ridge and two grooves above and below it (as the first rank of ligament). The four weak and faint ligamental groove-ridge couplets (as the second rank of ligament) extend about 5 mm in length in the dorsal side of the groove of the first rank ligament. Surface ornaments of the shell are unknown. Measurement Holotype: No. Ng4035; height 21 mm, length 10 mm, ratio H/L = 2.0. Etymology Yunkai, a regional geographic name in Guangdong Province. Comparison The present new species is distinguished from the genotype, Yunannia gankenensis (gen. et sp. nov) by the narrow ovate shape with a H/L ratio of 2, and the slightly incurved umbo. In contrast to the latter species’ mytiliform shape and a H/L ratio less than 1.5. On the respect of shell shape, the present species is more or less similar to Mytilarca chenmungensis (Conrad) (Pojeta, 1966, pl. 37, figs. 8–18; pl. 38, figs. 1–5, 10), but it differs from the latter species by having a subumbonal edentulous area, a posterior row of taxodont teeth, a large anterior adductor muscle scar, and large anterior pedo-byssal retractors laid on unmbonal septum. Occurrence Gankeng Village, Yunan County, Guangdong Province, Dongchong Formation, Middle Ordovician.