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American Museum Published by the American Museum of Natural History Central Park West at 79Th Street New York, N.Y NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2685 OCTOBER 9, 1979 ROGER L. BATTEN Permian Gastropods from Perak, Malaysia Part 2. The Trochids, Patellids, and. Neritids AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2685, pp. 1-26, figs. 1-33 October 9, 1979 Gastropods from Perak, Malaysia Part 2. The Trochids, Patellids, and Neritids ROGER L BATTEN' ABSTRACT This is a continuation of a study of Permian recorded, 10 species of trochids, four new, and nine gastropods from the Misellina claudiae zone from species of neritids, three new, are described. Most of the H. S. Lee Mine no. 8 near Kampar, Perak, the neritids are similar to morphotypes found from Malaysia. The fauna constitutes one of the richest, Djebel Tebaga east to Mongolia. Key characters in most diverse, known in the Permian, providing a Trachydomia imbricata, new species, suggest an an- critical base in the eastern Tethys. New species of cestral relationship to Neritopsis. Wall ultrastructure Sallva Yochelson, 1956, and Dic hostasia Yochelson, is reported and discussed for Trachyspira delphinu- 1956, record their presence for the first time outside loides Gemmellaro, 1889, and for the Recent of the west Texas region. One species of patellids is Neritopsis radula Linne, 1758. INTRODUCTION This study is a continuation of the descrip- Two genera, Sallya Yochelson, 1956, and tion of the Permian Malaysian gastropods. It is Dichostasia Yochelson, 1956, are included here based on the marine molluscan fauna found at a because they are trochid-like in appearance; single locality, Lee Mine no. 8 near Kampar, they belong to the superfamilies Pseudophora- Perak (Map of Malaysia sheet 2n/9 old series, cea and Caspedostromatacea, respectively. [MR 90356], see Batten, 1972, p. 5 for de- These superfamilies have been thought to be tails). The fauna is dominated by gastropods problematic and probably polyphyletic. It has but corals, scaphopods, bivalves, brachiopods, not been possible in the past to assign them to and cephalopods are present along with any known order. They generally are placed in fusulinids and other Foraminifera. Based on the Archeogastropoda with a query (Knight, fusulinids, the unnamed white-colored lime- Batten and Yochelson, 1960). I would be in- stone is dated as Late Artinskian-Early clined to refer them, as aberrant groups, to the Guadalupian (Jones, Gobbett and Kobayashi, Suborder Trochina. Until a thorough analysis of 1966, p. 328), and is included as a part of the these groups are made, I provisionally accept Misellina claudiae zone. As mentioned in the them as being internally consistent on mor- earlier study, this fauna is, with one exception, phological grounds. This would be particularly the richest known in the Permian. apropos for the Pseudophoracea since some of 'Curator, Department of Invertebrates, American Museum of Natural History. Copyright (© American Museum of Natural History 1979 ISSN 0003-0082 / Price $2.00 2 AMERICAN MUSEUM NOVITATES NO. 2685 the genera have been reported to have nacre of primary noding, the construction of the aper- (Knight, Batten and Yochelson, 1960, p. 297). tural complex and the early development of the Sallya and Dichostasia are of particular interest basic ornament pattern. The details of ornament from the paleobiogeographical view because and the basic apertural design in some of the they were originally described from the Per- variant patterns of T. gobbetti, new species, mian of the west Texas region and were previ- suggests that the morphotype represented by ously known only from that region. This this species could also have been the source for disjunct distribution was noted in the case of Hungariella. two pleurotomarian genera, Lacunospira Bat- Trachyspira Gemmellaro, 1889, on the other ten, 1958, and Lamellospira Batten, 1958. It is hand, is primarily known from the Middle Per- interesting to note that the four genera in ques- mian Sosio beds where five species have been tion are highly unique forms. Further, the spe- described. The genus is represented in the Ma- cies between the two areas are remarkably laysian sample by 12 specimens of T. delphinu- similar in details. loides Gemmellaro, 1889, and is moderately The patellids are represented by a single abundant in the Djebel Tebaga. The only other specimen of the genus Lepetopsis Whitfield, occurrence of Trachyspira that I am aware of is 1882, reflecting the fact that, in general, the from the Middle Permian of the west Texas patellids are not a common element in Upper region. Paleozoic faunas. Finally, the radiation of the Neritopsidae The neritacean species are similar to those during the middle Permian and into the Triassic found in the Sosio beds of Sicily, the Permian parallels that of the early radiation of the of Timor and Sumatra, the Salt Range of Kash- caenogastropods which is first encountered in mir, Yunnan, Mongolia, the Djebel Tebaga of the Malaysian fauna. In the latter radiation, a southern Tunisia, and the west Texas Permian number of neomorphs appear in the Malaysian and can be identified as typical of the Tethyan fauna which are the earliest representatives of faunal realm. The neritacean genus Tra- several important Mesozoic groups first encoun- chydomia Meek and Worthen, 1866, is wide- tered in the Mid to Upper Triassic (the families spread in distribution and is known from many Coelostylinidae, Procerithiidae, Potamididae (?) Upper Paleozoic faunas. It is not particularly and Nerineidae). abundant except in the Djebel Tebaga (see Ter- Naticopsis M'Coy, 1844, is represented by mier, Termier and Vachard, 1977) and the four species belonging to four of eight mor- Sosio faunas. The earliest reported occurrence photypes recognized in the Middle American of Trachydomia is in the Lower Pennsylvania Pennsylvanian (see Knight, 1933, pp. 366-367). of the midwestern United States. It was proba- These species are in the middle of the mor- bly derived from some group like the Lower phological spectrum of the genus and are diffi- Carboniferous Turbinitella de Koninck, 1881, cult to treat owing to a paucity of distinctive which shares such characters as a concave sub- features. Most Permian species seem to be sutural ramp and a thickened apertural lip with fairly restricted to given sub-provinces of the a concave trough. Tethys. For example, N. depressa (Guembel), Two species of Trachydomia resemble the 1877, has only been reported from the Bellero- Triassic genera Hungariella Kutassy, 1934, and phonkalk and associated strata in Europe. N. Neritopsis Grateloup, 1832. The latter genus is khurensis Waagen, 1880, is widely reported represented today in the Indo-pacific fauna by a from central Asia; at Karakorum and the Salt single species N. radula (Linne), 1758, the Range; from Malaysia to Sumatra and north only extant species of the family Neritopsidae; through Yunnan and Kweichow to Mongolia. this family includes 14 extinct genera distrib- As mentioned in an early report of the Ma- uted primarily in the Late Paleozoic and Meso- laysian pleurotomarians (Batten, 1972, p. 5), zoic. Neritopsis must have been derived from the preservation of the fauna is unusual in that an ancestor very close to that of Trachydomia specimens are powdery and fragile and mainly imbricata, new species; the details of the com- natural casts. They are found in a matrix of plex lamellate ornament are identical (see fig. decomposed limestone partly resulting from 21) as well as the formation and development baking by an intrusive underlying the mine. 1979 BATTEN: GASTROPODS 3 Batten (1972, p. 5) stated that no wall ultra- or can be transformed from, a complex pris- structure was found in the pleurotomarians, matic structure. Based on observations within suggesting that the specimens were natural more advanced groups, there is consistency of casts. While examining the sample of Tra- wall structure within each group (even at the chyspira delphinuloides Gemmellaro, 1889, it superfamily level). It would not be likely to was found that several specimens showed some find these different types in the same sub- ultrastructure preserved (see fig. 32). The struc- family. The problem is not soluble at present. ture was observed on a freshly broken portion As defined here, a morphotype is a relative of the adult whorl and not on breaks that oc- term describing a group of specimens, usually a curred before lithification of the lime mud species, possessing a distinct set of characters when the broken surface could have been im- which recur in different groups in space and/or pressed into the mud to form a mold, which time, allowing for convergence but without im- subsequently could have been filled by second- plying, necessarily, direct genetic relationships. ary calcite. SEM examination revealed that the wall is composed of a single massive complex ACKNOWLEDGMENTS prismatic layer reminiscent of the wall found in Bellerophon Montfort, 1808 (see MacClintock, I again thank Dr. Derek J. Gobbett of 1967, pp. 94-107). The apices of the prisms Cambridge University (formerly of the Geology undulate with the growth lines, indicating that Department, University of Malaya) for bringing there is no outer layer of wash formed by the this very important fauna to my attention and mantle. for permitting me to study it. Dr. Keiji I was unable to obtain specimens of the Nakazawa of Kyoto University deserves special Triassic species of Neritopsis but I sectioned thanks for sending vital additional specimens the Recent species N. radula (Linne), 1758, collected during the Southeastern Scientific Ex- which is composed of the relatively thin outer pedition made by members of the Ozaka City spiral crossed-lamellar layer and an inner col- and Kyoto universities. I gratefully thank Mr. labral crossed-lamellar layer, having the same G.
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