AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2829, pp. 1-40, figs. 1-62 October 21, 1985

Permian Gastropods from Perak, Malaysia. Part 3. The Murchisoniids, Cerithiids, Loxonematids, and Subulitids

ROGER L. BATTEN'

ABSTRACT This is the final part of a study of Permian gas- Paleozoic. The presence of these and other dom- tropods from the Misellina claudiae zone from the inant post-Triassic families and the presence of H. S. Lee Mine No. 8, near Kampar, Perak, Ma- Paleozoic groups in the Triassic serve as additional laysia. The fauna constitutes one ofthe richest and evidence that a catastrophic event did not affect most diverse known in the Permian-91 species gastropods at the end of the Permian as it might of 52 genera. Included in this portion ofthe study have in nonmolluscan groups. Of interest is the are analyses of species ofthe murchisoniids, loxo- recognition of a number of species which have nematids, cerithiids, subulitids, nerineids, pyram- very similar morphologies to species from the idellids, and trochids. This study now provides North American Pennsylvanian, particularly the best documented Permian gastropod fauna of among Retispira, Naticopsis, Trachidomia, Wor- the Eastern Hemisphere. There are 33 new species thenia, Orthonema, and Pseudozygopleura. The ofa total of41 and 4 new genera described herein. presence of an unusually large number of siphon- The genera belong to the Murchisoniidae, Loxo- ate species (18) in the fauna may indicate the be- nematacea (incertae sedis), Procerithiidae, and the ginning ofthe exploitation ofinfauna habitats, per- Nerineidae; the latter two families are important haps in response to the origin of predation. Mesozoic groups reported for the first time in the INTRODUCTION This study completes the description ofthe Perak [map ofMalaysia sheet 2n/9, old series Permian Malaysian gastropods. It was based (MR 90356)]; see Batten, 1972, p. 5 for de- on the marine molluscan fauna found at a tails. As stated in the first part of this study single locality, Lee Mine No. 8 near Kampar, (Batten, 1972, p. 5), the preservation is quite ' Curator, Department of Invertebrates, American Museum of Natural History.

Copyright K American Museum of Natural History 1985 ISSN 0003-0082 / Price $3.95 2 2AMERICAN MUSEUM NOVITATES NO. 2829 uneven; much of the surface detail is pre- families. Mention should again be made of served in fine, powdery calcite dust which is the disjunctive distributions of Sallya, Di- so fragile that merely touching the surface chostasia, Lacunospira, and Lamellospira may destroy details. In consequence, there known only from this Lee Mine locality and are fairly large numbers (more than 300 of a in several horizons in the Permian of the total of 646 studied) of specimens which can southwestern United States. There are sev- be placed in high-level categories only. Many eral examples of disjunct distributions in- ofthese are high-spired, many-whorled forms volving distances of thousands of miles in that suggest placement in the Cerithiacea and the Paleozoic marine faunas. Another ex- do not seem to fit into any of the groups ample is the Cambrian molluscan (Hypse- described herein. Other specimens appear to loconus) fauna found in the north central belong to some of the trochid and subulitid United States and the Ellsworth Mountains families. On the whole, this gives the fauna of Antarctica. In this case about 10 genera a far more Mesozoic appearance than would are found in common; in fact, the species are appear to be true from the illustrations and nearly identical (Yochelson, personal com- synoptic classification. I cannot overempha- mun., 1983). size the importance of these indescribable An important observation made during the specimens injudging the nature ofthis fauna. course of this study was to find a number of There is an unusually large number of si- iterative morphotypic species between North phonate species in this fauna (some 18) com- American Pennsylvanian and the Malaysian pared to all other known Permian faunas Permian of the genera Naticopsis McCoy, where one or two may be present. Some of 1844, Trachydomia Meek and Worthen, the siphonate species are in groups such as 1866, Retispira Knight, 1945, Worthenia the murchisoniids which are always holos- DeKoninck, 1883, Orthonema Meek and tomous (with apertures which do not have Worthen, 1862, and Pseudozygopleura siphonal notches indicating siphons). This Knight, 1930. Bayer and McGhee (1984) have may reflect an early trend into the infaunal shown that repeated morphologies of am- realm (see the conclusions for a more detailed monites are intimately correlated with direc- discussion). tional changes in the physical environment The fauna is dominated by gastropods, but in the Middle Jurassic Basin of Germany. corals, scaphopods, bivalves, brachiopods, These morphologies are not derived from and cephalopods are present along with fu- evolutionary lineages because they are re- sulinids and other Foraminifera. Based on peated in unrelated lineages involving en- fusulinids, the unnamed white-colored lime- demic and immigrant groups. Yet, in each of stone which contains the fauna is dated as three cycles the morphological sequence is Late Artinskian-Early Guadalupian and is exactly repeated. I suspect that the repeated included in the Misellina claudiae zone species morphologies ofthe above genera are (Jones, Gobbett, and Kobayashi, 1966, p. not directly linked genetically to the North 328). As mentioned in the earlier reports, this American species but are convergent on them Tethyan fauna containing 92 gastropod and reflect similar environments in different species is, with the exception of the Lower parts of the Tethys. Getaway Limestone fauna of the Guadalupe In the case of the Malaysian species there Mountains, Texas, the richest known thus far are, in addition to some identical features of in the Permian. The fauna is composed of North American Pennsylvanian species, suf- both endemic and cosmopolitan species and ficient differences to recognize them as being has several unusual and important charac- distinct. For example, ifI should throw species teristics, such as the presence of the earliest from both areas together I would have no known nerineid, Prodiozoptyxis permiana trouble separating them. In critical features (new genus and species). The endemic forms such as the two spiral ribs defining the outer Loxosonia (new genus), Acrospira (new ge- whorl face, Orthonema rectimurum n. sp. is nus), and Sinozyga (new genus), have exag- identical to Orthonema salteri Meek and gerated morphological features, such as si- Worthen, 1860, yet in details of the second- phons, unknown in other members of the ary ornament such as the threads near the 1985 BATTEN: PERMIAN GASTROPODS 3 sutures, the two species are distinct. The Ma- phonal notch and features shared with Ju- laysian Orthonema nakazawai n. sp., 0. rec- rassic cerithids, including nearly vertical, timurum n. sp., 0. tirusum n. sp., and 0. straight growth lines. This represents a major knighti n. sp. have the following iterative systematics change. counterparts in the North American Penn- There is a serious systematics consequence sylvanian: 0. marvinwelleri Knight, 1934, 0. of this discovery-over 50 species of Car- liratum Sayre, 1930, 0. salteri Meek and boniferous and Permian pseudozygopleurids Worthen, 1860, and 0. inornatum Knight, have been assigned to this genus. They are 1934. Knight (1934) was the first and only clearly holostomous and firmly placed as dis- reviewer of the genus Orthonema which, at tinct members ofthis family. They are some- that time, was known only by 10 species from what skimilar to Paleostylus, namely in having the North American Pennsylvanian. Because vertical ribbing on flattened whorls and a species of the genus are represented by only high-spired many-whorled shell. This ribbing a single or few specimens at any locality, in- ranges from typical slightly curved pseudo- ter- and intraspecific variation remained un- zygopleurid ornament to the straight ribbing known. This study furnishes some introspec- seen in the cerithids. See page 11 for a more tion into this variation. detailed discussion. It is difficult to compare the composition Finally, I should point out that the recog- ofthe known Tethyan Permian faunas owing nition of a new genus of nerineids in the Pa- to the variation in time, ecology, and geo- leozoic Prodiozoptyxis may appear difficult graphic distribution. The physically closest to justify. However, the presence of apo- faunas are those described by Delpey (1942) morphic nerineid features, such as internal in Cambodia and by Yin (1983) and Wang folds, in this new genus suggests placement and Xi (1980) from Yunnan and Guizhou. in this family rather than in the campanilids The Cambodian fauna is almost identical to as discussed in the systematics of Prodiozop- the Malaysian fauna in many groups, but it tyxis. The nerineids undergo a rapid diver- lacks the diversity; unfortunately, Delpey did sification in the Jurassic and Cretaceous. not report the size of her samples and we When the first species appeared in the Ba- cannot compare frequencies. The Sosio and jocian, they already had parietal and colu- Tunisian faunas are quite similar and are re- mellar folds. Huddleston (1889, p. 497) re- lated to the Western Hemisphere faunas rath- ports 26 species from the Inferior Oolite and er than to the eastern Tethyan faunas. The these show a range from simple folds to rather Yunnan and Guizhou faunas lack many of complex infolding of the principal folds. the species represented in the Malaysian fau- The literature does not indicate the pres- na. Obviously, this difference is due to the ence of nerineids in the Triassic; however, more complete sampling of the Malaysian there is an undescribed species in the Norian Permian because ofthe unique preservation. fauna of Idaho (Seven Devils Formation). In The Salt Range fauna has a few conservative that species there is a single mid-whorl pa- bellerophontid genera and many endemic rietal fold. The shell is very high-spired with species, making comparisons difficult. flattened, slightly concave whorls much like One of the most important observations the common Jurassic genus Nerinea De- made in this study was an incurrent siphonal shays, 1827. The only real gap is the absence notch in Paleostylus as represented by the ofa nereinid in the Middle Triassic (none are Asian Tethyan species. In consequence, Pa- present in the St. Cassian fauna). leostylus senso stricto (the type species P. pu- I cannot judge from Huddleston's illustra- poides Mansuy, 1914 was described from tions whether all of the 26 species he de- Southeast Asia) must be removed from the scribed belong to Nerinea. I suspect they do Loxonematacea, which are exclusively ho- not because he was extremely conservative lostomous. The concept of Paleostylus as a in using generic determinations; for example, member of the loxonematid pseudozygo- he used Pleurotomaria where none of the pleurids was introduced by Knight in 1936. species are currently assigned to that genus. My conclusion is to place the genus in the In fact, I doubt that all of the species would Cerithiacea based on the presence of the si- be considered valid by today's standards. In 4 AMERICAN MUSEUM NOVITATES NO. 2829 any event, the illustrations show an interest- SYMBOLS AND ABBREVIATIONS ing sequence ofincreasingly complex internal The following abbreviations are used: AMNH folds. The Permian species falls well within American Museum of Natural History. H shell the range of that Inferior Oolite sequence. height. In most cases this measurement is esti- ACKNOWLEDGMENTS mated because the protoconch and earliest whorls are frequently broken off. The height is approxi- I again thank Dr. Derek J. Gobbett ofCam- mated by measuring the distance between the base bridge University (formerly ofthe University of the shell and the intersection of eye-piece go- of Malaysia) for sending me the specimens niometer cross-hairs, which are touching the later that form the basis of this study. Dr. Keiji whorls to measure the spiral angle, and are ex- Nakazawa ofKyoto University deserves spe- tended above the top of the shell at the position cial thanks for sending vital additional spec- of the tip of the protoconch. In many cases the protoconch height is not related to the logarithmic imens collected during the Southeastern Sci- progression ofthe teleoconch; it may be higher or entific Expedition made by members of the lower. W shell width. WhH whorl height. WhW Ozaka City and Kyoto universities. I thank whorl width. SpAng the spiral,angle. Mr. G. Robert Adlington and Mr. Peter Har- The numbers in parentheses in the synoptic clas- ries for the bulk of the photographs made of sification are the numbers of specimens of each poorly photogenic material. species studied. SYNOPTIC CLASSIFICATION Class Gastropoda Subclass Prosobranchia Order Archeogastropoda Suborder Murchisoniina Superfamily Murchisoniacea Koken, 1896 Family Murchisoniidae Koken, 1896 Genus Murchisonia d'Archiac and deVerneuil, 1841 Subgenus M. (Donaldospira) Batten, 1966 M. (D.) malaysia, new species ...... (21) Genus Stegocoelia Donald, 1892 Subgenus S. (Taosia) Girty, 1939 S. (T.) bengkaka, new species ...... (10) Genus Goniasma Tomlin, 1930 Goniasma, species ...... (1) Genus Cibecuia Winters, 1963 C. divarica, new species ...... (4) Genus Loxosonia, new genus L. hormotoma, new species ...... (6) L. zygopleuroides, new species ...... (3) Order Mesogastropoda Theile, 1925 Superfamily Loxonematacea Koken, 1889 Family Pseudozygopleuridae Knight, 1930 Genus Pseudozygopleura Knight, 1930 Subgenus P. (Pseudozygopleura) Knight, 1930 P. (P.) pleurozyga, new species ...... (8) P. (P.) obliqua, new species ...... (4) P. (P.) convexus, new species ...... (6) P. (P.) lirata, new species ...... (12) P. (P.), species ...... (2) Subgenus P. (Leptozyga) Knight, 1930 P. (L.) cf. venustus Hoare and Sturgeon, 1981 ...... (4) Subgenus P. (Stephanozyga) Knight, 1939 P. (S.), species ...... (1) 1985 BATTEN: PERMIAN GASTROPODS 5

Genus Hemizyga (Hemizyga) Girty, 1915 Hemizyga globosa, new species . (1) Subgenus H. (Plocezyga) Knight, 1930 H. (P.) serocorona, new species. (3) Genus Microptychis Longstaff, 1912 Microptychis permiana, new species. (1) Superfamily Loxonematacea, incertae sedis Genus Acrospira, new genus Acrospira cristata, new species. (9) A., species . (3) Superfamily Cerithiacea Fleming, 1822 Family Turritellidae Woodward, 1851 Genus Orthonema Meek and Worthen, 1862 0. rectimurum, new species. (18) 0. nakazawai, new species . (17) 0. tirusum, new species . (5) 0. knighti, new species . (41) Family Procerithiidae Cossmann, 1905 Genus Paleostylus Mansuy, 1914 P. pupoides Mansuy, 1914 . (37) P. dussaulti Mansuy, 1914 . (8) P. delpeyae, new species ...... (2) P. pseudozyga, new species . (7) Genus Protostylus Mansuy, 1914 Protostylus lantenoisi Mansuy, 1914. (5) Genus Sinuozyga, new genus S. ajoka, new species . (6) Family Coelostylinidae Cossmann, 1909 Genus Omphaloptychia Ammon, 1892 0. paleozoica, new species . (74) 0. cingulata, new species . (17) O., species . (15) Genus Trypanostylus Cossmann, 1895 T. triadicus Kittl, 1894 . (2) Superfamily Subulitacea Lindstrom, 1884 Family Meekospiridae Knight, 1956 Genus Meekospira Ulrich, 1897 M. melanoides, new species . (18) M. ligoni, new species . (17) Family Subulitidae Lindstrom, 1884 Subfamily Soleniscinae Wenz, 1938 Genus Strobeus deKoninck, 1881 Strobeus, species. (6) Genus Soleniscus Meek and Worthen, 1861 S. elegans Gemmellaro, 1889. (13) Genus Cylindritopsis Gemmellaro, 1889 Cylindritopsis, species. (3) Subfamily Subulitinae Lindstrom, 1884 Genus Ceraunocochlis Knight, 1931 Ceraunocochlis, species. (1) Superfamily Nerineacea Zittel, 1873 Family Nerineidae Zittel, 1873 Genus Prodiozoptyxis, new genus 6 AMERICAN MUSEUM NOVITATES NO. 2829

P. permiana, new species ...... (37) P., species ...... (10) Order Opisthobranchia Superfamily Pyramidellacea d'Orbigny, 1840 Family Streptacididae Knight, 1931 Genus Donaldina Knight, 1931 Donaldina, species ...... (3) Genus Streptacis Meek, 1872 Streptacis, species ...... (1) Genus Spirocyclina Kittl, 1894 Spirocyclina, species ...... (1) Addendum Suborder Trochina Superfamily Trochacea Rafinesque, 1815 Family Turbinidae Rafinesque, 1815 Subfamily Liotiinae Adams and Adams, 1854 Genus Eucycloscala Cossmann, 1893 E. asiatica, new species ...... (9) E. medionodosus, new species ...... (31)...... Total specimens studied (501)

ORDER ARCHEOGASTROPODA archeogastropods and the mesogastropods are polyphyletic since the trochids also appear to SUBORDER MURCHISONIINA have given rise to a number of mesogastro- COX AND KNIGHT, 1960 pod families. DISCUSSION: Cox and Knight (1960) con- FAMILY MURCHISONIIDAE KOKEN, 1896 structed this suborder because of some mor- phological differences between the pleuroto- DISCUSSION: Knight et al. (1960) have dis- marians and the high-spired murchisoniids cussed the morphological range ofthe genera and also because they recognized that the included in this family. They concluded that murchisoniids were ancestral to the loxo- one of the most important morphological nematids and the cerithids and hence distinct complexes in the family was that ofthe place- from the pleurotomarians which they resem- ment of the selenizone. For example, Gon- ble. Some genera have incipient abapical iasma Tomlin, 1930 has a selenizone im- apertural canals whereas the pleurotomarians mediately underlying a centrally located keel are holostomous. They have a three-layered or periphery, while S. (Hypergonia) Donald, crossed-lamellar wall as opposed to the na- 1892 has a selenizone immediately overly- croprismatic wall of all known pleurotomar- ing the periphery. M. (Donaldospira) Batten, ians, which serves, in part, to separate the 1966 has a selenizone on the periphery so two suborders. The crossed-lamellar struc- that half of it is above and half below, re- ture is ofthe form ofshort first-order lamellae sulting in a convex selenizone. The family is with opposing sets of second-order lamellae relatively conservative with a low speciation at relatively low angles. This type ofstructure rate in the Upper Paleozoic. Inter- and in- is commonly encountered in many archaeo- trapopulation variation is low. Only a few gastropod groups, such as the euomphalids, specimens of a single species occur in most bellerophontids, and neritaceans (see Batten, faunas. At AMNH locality 512 in the lower 1984, p. 12). The centrally located selenizone Getaway limestone at Pine Springs, Guada- indicates that they retained paired aspido- lupe Mountains, Texas, the richest Permian branch ctenidia and rhipidoglossate radula. fauna yet to be found has about 100 speci- With this group as the ancestral stock of two mens belonging to five species. The Lee Mine mesogastropod superfamilies (Cerithiacea fauna has 43 specimens belonging to six and Loxonematacea), it is clear that both the species. 1985 BATTEN: PERMIAN GASTROPODS 7

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-0, 41,:1* _.f 1 :.A& 2 3V 4 FIGS. 1-4. Murchisonia (Donaldospira) malaysia. 1. Holotype, AMNH 42782, side view, x 1.5. 2. Paratype, AMNH 42784, side view with low-angled lighting to show the selenizone bisected by the angular periphery; this is a more rounded variant, x 1.5 mm. 3. Paratype, AMNH 42785, side view of a turriculate variant, x 2. 4. Paratype, AMNH 42788, apertural view of a fragment showing the well- developed siphonal trough; the unorthodox side lighting from the right is necessary to cast a shadow on the trough, x 2.

Murchisonia (Donaldospira) Batten, 1966 the position of the selenizone, along with its associated characteristics, can be used to dis- TYPE SPECIES: Murchisonia pertusa de- tinguish them. Koninck, 1883, p. 15, pl. 33, figs. 50, 5 1; from the v3b (=D2) zone at Vise, Belgium. DISCUSSION: Batten (1966, p. 68) recog- Murchisonia (Donaldospira) malaysia, nized seven species in the Devonian and new species Lower Carboniferous (confined to western Figures 1-4 Europe) as belonging to this subgenus; thus DIAGNOSIS: Large, turritelliform snails with far it has not been identified in the Pennsyl- a convex selenizone forming the periphery; vanian or elsewhere in the Permian. The early whorls angulate with prominent pe- preservation of these specimens is such that ripheries, older whorls rounded with pe- little surface detail, such as growth lines, is ripheries less well defined; selenizone orna- available for analysis. For example, the cu- mented with fine spiral threads and with rious patterned punctation found in the type collabral nodes; upper whorl surface convex, species M. (D.) pertusa was not observable. flat, or slightly concave with two or more It is possible that some of the silicified spec- spiral threads; alveozone flat to concave with imens identified as Stegocoelia (Donald, one medial spiral thread; basal margin 1889) in the Permian of west Texas may be marked by a prominent spiral rib; base with Donaldospira, but the coarse silicification has one or more spiral threads or ribs. Siphonal destroyed fine growth lines in every specimen groove well developed. examined, hence preventing proper taxo- DISCUSSION: The population sample aver- nomic assignment. The shell shapes, position aged about twice that of the type species M. of the periphery on the whorl, and other fea- (D. ) pertusa. It differs from Murchisonia dus- tures are nearly identical in both groups. Only saulti Mansuy, 1914 in having a broad, flat 8 AMERICAN MUSEUM NOVITATES NO. 2829 upper whorl surface, with a more rapid whorl base is somewhat flattened and minutely expansion rate. The fine spiral ornament is phaneromphalus. The columellar lip has an more numerous on the base and has a well- elongate, centrally located callus which developed siphonal groove. The first known slightly reflexes the lip. A spiral thread is lo- specimen of the subgenus to have the aper- cated in the middle ofthe base; no other spi- ture preserved is shown in figure 4. The si- ral ornament was observed. phonal canal is more tapered and narrow than DISCUSSION: This specimen is virtually in any related group. The nature of the se- identical to S. (T.) crenulata Girty, 1939, a lenizone is unknown in M. (D.) dussaulti. I ubiquitous species in the Pennsylvanian and have not recognized the species in other fau- Permian ofthe Western Hemisphere. It is not nas. preserved well so that the growth lines and The early whorls of this species are turric- perhaps fine ornament are not observable. ulate with prominently developed peripher- However, the basic morphology cannot be ies; older whorls are more rounded with the separated from that of S. (T.) crenulata. To periphery less well defined. This feature var- the best of my knowledge, this subgenus is ies among specimens; some are more turric- thus far unknown in the Eastern Hemisphere ulate throughout ontogeny. Twenty-one spec- with the exception ofa specimen I have iden- imens. tified from the highest Permian in Greece. MEASUREMENTS: Holotype AMNH 42782, SPECIMENS: Ten; two other specimens are H 47.1 (e) mm,W 17.1 mm, SpAng 130; para- from the Sin Thong Hing Mine near Kampar type AMNH 42783, H 40.2 (e) mm, W 13.8 and marked "Upper Paleozoic" on the De- mm, SpAng 14.50; paratype AMNH 42784, partment ofGeology label (University ofMa- H 58.2 (e) mm, W 18.7 mm, SpAng 16.00; laysia). paratype AMNH 42785, H 59.2 (e) mm, W MEASUREMENTS: Holotype AMNH 42789, 20.2 mm, SpAng 16.00; paratype AMNH H 8.0 mm, W 2.9 mm, WhH 1.5 mm, WhW 42786, H 51.6 (e) mm, W 20.6 mm, SpAng 2.9 mm; large paratype AMNH 42790, H 19.00; paratype AMNH 42787, H 61.9 (e) 21.0 mm, W 8.9 mm, WhH 4.2 mm, WhW mm, W 28.0 mm, SpAng 14.50; paratype 8.9 mm. AMNH 42788, H 85.1 (e) mm, W 26.8 mm, ETYMOLOGY: Bengkak, Malay for swelling. SpAng 14.50. (e) = estimated height, the spec- imen is broken. Stegocoelia (Stegocoelia) sp. Figure 7 Stegocoelia (Taosia) Girty, 1939 DESCRIPTION: High-spired, turriculate shells TYPE SPECIES: S. (T.) copei (White), 1881, with the periphery below midwhorl; the up- p. 31, pl. 3, figs. 1Oa, b. per selenizone margin forms the periphery; DISCUSSION: This subgenus is widely dis- the lower selenizone margin defines a narrow, tributed in upper Paleozoic rocks but only flat selenizone. The upper whorl face is slight- one or several specimens occur at any one ly convexoconcave; the lower whorl face is locality. about one-third the size of the upper whorl face and the sutures are sharply defined. Stegocoelia (Taosia) bengkaka, new species DISCUSSION: There is only a single speci- Figures 5-6 men ofthis taxon and the preservation ofthe turricu- shell is so poor that nothing can be discussed DIAGNOSIS: Relative high-spired, ofthe ornament or other morphology except late shells with a strongly noded periphery the selenizone and associated characters. One located low on the whorl. Sutures are located specimen. just below the periphery. A well-defined spi- MEASUREMENTS: AMNH 42791, H 14.7 ral cord is adjacent to the upper suture. There mm, may be fine collabral threads developed on W 5.7 mm, SpAng 290. the periphery and upper spiral cord. The se- 1930 lenizone is located midway between the su- Genus Goniasma Tomlin, ture and the periphery on the flattened or TYPE SPECIES: Murchisonia lasallensis slightly concavoconvex upper whorl face. The Worthen, 1930. 1985 BATTEN: PERMLAN GASTROPODS 9

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d*- t. ' .'' 7 _gH... 8 6 FIGS. 5-8. 5. Stegocoelia (Taosia) bengkaka. Holotype, AMNH 42789, side view, x 6.6. 6. Paratype, AMNH 42790, apertural view, note strong rib on periphery, x 2.5. 7. Stegocoelia (Stegocoelia) sp., AMNH 42791, side view of fragment, x 3. 8. Goniasma sp., AMNH 42792, side view, x 4.

DISCUSSION: As far as I am aware this is DISCUSSION: C. divarica, new species, is the the first report of the genus outside North only other species ofthe genus. Cibecuia pre- America. The type species is from the Penn- viously has been known only from the Perm- sylvanian ofIllinois. Three species have been ian Kaibab Formation ofeastern Arizona and described from the Pennsylvanian and Perm- from a single fragment from the Bone Spring ian of the western United States. The genus Formation, Sierra Diablo Mountains ofwest- is based on the selenizone location; it is just ern Texas. This is yet another example of a below the periphery, with the upper seleni- disjunct distribution involving the south- zone margin forming the periphery. western United States and Malaysia.

Goniasma sp. Cibecuia divarica, new species Figure 8 Figure 9 DISCUSSION: One specimen in this fauna is DIAGNOSIS: shells with whorl a member of this genus. The preser- High-spired clearly profile smooth and whorl face flat; selenizone vation is too poor to describe the species above the lower selenizone characteristics. The whorls are evenly convex just midwhorl, marks its selenizone mar- and there are several spiral cords. The upper margin midwhorl; margin ofthe selenizone marks the periphery. gins impressed; base delineated by a spiral numerous nodes on The selenizone is strongly concave and the keel; collabral, elongated margins rounded and well developed. The upper whorl surface adjacent to the suture; collabral threads are formed on the outer upper whorl surface has a well-developed spi- mar- ral cord in the center. The whorls embrace whorl surface from the lower selenizone to the basal whorls embrace on a spiral cord which forms the edge of the gin periphery; base. The base is narrowly phaneromphalus. just below the spiral keel; base ornamented One specimen. by two equally spaced and developed ribs; base is flat to slightly concave. MEASUREMENTS: AMNH 42792, H 12.3 DISCUSSION: This species is remarkably mm, W 5.6 mm, SpAng 210. similar to the type and the previously only known species, C. cedarensis Winters, 1963. Cibecuia Winters, 1963 It differs in that the whorl profile is less TYPE SPECIES: C. cedarensis Winters, 1963, stepped than that of C. cedarensis because p. 38, pl. 4, figs. 5a-6b. the lower part of the whorl is more fully de- 10 AMERICAN MUSEUM NOVITATES NO. 2829

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k.1. .1 9 0 9b 9c FIG. 9. Cibecuia divarica, holotype, AMNH 42793; (a) side view, x4.5; (b) apertural view, x4.5, low-angled lighting was used to bring out details of selenizone and subsutural ornament; (c) side view, x 4.5 with low-angle light to bring out details. veloped than that of the whorl near the su- DISCUSSION: The shell shape and the ture. The sutural nodes are more numerous collabral ornament of this genus is highly and finer than that of the type species. In all convergent on the fusiform shells of the lox- other features the two species are identical. onematids, but the presence of a well-devel- Four specimens. oped selenizone and siphonal canal is typical MEASUREMENTS: Holotype AMNH 42793, of the murchisonids. Loxisonia differs from H 15.00 mm, W 4.3 mm, WhH 3.0 mm., the murchisonids Aclisina and Stegocoelia in WhW 4.3 mm, SpAng 120; paratype AMNH having the selenizone much higher on the 42794, H 8.00 mm, W 3.5 mm, WhH 2.1 whorl (near the suture), in having a siphonal mm, WhW 3.5 mm, SpAng 12°; paratype canal, and by having rounded whorls with no AMNH 42795, WhH 3.2 mm, WhW 5.5 mm. spiral ornament, keels, or sharply defined pe- ETYMOLOGY: Divaricatus, Lat. spread. riphery. I have found specimens ofthe genus illustrated in an unpublished manuscript il- Loxisonia, New Genus lustration of the Permian of Crimea written by 0. Toumansky in the late 1930s or early DIAGNOSIS: Moderately high-spired, fusi- 1940s. form shells with a selenizone located near the ETYMOLOGY: Loxos, Gk. sloping; sohn, Gk. suture on the upper whorl surface; whorls related to. elongate, rounded; sutures embrace whorls below the rounded peripheral region; seleni- Loxisonia hormotoma, new species zone margins sharply defined by narrow Figures 10-11 grooves; ornament, ifpresent, consists ofcol- labral ribs or threads, is more developed on DIAGNOSIS: A moderately high-spired shell the outer whorl face; columellar lip reflexed with elongate, rounded to somewhat flat- into a narrow funicle or callus; siphonal canal tened whorls with a relatively narrow seleni- is short and shallow. zone, apparently without margins, located 1985 BATTEN: PERMIAN GASTROPODS I1I near the suture; the suture is sharp and deeply SPECIMENS: Three; all specimens are bro- impressed; ornament is faintly developed ken so that only the last two whorls are pres- collabral threads or ribs which are barely ent. raised above the whorl surface and are round- MEASUREMENTS: Holotype AMNH 42800, ed to somewhat flattened; whorls embrace H 7.0 mm, W 3.2 mm, SpAng 140; paratype well below the rounded periphery located at AMNH 42801, H 6.0 mm, W 2.8 mm, SpAng midwhorl; the base is rounded and contin- 120; paratype AMNH 42802, H 12.0 mm, W uous with the outer whorl surface; siphonal 6.0 mm, SpAng 200. canal well developed and narrow; hemiom- ETYMOLOGY: Zygon, Gr. yoke; pleuron, Gr. phalus to anomphalus. rib. DISCUSSION: This species differs from L. zygopleuroides in being an order of magni- ORDER MESOGASTROPODA tude larger and with the ornament faintly de- THIELE, 1925 veloped; the shell shape is more variable, ranging from a compressed form with more DISCUSSION: Extinct members ofthis order, tightly coiled, flattened whorls to a loosely particularly of the Paleozoic, are recognized coiled form with more inflated whorls and a by comparison of shell morphology with ex- rounded base. tant groups. This is reinforced by assuming SPECIMENS: Six; all specimens are broken that extinct members of higher taxa share but with from five to nine whorls preserved. ordinal features with their living sister groups. MEASUREMENTS: Holotype AMNH 42796, Occasionally, a living relict ofan extinct group H 10.5 mm, W 3.55 mm, SpAng 210; para- is discovered such as the monoplacophoran type AMNH 42797, H 5.1 mm, W 2.01 mm, Neopilina. Abyssochrysos melanoides Tom- SpAng 170; paratype AMNH 42798, H 4.20 lin, 1927, a living species, was found which mm, W 1.53 mm, SpAng 160; paratype may possibly be a member of the otherwise AMNH 42799, H 3.20 mm, W 1.10 mm, extinct superfamily Loxonematacea Koken, SpAng 140. 1889 (Houbrick, 1979). Thiele (1931), Wenz ETYMOLOGY: Hormos, Gk. chain; tomas, (1938), Taylor and Sohl (1962), and others Gk. cut. had placed the family Abyssochrysidae in the superfamily Cerithiacea. The Cerithiacea Loxisonia zygopleuroides, new species have living representatives so that the taxo- Figure 12 nomic placement within the living mesogas- DIAGNOSIS: Shell with smooth, rounded tropods is secure. whorls except at the selenizone and upper whorl surface above it which are flattened FAMILY PSEUDOZYGOPLEURIDAE relative to the rest ofthe whorl; the selenizone KNIGHT, 1930 is flat to concave with asymmetric lunulae; DISCUSSION: This family and others of the the selenizone is located higher on the whorl; Loxonematacea have been suggested as me- ornament is strongly developed collabral ribs sogastropods based on the recognition of the which are rounded between the suture and living Abyssochrysos melanoides Tomlin, the selenizone; the ribs are strongly arcuate 1927 as being loxonematid rather than cer- on the lower whorl face forming a deep sinus ithiacean (Houbrick, 1979) as previously be- adjacent to the selenizone; ornament absent lieved. This is based on the presence ofclosed on the base; siphonal notch short; anom- pallial gonoducts and a large penislike organ phalus. which excludes it from the cerithids which DISCUSSION: This species is strongly con- have open pallial gonoducts and an aphallic vergent on the pseudozygopleurid P. sinuosia condition in males (Houbrick, 1979, pp. 14- Knight, 1930 in the shape of the shell, the 15). whorl profile, the collabral ribs, the shape of There is a persistent recognition problem the base, and the reflexed columellar lip. It involving the families Zygopleuridae Wenz, differs from the pseudozygopleurids in hav- 1938, Pseudozygopleuridae Knight, 1931, and ing a well-developed selenizone which inter- Palaeozygopleuridae Horn, 1955 (see Hoare rupts the collabral ribs above the midwhorl. and Sturgeon 1978, pp. 850-852 for further 12 AMERICAN MUSEUM NOVITATES NO. 2829 discussion). Knight (1930, pp. 8-12) recog- in Tunisia, Sicily, or Greece. Even in the Tex- nized the Paleozoic pseudozygopleurids as as Permian it is poorly represented. being distinct from the Mesozoic zygopleu- rids on the basis that the first four whorls are P. (Pseudozygopleura) pleurozyga, ornamented by features differing from those new species of an adult shell. Hornm (1955) discovered Figure 13 several genera in the Devonian of Czecho- slovakia which are identical to adult pseu- DIAGNOSIS: Shell high-spired with very dozygopleurid shells except that the early weak spiral ornament on the periphery and whorls are unornamented. Subsequent to with slightly sigmoid collabral ribbing that is these finds, other species in the late Paleozoic well developed across the whorl. The convex have been found to have unornamented early whorl profile has a slight depression forming whorls as well, and these have been called by a troughjust below the suture but is separated various pseudozygopleurid generic names from it by poorly developed interference (Hoare, 1980). The problem is that without nodes on the ribs adjacent to the suture. Rib- the early whorls preserved (the usual state), bing is weakly developed in the trough. A it is impossible to place shells in either fam- weakly formed periphery is present on the ily. In addition, the specter ofhomeomorphy lower whorl face. The base is smoothly is present. I see no reason why this problem rounded. The ornament is evenly developed could not be simply resolved by broadening on all whorls except for the nuclear whorls the definition of the pseudozygopleurids to and the base where it is subdued. include the forms with smooth early whorls, DISCUSSION: The presence of a shallow hence synonymizing the palaeozygopleurids. troughjust beneath the suture is an important In the case of the Mesozoic zygopleurids, and unique feature. This and the related fea- the main phyletic path has evolved away from ture ofthe disappearance of ornament in the the pseudozygopleurid bauplan. Only Kato- trough make this a distinctive species unre- sira Koken, 1892, Zygopleura Koken, 1892, lated to any others described. There is vari- and Tyrsoecus Kittl, 1892 are similar to the ation in the development ofthe collabral rib- Paleozoic families. Hence, there is merit to bing from evenly formed ribs to those that retaining the zygopleurids as a distinct entity. are well developed only at the suture and on I would suggest that the above three genera the outer portions of the periphery. In some could be placed in the Pseudozygopleuridae, specimens, the trough is very faint and only but it is beyond the scope of this study to a slight weakening of the ribbing marks the make such revisions. position of the trough. Eight specimens. MEASUREMENTS: Holotype AMNH 42803, Genus Pseudozygopleura Knight, 1930 H 20.2 mm,W 8.4 mm, WhH 4.1 mm, WhW TYPE SPECIES: Loxonema semicostatum 7.3 mm; paratype AMNH 42804, broken Meek, 1872. WhH 3.7 mm, WhW 6.7 mm. DISCUSSION: Knight (1930) originally set ETYMOLOGY: Pleura, Gk. ribs; zygon, Gk. up four subgenera based on shell shape and yoke. ornament: P. (Stephanozyga), P. (Pseudo- zygopleura), P. (Leptozyga), and P. (Progo- P. (Pseudozygopleura) obliqua, new species zyga). Since that time two major revisions of Figure 14 the genus were effected by Knight et al. (1960) DIAGNOSIS: High-spired shell with mod- and Hoare and Sturgeon (1981). The former erately, but evenly, inflated whorls, with the placed the four subgenera in the genus Pa- upper whorl surface slightly flattened. The leostylus. This cannot be supported by cur- collabral ribbing is straight but oblique to the rent knowledge as documented by Hoare and suture. Each whorl has ribbing aligned with Sturgeon (1981, p. 572). ribbing in adjacent whorls. Ribs are fully de- Pseudozygopleura is not found in most fau- veloped from suture to suture. Ribbing may nas of the Tethyan Permian and, hence, we be fine or coarse. Ornament weakens during know little of its distribution. It is found in growth so that earlier whorls are more strong- Crimean Permian but is curiously absent from ly ornamented, the final two whorls with faint, the larger, better known faunas such as those rounded ribs or growth lines only. The pe- 1985 BATTEN: PERMIAN GASTROPODS 13

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11 -. FIGS. 10-12. 10. Loxosonia hormotoma, holotype, AMNH 42796, apertural view, note siphonal trough, xO.5. 11. Paratype, AMNH 42799, side view of fragment, xl. 12. Loxosonia zygopleura, holotype, AMNH 42800, apertural view, x 8. riphery is low on the whorls and the base is with a midwhorl periphery and vertical, somewhat flattened. The whorls embracejust straight collabral ribs. The ribbing is well de- below the periphery. The ornament on the veloped and sharply delimited with full de- base consists of faint collabral threads. velopment across the whorl. Weakly rein- DIscuSSION: Except for P. (P.) semicosta- forced interference nodes are found on tum (Meek), 1872 the type species, and P. collabral ribs immediately adjacent to the su- (P.) terebra Knight, 1930, most species have ture. The sutures are sharply defined and deep. the ornament fully developed throughout on- Whorls embrace on the edge of an ill-defined togeny. Some specimens may lose ornament low periphery. The base is evenly rounded. in species that normally retain full ornamen- DISCUSSION: The variation noted in this tation. I doubt that it has phyletic signifi- species involves the whorl expansion rate cance. The ribbing in this species and the very causing some specimens to be more trochoid evenly developed whorl profile is quite dis- or lower spired than most other species in tinctive. It most closely resembles P. (P.) mu- shell shape. Some specimens have somewhat cronotus Hoare and Sturgeon, 1981, in the finer ribs than that of the holotype. This obtuse ribbing and evenly formed ribs; it dif- species resembles P. (P.) inornata Knight, fers from that species by having more inflated 1930 in the development of the ribbing, but whorls and straight ribs. Four specimens. the whorl profile is more inflated and the base MEASUREMENTS: Holotype AMNH 42805, more rounded. It is similar to P. (P.) pugonis H24.0mm,W8.4mm,WhH3.8 mm,WhW Hoare and Sturgeon (1981) in the shape of 7.2 mm, SpAng 180; paratype AMNH 42806, the whorl but has finer ribbing. There does H 15.2 mm,W 2.9 mm, WhH 2.9 mm, WhW not seem to be any other described species 6.2 mm, SpAng 180. outside North America that is similar to this ETYMOLOGY: Obliquus, Lat. slanting. one. Six specimens. MEASUREMENTS: Holotype AMNH 42807, P. (Pseudozygopleura) convexus, H 8.1 mm, W 3.9 mm, WhH 2.2 mm, WhW new species 3.9 mm, SpAng 260; paratype AMNH 42808, Figure 15 broken WhH 3.6 mm, WhW 6.2 mm, SpAng DIAGNOSIS: The shell is moderately high- 240. spired with an evenly convex whorl profile ETYMOLOGY: Convexus, Lat. arched. 14 AMERICAN MUSEUM NOVITATES NO. 2829

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i 1.4 )i t i.. II I ;e".1$ 1. . CO .- .,,3. " .;0*. Am,., - -, I.- xmlw- " 144r-l 17I '1% 1 5 16 14 FIGS. 13-17. Pseudozygopleura (Pseudozygopleura). 13. P. (P.) pleurozyga, holotype, AMNH 42803, side view, x 2. 14. P. (P.) obliqua, holotype, AMNH 42805, apertural view (aperture broken), x 2. 15. P. (P.) convexus, holotype, AMNH 42807, side view, x 4.5. 16. P. (P.) lirata, holotype, AMNH 42809, side view, x 8. 17. P. (P.) sp., AMNH 42811, side view, x 12.5.

P. (Pseudozygopleura) lirata, new species H 4.6 mm,W 2.3 mm, WhH 0.9 mm, WhW Figure 16 1.7 mm, SpAng 160. ETYMOLOGY: Lira, Lat. for ridge. DIAGNOSIS: A small, turriculate shell with flattened whorl profiles and flattened base P. (Pseudozygopleura), species with a lira marking the lower margin of the Figure 17 outer whorl face. The outer whorl face slopes outward to a sharply defined periphery low DISCUSSION: Two specimens are sufficient- on the whorl. Collabral ribs are straight, mod- ly distinct to include as a separate species. erately well developed, and nearly vertical The whorl profile is evenly inflated with a flattened outer whorl face with well-devel- over the whorl face. Ornament is uniformly oped, slightly arched, collabral ribs which are developed over the entire shell. The suture distinct in being fully developed from suture is sharply defined but shallow. The base is to the base. The suture is deeply incised. None flatly rounded with reduced to absent orna- of the other species have this unique rib de- ment. velopment. Since only the illustrated speci- DISCUSSION: The most distinguishing fea- men is well preserved, I have refrained from ture of this species is the flattened base de- formalizing this species. Two specimens. limited by a lira which forms the boundary MEASUREMENTS: AMNH 42811, H 3.7 mm, between the base and the outer whorl face. W 1.7 mm, SpAng 380. To my knowledge, this is a unique feature to the genus. There is some variation in the de- P. (Leptozyga) Knight, 1930 velopment of the collabral ribs from numer- TYPE SPECIES: P. (Leptozyga) minuta ous fine ribs to fewer coarser ribs. The whorl Knight, 1930, p. 63, pl. 4, fig. 2. profile tends to be flattened but may be evenly convex and sloping toward the low-lying pe- P. (Leptozyga) cf. venustus riphery. The species somewhat resembles Hoare and Sturgeon, 1981 several flat-based species described by Hoare Figure 18 and Sturgeon (1981) such as P. (P.) gradatus and base but P. Leptozyga venustus Hoare and Sturgeon, 1981, in the flattened whorl profile p. pl. figs. 13-14. differs in having a basal peripheral lira and 584, 1, an unornamented base. Twelve specimens. DISCUSSION: Several specimens are so close MEASUREMENTS: Holotype AMNH 42809, in morphology to this species that I can detect H 4.8 mm, W 1.9 mm, WhH 0.8 mm, WhW no important differences between them. They 1.7 mm, SpAng 180; paratype AMNH 42810, show the same evenly convex whorl profiles 1985 BATTEN: PERMIAN GASTROPODS 15 and fine, numerous collabral threads. The base is subdued on the base and it is cryptom- is rounded and anomphalus. One tiny spec- phalus. It resembles the Silurian Auriptygma imen has coarser threads than those illus- Perner, 1903 in the shape of the aperture, trated by Hoare and Sturgeon. Four speci- ornament pattern, and thinness of the shell. mens. It differs from most species of Hemizyga in MEASUREMENTS: AMNH 42812, H 11.5 lacking revolving ornament. However, the mm, W 4.7 mm, WhH 1.9 mm, WhW 3.0 earliest whorls are not preserved so that it is mm, SpAng 2.50. possible that there might be revolving or- nament on the earlier whorls as in some P. (Stephanozyga) Knight, 1930 species. I have found no other species similar to this specimen. Two specimens. TYPE SPECIES: P. (Stephanozyga) nodosa MEASUREMENTS: Holotype AMNH 42814, (Girty), 1915. H 22.4 mm, W 13.6 mm, SpAng 340; para- type AMNH 42815, H 22.4 mm, W 13.5 P. (Stephanozyga) sp. mm, SpAng 440. Figure 19 DISCUSSION: A single, poorly preserved Hemizyga (Plocezyga) Knight, 1930 specimen is probably a member of this sub- TYPE SPECIES: Plocezyga corona Knight, genus. It resembles the type species in having 1930. a concavo-convex whorl profile with a very DISCUSSION: Knight considered this sub- low periphery. Suture embrace beneath the genus as belonging to Hemizyga based on the periphery. The periphery is noded with col- presence of spiral ornament, an arcuate col- labral ribs which disappear just above the umella, and a subcircular whorl cross section. peripheral ridge. Sutures are shallow and Hoare and Sturgeon (1981, p. 171) discov- sharply defined. The subgenus is very rarely encountered, known only from a few hori- ered that the genus Hemizyga does not have zons in the Pennsylvanian Des Moines Series a pseudozygopleurid embryonic whorl, based in the Mississippi Valley. One specimen. on examination of the type H. (H.) elegans MEASUREMENTS: AMNH 42813, H 4.6 mm, (Girty), 1915. They defined the new genus W 2.6 mm, WhH 1.1 mm, WhW 2.3 mm, Gamizyga to include all species having pseu- SpAng 220. dozygopleurid embryonic whorls and having the characteristic spiral ornament identical Hemizyga (Hemizyga) Girty, 1915 to that ofHemizyga, placing Plocezyga in the new genus plus 18 new species. They did not TYPE SPECIES: H. elegans Girty, 1915. specify the nature of the embryonic whorls did in the original descriptions). Hemizyga globosa, new species (nor Knight Figures 20a, b Again, a dual system does not seem logical. I will continue to be conservative, believing DIAGNOSIS: Moderately high-spired forms that the presence or absence of the intricate with globose whorls with numerous fine, embryonic whorls is not sufficient to con- evenly developed collabral ribbing on all struct a dual classification and will retain this whorls. Sutures are sharply defined and rel- subgenus in Hemizyga. atively deep. A sutural ramp is narrow and slanted toward the suture. The early whorls H. (Plocezyga) serocorona, new species have coarse ribs which become finer in later Figures 21a, b whorls. The adult whorl may be unorna- mented. Whorls embrace well below the DIAGNOSIS: Trochiform shells with strong- rounded periphery located at midwhorl. ly formed collabral ribs with faint spiral DISCUSSION: Two well-preserved speci- threads which appear to be confined to the mens with inflated, convex whorls have nu- whorl surface between the ribs, but this may merous, rounded collabral ribs; are fine ex- be a matter of preservation. The ribs are cept for the unornamented adult whorl. The evenly developed over the whorl surface ex- shell is thin and the aperture is holostomous. cept immediately adjacent to the suture. The columellar lip is straight. The ornament Whorls embracejust below the periphery. The 16 AMERICAN MUSEUM NOVITATES NO. 2829

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22 FIGS. 18-22. 18. P. (Leptozyga) cf. venustus Hoare and Sturgeon, 1981, AMNH 42812, oblique side view, x 12.5. 19. P. (Stephanozyga) sp., AMNH 42813, side view, x 7. 20. Hemizyga (Hemizyga) globosa; (a) holotype, AMNH 42814, oblique side view, x 2.5; (b) paratype, AMNH 42815, side view of adult whorl in low lighting to show lack of ornament, x 3. 21. Hemizyga (Plocezyga) serocorona; (a) holotype, AMNH 42816, oblique basal view, note open umbilicus, x 7; (b) paratype, AMNH 42817, side view lightly coated to show lack of ornament or ribs, x 8. 22. Microptychis permiana, holotype, AMNH 42818, side view with low lighting to show the sinus just under the subsutural nodes, x 4.5. base is somewhat flattened and phanerom- H 4.4 mm, W 2.6 mm, SpAng 330; paratype phalus. The columella is arcuate. The shell is AMNH 42817, H 30.0 mm, W 22.5 mm, relatively thin and the sutures are sharply SpAng 420. incised. ETYMOLOGY: Serus, Lat. late; corona, Lat. DISCUSSION: The whorl profile varies from crown. globose to somewhat flattened. The shell shape ranges from low trochiform to almost 1912 turbinform. This species comes closest to the Microptychis Longstaff, type species H. (Plocezyga) corona in the rel- TYPE SPECIES: Microptychis wrighti Long- ative intensity of the collabral ribs, the shell staff, 1912, p. 307, pl. 30, figs. 6a, b. shape, and whorl profile. Three specimens. DISCUSSION: This is the first described MEASUREMENTS: Holotype AMNH 42816, species of the genus in the Permian. It is rel- 1985 BATTEN: PERMIAN GASTROPODS 17

whorls more inflated, with weaker, more elongate collabral nodes and no spiral thread adjacent to the suture. The sinus is deeper than that of the Crimean species. There ap- pear to be several species in the Crimean Permian. This species resembles M. subcon- stricta (DeKoninck), 1881 from the European Carboniferous in shell shape and whorl pro- file but differs in having coarser ribbing and a deeper sinus. This beautifully preserved fragment is so distinctive that there is no . 7 < 4~~2 question that it represents a new Permian 23 species of the genus. One specimen. MEASUREMENTS: AMNH 42818, H 2.7 mm, FIGS. 23-24. 23. Acrospira cristata, holotype, W 4.3 mm. AMNH 42819, side view, note varix formed at ETYMOLOGY: Permian, a geological system. the outer lip, x 10.5. 24. Acrospira sp., AMNH 42821, side view, x 10.5. SUPERFAMILY LOXONEMATACEA, INCERTAE SEDIS atively common in the European Lower Car- DISCUSSION: A new genus, Acrospira, which boniferous but rare in the Pennsylvanian. The is almost identical in external morphology to Crimean species has strong noding adjacent Procerithiopsis Mansuy, 1914, but lacks col- to the suture, identical to that of Paleostylus umellar folds, is assigned to the loxonema- suggesting that it is a transitional form be- tids. The dominance of collabral ribbing in tween the two genera. It helps to clarify the this superfamily, along with a high-spired relationship between them and the pseudo- shell, has influenced workers to assign species zygopleurids which lack a sinus. Hoare (198 1, to this group even though, I suspect, they do p. 1039) described the new genus Anozyga to not belong. A case in point was the accep- include forms which he described as being tance for many years ofPaleostylus as a mem- Microptychis -like with pseudozygopleurid ber, although clearly it belongs to Cerithiacea embryonic whorls. His illustrations seem to because it is siphonate. In any event, I too indicate that it lacks a sinus and has fewer, am influenced by the presence of collabral more inflated whorls than Microptychis sensu ribbing in Acrospira and place it in this group. stricto. Acrospira, New Genus new species Microptychis permiana, DIAGNOSIS: Trochoid shells with collabral Figure 22 ribbing and adult varix. The shell is mod- DIAGNOSIS: High-spired shells with slightly erately high spired with about seven whorls. inflated whorls with elongate collabral nodes The whorl profile is evenly convex with well- immediately adjacent to the suture. Sutures developed collabral ribbing which is more sharply impressed. Whorls embrace just be- fully developed in the broad peripheral re- low a lowly positioned periphery. Growth gion. The suture is deeply impressed and lines are sigmoidal, forming a broad sinus whorls embrace below the broad periphery. just below the ring of collabral nodes. Elon- The aperture is holostomous and the adult gate collabral nodes disappear just above the aperture has a greatly thickened outer lip sinus. The whorl face is flattened in the sinus formed into a large varix. The base is rounded region but becomes convex and sloping with no ornament and is anomphalus. downward to the periphery. The base is DIscuSSION: This genus is quite close to rounded, without ornament. Possibly mi- that ofProcerithiopsis Mansuy, 1914, in many nutely phaneromphalus. features but has an adult apertural varix and DISCUSSION: This species differs from the lacks the two columellar folds which are ge- undescribed Crimean species by having the neric level determinates. I have not found 18 AMERICAN MUSEUM NOVITATES NO. 2829 any other species that resembles these shells. SUPERFAMILY CERITHIACEA FLEMING, 1822 Further, as far as I am aware, Procerithiopsis FAMILY TURRITELLIDAE WOODWARD, 1851 has not been reported elsewhere other than by Mansuy; yet the genus is valid. The Paleozoic genera, Acanthonema Sher- ETYMOLOGY: Akron, Gk. top; spira, Gk. zer and Grabau, 1908, Callispira Nelson, twist. 1947, and Orthonema Meek and Worthen, 1862 previously placed in this family contain Acrospira cristata, new species species that all differ from Mesozoic and Ce- Figure 23 nozoic genera by having an aperture with the outer lip straight and nearly vertical. All post- DIAGNOSIS: Same as that of the genus. Paleozoic genera either have broad sinuses DISCUSSION: The most notable variation among the nine specimens is the change in with the greatest depth at the center of the the whorl profile from evenly rounded to flat- aperture or have anal notches near the suture. tened. The collabral ribbing varies somewhat Knight (1934, p. 436) first observed this sinus in the number (hence intensity) of ribs per in 0. bilineatum Mark, 1912, 0. schucherti whorl, a feature also noted in species ofpseu- Knight, 1934, and 0. inornatum Knight, dozygopleurids. The whorls embrace just be- 1934. Most species, such as 0. conicum Meek low the periphery, but if the whorl profile is and Worthen, 1866, have a straight, nearly flattened, then the embracement position will vertical, outer lip with some specimens show- cause the sutures to appear more deeply in- ing a slight backward oblique swing near the cised. Nine specimens. suture marking an incipient anal notch. Even MEASUREMENTS: Holotype AMNH 42819, though the sinuosity is the most defining fea- H 4.6 mm, W 2.7 mm, WhH 1.1 mm, WhW ture of the Turritellidae, the absence of the 2.0 mm, SpAng 380; paratype AMNH 42820, sinuous outer lip in some species has not been H 3.9 mm, W 2.2 mm, WhH 1.0mm, WhW regarded as significant in Paleozoic species. 0.8 mm, SpAng 400. It is interesting to note that no species of ETYMOLOGY: Crista, Lat. crest or ridge. turritellids are to be found in the uppermost Permian and throughout the entire Triassic! Acrospira, species The earliest occurrence of a Mesozoic turri- Figure 24 tellid is from the Jurassic Inferior Oolite of DESCRIPTION: The shell with seven to eight Great Britain (Huddleston, 1889). Those whorls with numerous fine collabral threads species are high-spired with rounded whorls which are collabral sloping downward from having a few spiral ribs and a broad, shallow the suture to the aperture. Whorls embrace sinus. at periphery and are inflated with gently con- vex outer whorl face which slopes downward and outward to a periphery that is low on the Genus Orthonema whorls. The base is flatly rounded and anom- Meek and Worthen, 1862 phalus. TYPE SPECIES: Eunema? salteri Meek and DISCUSSION: This species differs from A. Worthen, 1961. cristata in having less inflated whorls, with DISCUSSION: Orthonema is seemingly a finer ribbing and only a moderate amount of conservative genus, being represented in the thickening at the aperture and no sharply de- upper Paleozoic by 16 species. In most oc- fined varix. Whorls embrace higher on the currences it is known by a single specimen at shell, at the periphery. Only three complete localities where large numbers of other gas- shells are known and they are not sufficiently tropod species are found. detailed to formalize. This species is strik- The basis for species recognition is the spi- ingly similar to Procerithiopsis ambigua ral ornament which usually consists of two Mansuy, 1914 but lacks an umbilicus and groups of carinae each near the top or base columellar folds. Three specimens. ofthe whorl and separated by a flat or slightly MEASUREMENTS: AMNH 42821, H 6.0 mm, concave outer whorl face. In some species W 3.5 mm, WhH 1.5 mm, WhW 2.8 mm, only a single carina may be present at the SpAng 410. base ofthe whorl, or there may be more than 1985 BATTEN: PERMIAN GASTROPODS 19

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FIGS. 25-28. Orthonema. 25. 0. rectimurum, holotype, AMNH 42822, side view, x 5. 26. 0. naka- zawai, holotype, AMNH 42824, side view offragment, x 7. 27. 0. tirusum; (a) holotype, AMNH 42827, side view, x4.5; (b) paratype, AMNH 42828, side view showing the principal spiral rib low on the outer, foreshortened whorl face, x 6. 28. 0. knighti; (a) holotype, AMNH 42830, side view, x 3; (b) paratype, AMNH 42831, side view with low lighting to emphasize the basal rib and reflexed columellar lip, x 3. one present at a site. The relative placement growth lines, and the nature of secondary or- and development ofthese spiral elements af- namentation discussed below. fects the shape ofthe whorl profile and hence The Malaysian species are much more the entire shell shape. Sutural placement on abundant than at any other known locality. the whorl is another theme of species varia- For example, at the richest known Permian tion, although we can recognize interspecific locality (AMNH no. 512, Getaway Lime- variation patterns; we have hardly any un- stone, Guadalupe Mountains, Texas), Ortho- derstanding of intraspecific variation owing nema is represented by 10 specimens while to the small size ofpopulation samples. How- other genera are known by thousands ofspec- ever, this current study allows us to examine imens. Schindel (oral commun.) reports that several patterns that do occur with popula- in the Yale collections from the Pennsylva- tions, such as the intensity of spiral element nian of North Central Texas, one kilogram development, changes in the shape of the ofshale will yield one or two specimens where 20 AMERICAN MUSEUM NOVITATES NO. 2829 hundreds of other gastropod specimens are MEASUREMENTS: Holotype AMNH 42822, recovered. WhH 1.9 mm, WhW 1.6 mm; paratype (larg- The morphotypes of Orthonema in the est) AMNH 42823, WhH 8.3 mm, WhW 8.8 Malaysian Permian duplicate rather closely mm. the range ofvariation noted by Knight (1934) ETYMOLOGY: Rectus, Lat. straight or up- in the North American Pennsylvanian, as right; murus, Lat. wall. discussed in the Introduction. There appears to be a species of Orthonema in the Permian of the Crimea of the 0. salteri morphotype Orthonema nakazawai, new species which is undescribed. Other than that occur- Figure 26 rence, I have not found any other reports of the genus in the Tethyan Permian. The il- DIAGNOSIS: The outer whorl face is concave lustration of 0. cf. salteri in Delpey (1941, with one or more threads unevenly distrib- fig. 63, p. 67) is so generalized that I am un- uted; the upper rib defines the outer whorl in face and has several threads above it near the able to place it even a suborder. suture. The lower rib is much more devel- IMPORTANT NOTE ON MEASUREMENTS: All oped and is rounded; one or two threads un- ofthe specimens of Orthonema have the ear- derlie the lower rib on the concave base where ly whorls missing; no complete specimens it forms interference nodes with basal threads. were recovered. Therefore, I have omitted Some collabral ornament is noted on the up- the total height measurement since it is per whorl face as well. Sutures embrace at the meaningless. Only the whorl height (WhH) point where the base flattens out from a con- and whorl width (WhW) of the penultimate cave region at the edge of the lower whorl whorl are given and these have been made face. on the last complete whorl preserved. DISCUSSION: 0. nakazawai differs from 0. rectimurum n. sp. in that the outer whorl face is concave and the lower and upper ribs are Orthonema rectimurum, new species more centrally located. The ribs of the latter Figure 25 species are much closer to the sutures, form- DIAGNOSIS: Two dominant spiral ribs mark ing a wider outer whorl face. The two species an outer whorl face that is flat or slightly are similar in having the outer whorl face concave and inclined toward the axis; it is defined by an upper and lower rib. 0. recti- ornamented by one to ten spiral threads; if murum differs from 0. salteri in having a by a single thread, it is medially located and concave outer whorl face with the suture em- prominent. An upper rib or thread marks the bracing farther down the base. In addition, upper margin of the outer whorl face; it is the ribs are not as well developed in 0. sal- separated from the suture by an additional teri, so that the shell shape is less steplike. It thread. The sutures are sharp and deep. The differs from the North American 0. marvin- lower rib defining the outer whorl face is more welleri in having the lower rib more fully de- strongly developed and it also is separated veloped, and with collabral ornament (this is from the suture by a thread. The base is flat- an autapomorphic feature). Also, the spiral tened with one or two ribs as ornament. The threads adjacent to the sutures are weaker in columellar lip is straight and the lower lip the Malaysian species and the main ribs are forms an acute angle with the columellar lip closer to the sutures. It is similar to the North bearing a sharply defined groove. American populations in having the outer DISCUSSION: This species differs from the whorl face occupy over one-halfofthe whorl; North American 0. salteri by having a less the visible upper whorl face has one or two well-developed upper rib and by having a flat threads and an exposed base which also may almost vertical outer whorl face as contrasted have one or more threads. to the concave face of 0. salteri. That species The most notable variation within this has a rounded, unornamented base. I have species is the relative development ofthe up- found no other species that has many of the per and lower spiral ribs that delineate the features noted in these two. Eighteen speci- whorl face. In the holotype they are about mens. equally well developed and the outer whorl 1985 BATTEN: PERMIAN GASTROPODS 21 face between them is nearly flat. In one vari- the selenizone ofthe murchisoniid Stegocoe- ant, the upper rib complex is more developed lia which it resembles in several features. than the lower and in still others the lower This species differs from 0. rectimurum by rib is more fully formed. A number of spec- having the lower subsutural carina placed imens show the latter condition which causes lower on the whorl and in being more weakly the outer whorl face to appear concave, thus developed so that the outer whorl face is slop- giving the shell a more turriculate shape. Oth- ing rather than being nearly vertical; this re- er variations include changes in the numbers sults in a shell shape that is more conical and development of the basal spiral orna- compared to the turriculate 0. rectimurum. ment and in the relative height of the whorl It differs from 0. nakazawai in lacking the (reflecting the placement of the sutures). full rib development of the upper and lower Some specimens show some collabral or- ribs, the lack of the basal carinae, and with nament in the form of faint nodes on the a flattened base. spiral ornament only-not observed on the Variation between specimens is noted in whorl face itself. Seventeen specimens. the relative development ofthe spiral carinae MEASUREMENTS: Holotype AMNH 42824, and lower rib. Ifthese are well developed, the WhH 3.0 mm, WhW 5.3 mm; paratype outer whorl face tends to be flat or concave AMNH 42825, WhH 2.0 mm, WhW 4.0 mm; and the base more flattened. If the spiral or- paratype AMNH 42826, WhH 1.3 mm, WhW nament is less well formed, then the outer 2.9 mm. whorl face will appear convex and the base ETYMOLOGY: Named for Professor Keiji more rounded, giving the shell a more conical Nakazawa. shape. Five specimens. MEASUREMENTS: Holotype AMNH 42827, WhH 2.7 mm, WhW 7.6 mm; paratype Orthonema tirusum, new species AMNH 42828, WhH 4.5 mm, WhW 2.0 mm; Figures 27a, b paratype AMNH 42829 WhH 3.2 mm, WhW 5.8 mm. Note: only whorl heights and widths DIAGNOSIS: Moderately small shells with were measured on the last whorl preserved two subsutural carinae, the lower one form- because all specimens are fragments; also, the ing the shoulder ofthe outer whorl face. The spiral angle could not be taken. outer whorl face occupies about one-third of ETYMOLOGY: Tirus, Malayan for conical. the whorl. The upper subsutural carinae or thread is located in the central portion ofthe upper whorl face. The outer whorl face may be flat, gently convex, or somewhat concave Orthonema knighti, new species depending on the relative development ofthe Figure 28 carinae. The lower whorl face-outer whorl DIAGNOSIS: High-spired many-whorled face boundary is marked by a large rib. The shell with flat to rounded whorls. The suture whorl profile, on the whole, slopes gently out- is sharply defined but shallow. There is a faint ward giving the shell a rather smooth cone spiral thread which marks the lower margin shape. Growth lines form a sinus resulting in ofthe very narrow, concave upper whorl face. the formation of a pseudoselenizone located The outer whorl face is flat to slightly convex. between the upper suture and the shoulder. Whorls embrace on the basal spiral thread From the shoulder, the growth lines swing which marks the lower margin of the outer strongly opisthocline on the outer whorl face. whorl face. In some forms there is a basal rib The base is flat and the columellar lip is re- marking the lower margin ofthe outer whorl flexed forming a callus or funicle. face. The base is somewhat rounded with a DISCUSSION: The sinus is the deepest and spiral thread in the middle portion. most sharply defined of any of the genera of DISCUSSION: Because of poor preservation the Turritellidae. The only other genus that of most specimens, it is not possible to ob- shows a slight sinus is the Pennsylvanian Cal- serve the details of ornament or any other lispira Nelson, 1947 which has a broad sinus feature. However, the whorl profiles are well in the middle of the whorl. The sinus of 0. defined and strongly suggest that there was tirusum is somewhat closer to the suture than no ribbing to begin with. Therefore, I am 22 AMERICAN MUSEUM NOVITATES NO. 2829

reasonably confident that these specimens Carboniferous and Permian ofSoutheast Asia constitute a distinct species. Again, this by Mansuy (1914a) and Delpey (1942). The species appears to have an iterative counter- species are moderately variable with the vari- part in 0. inornatum Knight, 1934, which ation involving ornament and, more impor- also lacks typical spiral ribbing but has more tantly, axial translation. This latter variation convex whorl profiles. alters shell shape; in P. pupoides the rate of There is some variation in the shape ofthe axial translation in some individuals slows whorl profile ranging from slightly convex, down relative to whorl expansion causing the flat-sided to convexoconcave with the outer pupoid (beehive) shape. In P. delpeyi, new face either parallel or inclined to the axis species, the axial growth speeds up relative forming a conical shell. The specimens hav- to whorl expansion causing a coeloconoid ing convex-sided whorls are an enigma since shape to the shell (fig. 31). Ornament ranges no species of Orthonema possess smooth, from well-developed axial ribs which are unornamented convex whorls. However, straight and evenly formed from suture to there are gradations between those specimens suture in P. dussaulti Mansuy, 1914a to a and others that have flattened whorl profiles. reduced ornament with collabral ornament The growth lines (see fig. 28a) that are pre- restricted to the upper portion of the whorl served are typical of the genus in being es- in P. pupoides. sentially vertical with a sharp turn immedi- In the Treatise, Knight et al. (1960) placed ately adjacent to the suture, marking a shallow this group as a subgenus ofPseudozygopleura anal notch. The only other group to which based on several homologies. However, the these forms could be assigned would be some range ofvariation, as in the Malaysian fauna, genus of loxonematids, but they all have a clearly demonstrates a generic separation- well developed, broad sinus. Forty-one spec- indeed, I suggest that Paleostylus is not a imens. loxonematid at all but rather a cerithiid based MEASUREMENTS: Holotype AMNH 42830, on the fundamental apertural characters. WhH 3.0 mm, WhW 5.2 mm; paratype Delpey (1942) placed it in the Terebrellidae AMNH 42831, WhH 1.6 mm, WhW 2.7 mm. and suggested that it might be a campanilid, ETYMOLOGY: Named for J. Brookes Knight. but I find few features that are shared with those groups and such a placement appears FAMILY PROCERITHIIDAE COSSMANN, 1905 quite farfetched. DISCUSSION: Delpey (1942, p. 69) placed The pseudozygopleurid relationship does the genus Paleostylus Mansuy, 1914 in the have merit; the strong collabral ribs found in family Terebrellidae, an imperfectly known Paleostylus are nearly identical to those found Mesozoic group. Paleostylus shares a number in several species ofthe pseudozygopleurids, offeatures with Knishbia Winters, 1963, par- such as P. tenuivirga Knight, 1930. In Hel- ticularly transverse nodes confined to the up- minthozyga the last several whorls become per portion ofthe whorls, a basal rib forming uncoiled, thus resembling the cerithiid Ver- the periphery, a high-spired shell, and an in- micularia which displays a change in the re- cipient siphonal notch. Hence, I will assume lationship between axial translation and whorl that this family should be the proper place expansion as seen in Paleostylus but not to for this genus. the degree seen in those other genera. The The procerithids are common elements of most striking example of the convergence of the Triassic and Jurassic faunas, but are Paleostylus and Pseudozygopleura is in Pa- known only by Spanionema from the De- leostylus pseudozyga, new species, which vonian and Knishbia, Paleostylus, and the new would be described as a species of Pseudo- zygopleura if the apertural features were genus Sinozyga from the Permian. missing. Genus 1914 A feature which separates Paleostylus from Paleostylus Mansuy, the pseudozygopleurids is the presence of an TYPE SPECIES: Paleostylus pupoides Man- umbilicus. This feature was used by Hoare suy, 1914a, p. 48, pl. 7, fig. 16. and Sturgeon (1981, p. 572) to reject the use DISCUSSION: Paleostylus is known from of Paleostylus for North American pseudo- three species which were described from the zygopleurids and to point out the similarity 1985 BATTEN: PERMIAN GASTROPODS 23

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33 34 32 35 FIGS. 29-35. 29. Paleostylus pupoides Mansuy, 1914, AMNH 42832, side view of specimen with a weak basal rib, x 3. 30. P. pupoides, AMNH 42833, side view of a specimen with a low outer whorl face and a well-developed basal rib, x 7.5. 31. P. dussaulti Mansuy, 1914, AMNH 42836, side view, 1914. 32. P. delpeyae, holotype, AMNH 42839, apertural view, x 7. 33. P. pseudozyga, holotype, AMNH 42840, apertural view, note the well-developed siphonal trough and subsutural nodes on the second whorl, x 2. 34. Protostylus lantenoisi Mansuy, 1914, AMNH 42843, side view, x 5. 35. Sinuozyga ajoka, holotype, AMNH 42844, apertural view, x 7.5.

of Spiromphalus Hayaska, 1939 to Paleos- is similar to Paleostylus and the siphonal tylus. notches are all shallow. In addition, C. wans- The most important feature which sepa- fordiae is shown to have a coeloconoid shell. rates Paleostylus from the pseudozygopleu- I am uncertain as to the systematics of the rids is the presence ofa siphonal notch at the Huddleston species and it is beyond the scope junction of the columellar lip and the lower of this study to assess their taxonomic posi- lip. Looking over the Mesozoic faunas, I note tion; suffice it to say that I am convinced that the presence of a large number of species de- Paleostylus should be considered a cerithi- scribed as "Cerithium" in the Inferior Oolite acean rather than a loxonematid. I find noth- (Jurassic) by Huddleston (1889). Such species ing in the Triassic faunas that I can associate as C. subglabrum Huddleston, 1889, C. be- Paleostylus with. Finally, this genus is ap- anii Huddleston, 1889, or C. wansfordiae parently restricted to the Tethyan Permian. Huddleston, 1889 all have ornament which I am certain that all of the species from the 24 AMERICAN MUSEUM NOVITATES NO. 2829

North American Upper Paleozoic currently whorl face; in figure 29 the whorl face is high identified as belonging to Paleostylus and causing a more turriculate shell shape. The having holostomous apertures should prob- unornamented lower half of the outer whorl ably be considered pseudozygopleurids as face may be concave so that the upper half suggested by Hoare and Sturgeon (1981). and the basal spiral rib ofthe preceding whorl Paleostylus does occur in the North Amer- form a rectangular unit producing a more tab- ican Permian. I have examined some of the ulate shell. In some specimens, the spiral rib specimens in the type lot of P. giganticus at the base of the outer whorl face may be Winters, 1963 from the Permian Supai For- absent. The base is variable in being flat to mation of eastern Arizona which have col- rounded. Thirty-seven specimens. labral ornament concentrated near the upper MEASUREMENTS: AMNH 42832, WhH 2.8 suture, steplike whorls, spiral ornament con- mm, WhW 7.2 mm; AMNH 42833, WhH fined to the upper whorl face where it forms 2.0 mm, WhW 5.8 mm; AMNH 42834, WhH interference nodes with collabral ornament, 1.3 mm, WhW 2.5 mm; AMNH 42835, WhH and a narrow umbilicus. They differ from 2.2 mm, WhW 4.0 mm. species of Knishbia (also described from the Supai Formation by Winters) which has Paleostylus dussaulti Mansuy, 1914 rounded whorls and a different ornament pat- Figure 31 tern. However,judging from the suite ofspec- P. dussaulti Mansuy, 1914, p. 49, pl. 4, fig. 21. imens in the type lots of the species of both P. inosinicus Mansuy, 1914, p. 49, pl. 4, fig. 20. genera in the Supai material, I am ofthe opin- ion that Knishbia should also be considered DISCUSSION: The most significant feature closely related to Paleostylus if not a syn- ofthis species is the full development of col- onym. This conclusion could reinforce my labral ribs, which are spread uniformly across belief that Paleostylus is a cerithiid, since the whorl surface, rather than restricted as in Knishbia was placed in the Procerithiidae by other species. Furthermore, in contrast to Knight et al. (1960) near Orthonema. most species ofpseudozygopleurids, these ribs are vertical (or slightly opisthocline) and Paleostylus pupoides Mansuy, 1914 straight. The whorls embrace above the base. Figures 29-30 As in other species, the inner lip is reflexed to form a cryptomphalid condition of the Paleostylus pupoides Mansuy, 1914, p. 49, pl. 7, umbilicus. The chief features separating this fig. 16. species from the pseudozygopleurids are the DESCRIPTION: High-spired forms with axial siphonal notch and an umbilicus. Also the growth which may become progressively slope ofthe whorl profile is rectangular rather slower relative to whorl expansion causing than inflated. Eight specimens. the shell to be pupaeform. The whorls are MEASUREMENTS: AMNH 42836, WhH 1.5 flattened to somewhat convex with a sharply mm, WhW 3.2 mm; AMNH 42837, WhH defined spiral rib at the base ofthe outer whorl 1.3 mm, WhW 2.6 mm; AMNH 42838, WhH face which forms the boundary of the flat- 0.9 mm, WhW 2.4 mm. tened base. Whorls embrace just below this spiral rib. There are three spiral threads evenly Paleostylus delpeyae, new species spaced on the outer whorl face. Collabral ribs Figure 32 occupy the upper half ofthe whorl and form Paleostylus pupoides Delpey, 1942, fig. 66. interference nodes at the intersection of the upper two spiral threads. The third spiral DIAGNOSIS: Coeloconoid shells with col- thread is centrally located on the lower half labral ornament restricted to the upper quar- of the whorl face. Cryptomphalus. ter of the whorl. There is a single spiral rib DISCUSSION: There is some variation in ax- just below the suture which causes larger in- ial growth; in several specimens the shells are terference nodes to form at the intersection pupaeform, but most shells appear to be nor- of collabral ribs which extend to just below mally orthostrophic. There is a considerable a spiral thread just below the nodes. Faint amount ofvariation in the height ofthe outer traces of these collabral ribs can be found on whorl face; figure 30 shows a relatively low the lower whorl face. There are 10 or more 1985 BATTEN: PERMIAN GASTROPODS 25 fine spiral threads just barely visible on the uous to straight collabral ribs which are fully lower whorl face. Sutures embracejust below developed from suture to suture across the a very prominent flange marking the lower whorl face. Sutures are shallow, but sharply margin of the outer whorl face. The base is defined. The upper whorl face is flat to slight- flat. Narrowly phaneromphalus. ly concave and the outer whorl face is convex. DISCUSSION: There are but two known Subsutural nodes are present on early whorls. specimens representing this species. The Whorls embrace just beneath a periphery morphological discontinuity from other which is low on the whorl. The columellar species is sufficient to cause me no hesitation lip is reflexed and forms an angle with the in recognizing this species as being distinct. inner lip which has a well-developed siphon. Delpey (1942, p. 71) was of the opinion that There is no spiral ornament and the rounded the specimen she illustrated (fig. 66, p. 71) base has diminished collabral ornament. represents one end of a range of variation in Cryptomphalus. the rate ofaxial growth causing a coeloconoid DISCUSSION: This species is highly conver- shell shape, the rate of axial growth slowed gent on the pseudozygopleurids in shell shape, in relation to whorl expansion rate. She il- whorl profile, and dominant collabral rib- lustrated (an ink sketch) the specimen as hav- bing. Without the presence of subsutural ing uniform collabral ribs as in P. dussaulti, nodes on the early whorls and the complete but she also illustrated a specimen of a pu- aperture and base, this species would be con- poid type of P. pupoides as having the same sidered a species ofPseudozygopleura. There uniform development. In both cases, the Ma- is some variation in the shape of the whorl laysian specimens have the collabral ribs re- from evenly convex with a medial periphery stricted to the upper portion of the whorl, to a flattened whorl face near the upper suture and the presence of spiral ornament not il- becoming convex to a low periphery. The lustrated by Delpey. The types illustrated by development of the ribs ranges from fine to Mansuy (1914a, pl. 6, fig. 16) have identical coarse. The shell shape also varies from high- ornament patterns to those of the Malaysian spired to an almost trochoid shape. Seven specimens, so that again I suspect that Del- specimens. pey's renditions may not be accurate. MEASUREMENTS: Holotype AMNH 42840, In any event, the flange demarking the base H 20.6 mm, W 6.1 mm, WhH 3.7 mm, WhW is quite distinct from the relatively weak spi- 5.0 mm, SpAng 130; paratype AMNH 42841, ral rib in the same position in P. pupoides H 10.9mm,W4.1 mm,WhH2.3mm,WhW and is lacking altogether in P. dussaulti. The 3.1 mm, SpAng 14.50; paratype AMNH flange emphasizes the very flat base not seen 42842,H 3.2 mm, W 2.1 mm, WhH 0.9 mm, in the other species. The most important dis- WhW 1.7 mm, SpAng 400. tinction (quite apart from the distinct shell ETYMOLOGY: Pseudos, Gk. lie; zygon, Gk. shape) is the presence of a single spiral rib at yoke. the top ofthe whorl compared to two or more spiral ribs consistently found in P. pupoides. Genus Protostylus Mansuy, 1914 The outer whorl face over the lower three- quarters ofthe surface is devoid ofornament TYPE SPECIES: P. lantenoisi, Mansuy, 1914a, other than very faint spiral threads. Two SD, Batten, 1956. specimens. DISCUSSION: Knight et al. (1960) placed this MEASUREMENTS: Holotype AMNH 42839, genus in Loxonematacea, genus Inquirendum H 6.0 mm,W 5.0 mm, WhH 1.4 mm, WhW because details of the aperture were un- 5.0 mm, SpAng 220. The other species av- known. The specimens from this Malaysian erage about 100. fauna clearly show a reflexed columellar lip ETYMOLOGY: Named for Genevieve Del- and a siphonal trough indicating that the ge- pey. nus should be placed in the cerithiids. Paleostylus pseudozyga, new species Protostylus lantenoisi Mansuy, 1914a Figure 33 Figure 34 DIAGNOSIS: Moderately large, turriculate Protostylus lantenoisi Mansuy, 1914a, p. 11, pl. 1, shells with inflated, convex whorls with sin- fig. 17. 26 AMERICAN MUSEUM NOVITATES NO. 2829

DISCUSSION: I have identified five shells as There is no other genus to which compar- belonging to this species. The shells are nearly isons can be made. It bears a resemblance to identical to that illustrated by Mansuy (1914a, Loxosonia zygopleuroides in having a sinus pl. 1, fig. i), but there are some variations in on the upper whorl face; however in that ge- the whorl profile. Three specimens have nus the sinus is quite deep and narrow and slightly inflated and evenly rounded profiles. closer to the suture. The closest genus may The other two specimens have less inflated be Paleostylus based on the presence of a re- whorls with a flattened upper whorl surface flexed columellar lip, a siphonal notch, dom- and a rounded periphery just below the mid- inant collabral ornament, and the cryptom- whorl. The columellar lip is reflexed and the phalus condition. Therefore, I consider it to lower lip shows a siphonal trough. All spec- be a member of the procerithiids. The func- imens are cryptomphalus. tion of the sinus that appears on the adult Mansuy also described P. dussaulti and his whorls is a puzzle. It may be an anal notch, illustration (1914b, pl. 1, fig. 18) is of a shell but it is not at the suture where it is always with a more flattened whorl profile. Based on located in siphonate forms and why is it found the variation I have noted above, I suspect only in the adults? In almost every group that that P. dussaulti is synonymous with the type possesses a sinus beneath the suture, such as species. Delpey (1941a) does not report this the pleurotomarians, there are paired organs genus from the Cambodian Permian, but it associated with a holostomous aperture. This is possible that her illustration of Trypano- genus is unknown elsewhere. stylus cf. cerithiformis (p. 51, fig. 44) could ETYMOLOGY: Sinus, Lat. curve; zygon, Gk. be of Protostylus. Five specimens. yoke. MEASUREMENTS: (of illustration specimen) AMNH 42843, H 8.2 mm, W 2.8 mm, SpAng Sinuozyga ajoka, new species 290. Figure 35 Sinuozyga, New Genus DIAGNOSIS: Moderately high-spired forms with at least nine whorls. The whorl profile DIAGNOSIS: Moderately high-spired si- is rounded with an ill-defined periphery either phonate forms with a reflexed columellar lip, at midwhorl or low on the whorl. The col- presumably cryptomphalus. The first several labral ribbing is well developed, becoming whorls have very faint spiral ornament; the stronger with growth. The base is rounded. next several have relatively straight, nearly DISCUSSION: The six specimens available vertical collabral ribs. The last two or three for study show some degree ofvariability per- whorls have a slightly depressed trough just mitting a study ofinterspecific variation. The beneath the suture on the upper whorl face. whorl profile is variably shaped from an Collabral ribs and growth lines abruptly bend evenly rounded, convex profile to one which backward into the trough forming a shallow has a recognizable periphery low on the whorl; asymmetrical sinus. In the trough the ribbing also the profile may be less inflated. Ribbing disappears. The whorl profile is evenly con- shows a range from relatively fine ribs to vex except at the trough region in the adult coarse ribs fewer in number on the whorl. whorls where there is an indentation. Some Six specimens. specimens have a poorly defined periphery MEASUREMENTS: Holotype AMNH 42844, low on the whorl; most specimens have a H 6.7 mm, W 2.5 mm, WhH 1.7 mm, WhW periphery in midwhorl. The base is rounded. 2.5 mm, SpAng 620; paratype AMNH 42845, DISCUSSION: Because of the presence of an H 5.9 mm, W 3.1 mm, WhH 2.1 mm, WhW umbilicus, reflexed columellar lip, shallow si- 3.1 mm, SpAng 600. phonal trough, and sinus, this genus cannot ETYMOLOGY: means mimic in Ma- be placed in the pseudozygopleurids in spite Ajoka of its close resemblance in shell shape, whorl layan. profile, and collabral ribbing. The ornament, in contrast to most species of Pseudozygo- FAMILY COELOSTYLINIDAE COSSMANN, 1909 pleura, becomes progressively stronger with Wenz (1938, p. 39) included a number of growth. genera in this family which are highly diver- 1985 BATTEN: PERMIAN GASTROPODS 27 gent in many basic features and by today's whorls embracejust below a broad, low-lying criteria are clearly unrelated. A revision of periphery. The base is flatly rounded. There the family is long overdue. Because Wenz is very faint ornament in the form of broad included Omphaloptychia Ammon, 1892 in collabral raised areas more fully developed the family, I will retain Wenz's scheme. Since on the base (see fig. 36c). Phaneromphalus to there are three species in the Malaysian fauna cryptomphalus. The columellar lip is reflexed I do not believe it is warranted to undertake and the parietal inductura is thin. A broad a revision. Wenz placed this family in the shallow sinus in the lower lip is located ad- Loxonematacea but because of the impor- jacent to the columella. tance I have placed on the siphonate condi- DIscuSSION: These very conservative shells, tion, I would suggest that this family be placed as in other species, lack general ornament near the Procerthiidae in the Cerithiacea. except for some ofthe specimens which have very faint subsutural nodes. The whorls are Genus Omphaloptychia Ammon, 1892 more inflated and the sutures embrace lower TYPE SPECIES: 0. nota Ammon, 1892, p. on the whorl than those without the nodes, 199. giving a more steplike shell shape; these re- DIscuSSION: There has been much discus- semble the St. Cassian Coelostylina crassa sion about the relationship and the status of Kittl, 1894. The others in this sample resem- Omphaloptychia and Coelostylina (see Haas, ble C. conica Kittl, 1894. Both ofthese Perm- 1953, pp. 131-132, 137). Haas decided to ian forms differ from the Triassic species by recognize both genera, placing the lower- having a more developed siphonal notch. spired more robust species in the former ge- Seventy-four specimens. nus and the higher-spired forms in the latter. MEASUREMENTS: Holotype AMNH 42846; The problem with recognition lies in the fact H 9.1 mm, W 4.8 mm, SpAng 340; paratype that these species are very conservative and AMNH 42847, H 9.4 mm,W 5.2 mm, SpAng are almost featureless with smooth, rounded 320; paratype AMNH 42848, H 9.6 mm, W whorls like beans. I have only used the lit- 5.2 mm, SpAng 430. erature for a survey of the species and con- ETYMOLOGY: Named for the Paleozoic Era. clude that all species should be placed in a single genus. Since Omphaloptychia is the Omphaloptychia cingulata, new species prior name, I shall use that. Figure 37 The genus is characterized by having a smooth shell with sigmoid growth lines but DIAGNOSIS: Moderately high-spired tur- without ornamentation, and medium high- biniform shells with a well-developed row of spired turbiniform shells which may be nodes adjacent to the suture. Whorls are gently phaneromphalus or anomphalus and with a rounded and sutures embrace at orjust below siphonal trough. In addition, I have found a low, ill-defined periphery. Growth lines are ornament in the two described species. 0. sigmoid but nearly vertical at the suture where cingulata has subsutural nodes and 0. paleo- elongated nodes occur. The columellar lip is zoica has faint basal collabral ornament. It is reflexed, forming a narrow, arcuate parietal widely distributed and one ofthe most abun- inductura. The lower lip has a broad, shallow dant snails found in the Triassic. Wenz re- siphonal trough. Cryptomphalus. ported that it is found in the Permian, but I DISCUSSION: This species resembles 0. pa- have not seen it in the literature or in collec- leozoica but has stronger subsutural nodes tions except for this one occurrence. and less inflated whorls. It sharply differs from all other known species by the presence of subsutural nodes. Anozyga Hoare, 1980, Omphaloptychia paleozoica, new species is moder- Figures c classified as a pseudozygopleurid, 36a, b, ately high-spired with a very similar shell DIAGNOSIS: Moderately high-spired tur- shape and a row ofsubsutural nodes. It differs biniform shells with convex to somewhat in- in being holostomous and having flattened flated whorls with or without weak subsutural embryonic whorls. An undescribed specimen nodes. The suture is deeply recessed. The in the Molengraaff Collection (housed in the 28 AMERICAN MUSEUM NOVITATES NO. 2829

Geological Museum of the Delft Technical SUPERFAMILY SUBULITACEA Institute) from the Permian of Nefalassi, Ti- LINDSTROM, 1884 mor, has flattened whorls which are slightly FAMILY MEEKOSPIRIDAE KNIGHT, 1956 concave in the upper half of the whorl and Genus Meekospira Ulrich, has a raised subsutural rib with nodes, the in Ulrich and Scofield, 1897 whorls embracing higher on the whorls. It should be included with this species. Sev- TYPE SPECIES: Eulima? peracuta Meek and enteen specimens. Worthen, 1861 (Ulrich, p. 1079). MEASUREMENTS: Holotype AMNH 42849, DISCUSSION: As far as I am aware, Mee- H 4.4 mm, W 2.6 mm, SpAng 380; paratype kospira has not been reported previously from AMNH 42850, H 5.2 mm, W 2.6 mm, SpAng Asia. A specimen identified as Holopella cf. 380; paratype AMNH 42851, H 5.0 mm, W trimorpha Waagen by Grabau (1931) from 3.8 mm, SpAng 470. the Permian of Mongolia quite possibly is a ETYMOLOGY: Cingulum, Lat. belt or zone. member of the genus, but it is a fragment. The illustration in Waagen (1880, pl. 10) is Omphaloptychia sp. apparently a steinkem ofa murchisoniid from Figure 38 the Productus Limestone of the Salt Range. DISCUSSION: A group of quite variable Meekospira melanoides, new species specimens belongs to a separate species but Figure 40 is not sufficiently well preserved to warrant formalization. They are characterized by a DIAGNOSIS: Many-whorled high-spired distinct angulate periphery very low on the Melania-like shells with sutures at an acute whorl. Figure 38 illustrates a lower-spired angle to the axis. The whorl profile is flattened form. Fifteen specimens. or very gently convex, giving the shell an MEASUREMENTS: AMNH 42852, H 6.8 mm, evenly fusiform shape. The aperture is nar- W 4.9 mm, SpAng 320. row, teardrop shaped, and holostomous. The columellar lip is reflexed. The base tapers to Genus Trypanostylus Cossmann, 1895 a narrow point at the bottom ofthe lower lip. Anomphalus. TYPE SPECIES: Eustylus militaris Kittl, 1894. DISCUSSION: Most species ofthe genus have DISCUSSION: Cossmann erected this genus fewer whorls (usually 6-8) than does M. mel- name to replace Eustylus Kittl, 1894, which anoides (with at least 10). The rate of axial is a homonym ofEustylus Schomherr, 1843. translation is much greater than for other known species and this is reflected in the Trypanostylus triadicus Kittl, 1894 sharply acute suture lines. This rapid rate also Figure 39 results in the more flattened whorls and the Eustylus triadicus Kittl, 1894, p. 1955 pl. 8. figs. narrow aperture. No other species is remotely 26-27. similar to this one, which represents an ex- DISCUSSION: Two specimens in the fauna treme variant for an elongated species within are so morphologically similar to this St. Cas- this genus. Eighteen specimens. I can find no of MEASUREMENTS: Holotype AMNH 42854, sian species that way recog- H 4.55 mm, W 1.28 mm, SpAng 60; paratype nizing differences. The whorl profile is flat AMNH 42855, H 3.91 mm, W 1.19 mm, with a deeply incised suture, but, important- SpAng 70. ly, there is a narrow siphonal trough on the Melania. outer margin of the base. The columellar lip ETYMOLOGY: Like the genus is reflexed and a siphonal notch is incised at the junction of the columellar lip and the Meekospira ligoni, new species lower lip. These are apomorphic features of Figure 41 the St. Cassian population. Two specimens. DIAGNOSIS: Inflated, fusiform shells with MEASUREMENTS: AMNH 42853, H 5.4 (e) suture lines normal to the coiling axis. The mm, W 2.1 mm, WhH 0.9 mm, WhW 0.8 whorl profile is broadly convex with a round- mm, SpAng 190. ed periphery at midwhorl. Whorls embrace 1985 BATTEN: PERMIAN GASTROPODS 29

,

.0.4 Is. I

T./ -rI

36a 36b

wf37 )3 38 39 FIGS. 36-39. 36. Omphaloptychia paleozoica; (a) holotype, AMNH 42846, side view, x 8; (b) para- type, AMNH 42848, apertural view, x 8; (c) side view, x 8. 37. Omphaloptychia cingulata, holotype, AMNH 42849, side view, x 6. 38. Omphaloptychia sp., AMNH 42852, side view, x 8. 39. Trypanostylus cf. triadicus Kittl, 1894, AMNH 42853, side view, x 8. well below the periphery. Whorl expansion ETYMOLOGY: Named for William Ligon, rate is moderate. Aperture oval shaped and Jr. the columellar lip is straight. DISCUSSION: This species, like M. mela- FAMILY SUBULITIDAE LINDSTROM, 1884 noides, has a rather high axial translation rate SUBFAMILY SOLENISCINAE, WENZ, 1938 compared to other species. It differs from the Genus Strobeus 1881 type species in having a higher whorl expan- deKoninck, sion rate with the same number of whorls TYPE SPECIES: S. ventricosus deKoninck, (eight). As a result, the shell shape is more 1881, pl. 3, figs. 26-27. fusiform than in the type. I have not found DISCUSSION: The Treatise on Invertebrate this species in other faunas. Seventeen spec- Paleontology, Part I, recognized Ianthinopsis imens. Meek and Worthen, 1866, as the valid name MEASUREMENTS: Holotype AMNH 42856, for this group. This was not a wise decision H 4.27 mm, W 1.37 mm, SpAng 11°; para- since the type is missing and the original de- type AMNH 42857, H 1.65 mm, W 0.65 scription and illustrations prevent firm rec- mm, SpAng 200. ognition ofthe taxon. Strobeus ventricosus is 30 AMERICAN MUSEUM NOVITATES NO. 2829

-

42 - 43 41 i.\I

-t"04 44

40 FIGS. 40-45. 40. Meekospira melanoides, holotype, AMNH 42854, apertural view with low side lighting to bring out details of the reflexed columellar lip, x 2. 41. Meekospira ligonensis, holotype, AMNH 42856, side view, x 10. 42. Strobeus sp., AMNH 42858, apertural view, x 10. 43. Soleniscus elegans Gemmellaro, 1889, AMNH 42859, side view, x6. 44. Cylindritopsis sp., AMNH 42860, side view with the aperture broken to reveal the columellar folds, x 1.75. 45. Ceraunocochlis sp., AMNH 42861, side view, x7. well known in Europe and the type is pre- MEASUREMENTS: (of illustrated specimen) served in Brussels, and hence should serve AMNH 42858, H 3.2 mm,W 3.0 mm, SpAng well as the name bearer, as discussed by Har- 840. per (1981). Genus Soleniscus Meek and Worthen, 1861 Strobeus sp. TYPE SPECIES: S. typicus Meek and Wor- Figure 42 then, 1861, p. 467 DISCUSSION: Six very small and not well Soleniscus elegans Gemmellaro, 1889 preserved specimens can be referred to this Figure 43 genus. Delpey described three specimens of what she called Soleniscus from Cambodia. DISCUSSION: Twelve tiny specimens of The Malaysian specimens most closely re- subulitids in this fauna lack columellar folds semble the illustrations she labeled Solenis- or apertural teeth. They vary considerably in cus welleri Knight, 1931. They are low-spired shell shape, whorl convexity, and whorl ex- whorls that are evenly inflated but with a pansion rate but lack of other significant fea- rounded periphery centrally located. Broken tures precludes further taxonomic assess- specimens reveal a columellar fold low on ment. The specimen illustrated comes closest the columella, the distinctive generic feature. to a described species of Girtyspira with a Six specimens. deeply set suture suggesting a ramp, similar 1985 BATTEN: PERMIAN GASTROPODS 31 to that of Girtyspira fusiformis deKoninck siphonal notch is well developed. One spec- (1881). The whorls are evenly convex and imen. whorls embrace just below the broad periph- MEASUREMENTS: (of illustrated specimen) ery. However, S. elegans ofGemmellaro also AMNH 42861, H 5.3 mm, W 2.1 mm, SpAng has deeply set sutures but lacks the ramp so 15°. that, to be conservative, I shall assign these specimens to that taxon. Thirteen specimens. MEASUREMENTS: AMNH 42859, H 4.7 mm, ?SUPERFAMILY NERINEACEA ZITTEL, 1873 W 2.5 mm, SpAng 390. FAMILY NERINEIDAE ZITTEL, 1873 DISCUSSION: I provisionally place the new Genus Cylindritopsis Gemmellaro, 1889 genus Prodiozoptyxis in the nerineids as dis- TYPE SPECIES: C. oovalis Gemmellaro, 1889. cussed below. The nerineids are a prominent Jurassic and Cretaceous group which have Cylindritopsis sp. not been firmly related to any other group. Figure 44 According to Cossmann (1909, vol. 21, p. 209), two genera, Pseudonerinea (Ceritelli- DISCUSSION: Three specimens can safely be dae) and Iturvia (Itieriidae) have hetero- assigned to this genus. They are relatively strophic nuclear whorls and, in consequence, high-spired and clearly have two distinct col- he erected an opisthobranch suborder, the umellar folds low on the columella. Most Entomotaeniata, for the superfamily. Knight species have an ovoid shell shape with the et al. (1960, p. I145) point out that the group exception of C. inflata Gemmellaro, 1889, possesses features not present in most of the from the Permian Sosio beds of Sicily. Spec- opisthobranchs, namely the short siphonal imens are rare in most faunas and spotty in canal and a narrow anal emargination of the distribution in the Upper Paleozoic. outer lip (as in the conidae), which gives rise Delpey described a new genus, Angkorella, to an anal fasciole. Yochelson (1956) sug- in 1942 from Cambodia and it clearly is syn- gested that the columellar folds resemble those onymous with this genus. The type species ofthe subulitacean Labridens but the absence A. sisophonensis is very low-spired and ovoid of an anal fasciole, the relatively low whorl but unrelated to the Lee Mine forms. Since translation rate, and shell thickening in that the three Malaysian specimens are not par- genus suggest convergence rather than a re- ticularly well preserved, I have refrained from lationship. However, I lean toward a subu- formalizing a species name. Three speci- litid derivation. Houbrick (1981, p. 282) states mens. that there is some doubt regarding the place- MEASUREMENTS: (of illustrated specimen) ment of the Cretaceous genus Diopzoptyxis AMNH 42860, H 1.79 mm, W 1.22 mm. in the nerineidae. This is based, in part, on the study of the campanilids by Delpey SUBFAMILY SUBULITINAE LINDSTROM, 1884 (1941b) in which she discussed the features Genus Ceraunocochlis Knight, 1931 of Diozoptyxis she considered to be campa- nilid in nature. The most important were a Ceraunocochlis sp. heterostrophic nucleus and an anal fasciole. Figure 45 However, the campanilids lack the complex internal folds that characterize the nerineids DISCUSSION: A single sinistral specimen and for this reason I will place Prodiozoptyxis clearly should be assigned to this genus. Its along with Diozoptyxis in that group. axis of coiling is curved and the shell is fu- The most improbable assignment of the siform, typical of the genus. The genus is nerineaceans was made by V. F. Pchelintsev found sporadically in the Upper Paleozoic, (1968) who placed them into the order Mur- usually a single specimen at each locality. The chisoniata (Mesogastropoda) along with the whorls are evenly convex; the sutures are murchisoniids (Archeogastropoda), ceri- deeply indented, forming a weak subsutural thiids, turritellids, caecids, and ianthiniids (all ramp. Whorls embrace beneath a broad pe- Mesogastropoda) along with many other riphery. The base is evenly convex and the groups. The lack of a selenizone is a clear 32 AMERICAN MUSEUM NOVITATES NO. 2829

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48b 48a 47

FIGS. 46-48. 46. Prodiozoptyxis permiana; (a) holotype, AMNH 42862, side view, x 8.5; (b) aper- tural view, x 8.5; (c) paratype, AMNH 42863, apertural view with some of the whiting removed from the spiral ribs to show collabral nodes, x 7.5. 47. Diozoptyxis monilifera (d'Orbigny), AMNH 42865, apertural view showing siphonal trough, x 0.75. 48. Prodiozoptyxis sp., from the Permian Colina For- mation, Warren, Ariz. (AMNH Loc. 2019); (a) AMNH 42866, side view, note pupoid shape, x 6; (b) AMNH 42867; (b) an apertural view of a fragment showing the deep siphonal trough, x 8. indication that they are not related to the zoptyxis monilifera (d'Orbigny) which has two murchisoniids. spiral ribs occupying most of the whorl sur- face except for a narrow trough separating Prodiozoptyxis, New Genus them in the midwhorl, and in bearing elon- TYPE SPECIES: Prodiozoptyxis permiana, gate axial costae or nodes. The height and new species. shape of the shell, the phaneromphalus, flat- DIAGNOSIS: High-spired, turriculate shells tened base, and the quadratic aperture are the with spiral angles less than 30°, two spiral same in both genera. ribs forming most of the external whorl sur- The principal difference is in the internal face, two internal folds located on the upper fold pattern, which is the characteristic of and lower internal whorl surfaces, phaner- most importance for generic recognition omphalus or cryptomphalus. within the family. In Diozoptyxis there is a DISCUSSION: This genus is highly conver- strong upper internal surface fold and a lower gent on the Cretaceous nerineid species Dio- internal fold as in Prodiozoptyxis. But, in ad- 1985 BATTEN: PERMIAN GASTROPODS 33

.4

r

49 50 51

FIGS. 49-51. 49. Donaldina sp., AMNH 42868, apertural view, x 17. 50. Streptacis sp., AMNH 42869, side view, x 17.5. 51. Spirocyclina sp., AMNH 42870, side view of fragment, x 15. dition, Diozoptyxis has two axial folds and a There is an undescribed species in the low parietal fold. Since the Permian genus is Permian Colina Limestone in the Mule the earliest appearing member of the family, Mountains of S.E. Arizona (AMNH Loc. it would be reasonable to assume that the 2019), figure 48. It has two large spiral ribs internal fold system should be the simplest, separated by a relatively narrow trough much as is the case with the earliest Jurassic neri- like P. permiana but differs from that species neans. No function has yet been assigned to in having a deeper trough and lower spires. this system. The axial translation rate appears to slow down with growth, causing a somewhat pu- Prodiozoptyxis permiana, new species poid shell shape. Figures 46-48 There is some variation in P. permiana in DIAGNOSIS: Turriculate shells with coiled, the whorl translation rate, which causes the rope-like whorl profiles, with eight or more shell shape to be more or less turriculate. The whorls; elongate axial costae or nodes on two ornament varies from a faint axial threading large spiral ribs are weakly developed and on the spiral ribs to moderately well-devel- have faintly developed collabral nodes. oped costae. Thirty-seven specimens. Phaneromphalus. Base flattened, two inter- MEASUREMENTS: Holotype AMNH 42862, nal folds present. H 11.0mm, W4.3 mm, WhH 2.2 mm, WhW DISCUSSION: This is the only species known; 4.3 mm, SpAng 250; paratype AMNH 42863, therefore most of the generic diagnosis ap- H 5.7 mm,W 3.6 mm, WhH 1.6 mm, SpAng plies to it. A sample of Diozoptyxis monili- 23°; paratype AMNH 42864, H 4.3 mm, W fera (d'Orbigny) (AMNH #2894/1) from the 2.4 mm, SpAng 280; Diozoptyxis monilifera Cretaceous Naaman beds of Syria, figure 47, AMNH 42865, H 12.6 mm, W 6.0 mm, shows a high degree ofvariability particularly SpAng 280. in the shape ofthe whorls. Shape changes are due to the relative development ofthe spiral ribs, the development of the upper rib at the Prodiozoptyxis sp. suture, and the development ofthe lower rib Figures 48a, b at the basal margin. In some specimens, the DISCUSSION: This is an undescribed species two ribs are weakly formed so that the whorl from the Permian Colina Limestone, in the face between them is broad and flat, with a Mule Mountains, one mile north of Warren, flat base. In other specimens the two spiral Arizona, U.S.A. (AMNH Loc. 2019). Ten ribs are massive and occupy all of the whorl specimens. face so that only a narrow trough separates MEASUREMENTS: AMNH 42866, H 6.8 mm, them. The result is that the whorl profile is W 4.2 mm, SpAng 410; AMNH 42867, WhH inflated as in the Permian genus. 1.9 mm, WhW 3.4 mm. 34 AMERICAN MUSEUM NOVITATES NO. 2829

ORDER OPISTHOBRANCHIA distinctive spiral threads are unevenly spaced on the whorl with the most prominent thread SUPERFAMILY PYRAMIDELLACEA forming a low-lying periphery on the inflated D'ORBIGNY, 1840 whorl. The columellar lip is reflexed and ob- FAMILY STREPTACIDIDAE KNIGHT, 1931 scures a narrow umbilicus. Laube and Kittl Genus Donaldina Knight, 1931 defined the genus and type species from the Middle Triassic St. Cassian beds of Cortina, TYPE SPECIES: Aclisina grantonensis Don- Italy, and Wenz (1938) placed the genus in ald, 1898, p. 60. the streptacids, a conclusion with which I concur. One specimen. Donaldina sp. MEASUREMENTS: AMNH 42870, WhH 1.4 Figure 49 mm, WhW 1.5 mm. DISCUSSION: Three fragments appear to be- long to this genus. The whorl profiles are ?SUBORDER TROCHINA evenly convex and there are spiral threads SUPERFAMILY TROCHACEA RAFINESQUE, 1815 ornamenting the lower halfofthe whorl. The FAMILY TURBINIDAE RAFINESQUE, 1815 growth lines are sinuous and the apertures This family ranges from the Triassic to Re- are holostomous. Their size also suggests that cent and is worldwide in distribution. Most they are members of the genus. The embry- genera have strong ornament dominantly ax- onic whorls are missing, as are the diagnostic ial or collabral and there is a considerable heterostrophic nuclear whorls. Three speci- range of shell shapes from almost planispiral mens. to high-spired and uncoiled like Vermicular- MEASUREMENTS: AMNH 42868, WhH 0.5 ia. mm, WhW 0.7 mm. SUBFAMILY LIOTIINAE Genus Streptacis Meek, 1872 ADAMS AND ADAMS, 1854 TYPE SPECIES: S. whitfieldi Meek, 1872, p. Genus Eucycloscala Cossmann, 1893 173. TYPE SPECIES: Trochus binodosus Munster, Streptacis sp. SD, COX, 1960. Figure 50 DISCUSSION: When I first observed the DISCUSSION: A single specimen clearly be- specimens I am now identifying as Eucy- longs to this genus. It has the diagnostic het- closcala, I believed them to be promathildid erostrophic nuclear whorls and the typical cerithiaceans based on the presence of a si- elongate, somewhat flattened whorl profiles. phonal trough, the shell shape, varices, and The growth lines are sinuous and the aperture other distinctive ornament features. These is holostomous. As is typical, the shell lacks specimens are convergent on Promathilda ornament. This is the first report of either tyrsoecus (Kittl), 1894, or P. decorata (Klip- Donaldina or this genus in the Tethyan stein) Kittl, 1893. But there is a dilemma Permian. One specimen. which I cannot resolve at this point. The MEASUREMENTS: AMNH 42869, H 2.4 mm, Permian specimens even more closely resem- W 0.8 mm, SpAng 200. ble many species ofTriassic Eucycloscala such as the type species. The problem is that the Genus Spirocyclina Kittl, 1894 Malaysian shells all have siphonal troughs and reflexed columellar lips that are never TYPE SPECIES: Turritella eucycla Laube, found in the trochids. However, they also 1869, p. 14, pl. 30. have orthostrophic nuclear whorls, whereas the promathildids have markedly hetero- Spirocyclina sp. strophic nuclear whorls. Since there are more Figure 5 1 elements in common with Eucycloscala, I will DISCUSSION: A fragment of a single whorl assume that the Permian specimens should is sufficient to place it in this genus. The very be assigned to that genus. The illustrations 1985 BATTEN: PERMIAN GASTROPODS 35

* ,.-.Ss

. I% -49 "'');s orm"I'l isb *t txLssRo 4 sSs;! *s S -u ,K,X,, ,^S.,#L s ^ 'tl bkr-:Fiif$ ;jw

> 53 ^< 52a 54

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r .A*--All .. I X

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FIGS. 52-57. 52-54.52(a) Eucycloscala asiatica, holotype, AMNH 42871, side view, x 8; (b) oblique basal view, x 8. 53. Paratype, AMNH 42871, side view showing well-developed collabral ribs on outer whorl face, x 8; 54. Paratype, AMNH 42873, apertural view showing varixlike ribs, x 7.5. 55-57. Eucycloscala medionodosa. 55. Holotype, AMNH 42874, apertural view, x6. 56. Paratype, AMNH 42875, side view, x 8.5. 57. Paratype, AMNH 42876, side view, x 12.5. of the Triassic species do not indicate the per whorl face, at the shoulder, and on the presence of a siphonal trough and this is the basal margin. There are several additional principal difference between the Permian and threads on the base that become progres- the Triassic species. sively weaker on the base. The columellar lip is reflexed. Cryptomphalus. The junction of the columellar and lower lips is the site of a Eucycloscala asiatica, new species siphonal trough. Figures 52-54 DISCUSSION: There is considerable varia- DIAGNOSIS: Relative high-spired, turricu- tion in the development ofthe varices. In one late shells with concave profiles on the upper specimen, the varices are sharply delineated third of the whorls. The outer whorl face is and evenly developed on the outer whorl face. flat to somewhat convex and slopes inward On another, the varices are more diffused toward the base. The base is flat to somewhat with the upper portion rounded almost into concave. The sutures may be wavy, reflecting a boss; in other specimens, the varices are the whorl embracement at the sites ofvarices. uniformly rounded to form an elongate node Whorls embrace at the lower margin of the on the outer whorl face. In some specimens outer whorl face. The periphery is the shoul- the outer whorl face is flat and the upper der marking the junction of the upper and whorl face is concave giving an angulate, tur- outer whorl faces. Five to seven varices are reted appearance; in others, the upper and developed mostly on the outer whorl face at outer whorl face may be convex giving the the shoulder. There are usually four spiral whorl a globose profile and a more trochoid costae; one adjacent to the suture, on the up- shell shape. Nine specimens. 36 AMERICAN MUSEUM NOVITATES NO. 2829

599

" -- I W-IT".., .U....14, , , i Popp ., . e '. icA ..j Aii k - .S. ,4 a 58

FIGS. 58-59. 58. Glyptospira sp., AMNH 42918, side view, x 16. 59. Phymatopleura sp., AMNH 42919, side view, x 24.

MEASUREMENTS: Holotype AMNH 4287 1, The predominant shell shape, however, is high H 5.8 mm, W 3.2 mm, SpAng 230; paratype turbiniform with a row of nodes on a pe- AMNH 42872, WhH 1.7 mm, WhW 3.2 mm. riphery low on the whorl. Thirty-one speci- mens. Eucycloscala medionodosa, new species MEASUREMENTS: Holotype AMNH 42874, Figures 55-57 H 7.2 mm,W 4.0 mm, WhH 1.7 mm, WhW 4.0 mm; paratype AMNH H 2.7 mm, DIAGNOSIS: Variable shells, medium to 42875, W 3.6 mm, WhH 2.7 mm, WhW 3.6 mm, high-spired with dominant spiral ribs located SpAng 460. midwhorl to somewhat lower, forming the ETYMOLOGY: Medius, Lat. middle; nodo- periphery. Spiral ornament consists of a sus, Lat. knot. prominent thread adjacent to the suture, a central spiral rib, and a spiral thread at the GENERAL CONCLUSIONS basal margin. The upper whorl surface is con- cave, flat, or convex. The lower whorl face is Now that this Tethyan fauna is known, it short and usually flat. Whorls embrace just is clear that the southeastern Asian gastropod below the basal spiral thread. The rounded fauna is the probable source for the Mesozoic or flat base has several spiral threads. Col- faunas that spread from that region to all of labral ornament is either threads, nodes, or the seas by Jurassic time. The elusive lower varices confined to the periphery and lower Triassic refugia were undoubtedly concen- whorl surface. Crytomphalus. trated in the center ofthe eastern Tethys and DISCUSSION: This is one of the most vari- should be looked for in eastern Tibet to east- able species encountered in the Malaysian ern Sichuan (Wang, 1978). The western Teth- Permian. The width and shape of the upper yan fauna, while having many common ele- whorl face, the strength of the collabral or- ments with the eastern, is related in a nament, and the base greatly alter the shell fundamental way with faunas ofthe Western shape. The region between the periphery and Hemisphere, particularly the southwestern the basal thread is the same as the outer whorl United States and western South America. face in E. asiatica and is occupied by orna- The presence ofthe nerineid Prodiozoptyxis, ment, in contrast to the well-developed and the trochid Eucycloscala, the neritid Trach- usually flat outer whorl face of the other ydomia, the murchisoniid Loxosonia, the species. Ornament tends not to be as well loxonematids Sinozyga and Acrospira, and developed in this species. There are from 10 the coelostylinids Omphaloptychia and Try- to 15 nodes per whorl compared to 5 to 7 panostylus are all on phyletic lines leading to varices in E. asiatica. The spiral thread next major elements of the Jurassic and Creta- to the suture is much more strongly formed ceous cosmopolitan fauna. and is consistently present. The height of the There are two aspects of the Permian fau- spire is also variable, ranging from almost nas as they relate to Mesozoic faunas: (1) the turbiniform to a high cerithid appearing shell. presence oflong-ranging, conservative Perm- 1985 BATTEN: PERMIAN GASTROPODS 37 ian genera in the Middle to Upper Triassic convergence by identifying the Cambodian and (2) "precursor" genera in the Permian of Permian species as those of Western Europe higher categories that become important and or North America. However, in such groups diverse groups in the Mesozoic. These as- as the pseudozygopleurids and in Orthone- pects give the Triassic faunas a distinct Pa- ma, a cladistic analysis shows that apo- leozoic appearance that I have discussed pre- morphic characters group East Asian Teth- viously (Batten, 1973). At that time, I yan species more closely, thus providing observed that the last of the diverse marine sufficient evidence to warrant recognition of gastropod faunas in most of the world oc- the Malaysian species as being distinct. curred at the end ofGuadalupian (Kazanian) Another aspect of this Malaysian fauna is time. Since then, two localities have been dis- the presence of a number of species (18) that covered which have diverse gastropod as- are siphonate; usually only one or two species semblages in the Late Permian. A fauna col- ofthe siphonate Orthonema would be found lected by Richard Grant in the highest in a normal Upper Paleozoic fauna. There Permian (Dzulfian) ofGreece contains a nor- has been discussion that gastropods began mal marine fauna but with numerous species invading the infaunal realm by Jurassic time of zygopleurids and other mesogastropods (Signor, 1982). It has been suggested that this which have a distinct Triassic appearance. In was in response to the evolution ofpredators. several places in south China in the latest The development of siphons was an impor- Changxingian there are diverse gastropod tant factor in allowing migration into infau- faunas (Wang, 1978; Pan, 1983). Thus the nal niches. I am suggesting that the move- gap between the Middle Permian and Middle ment into the infauna was in response to the Triassic normal marine faunas has been origin ofpredators in the Permian rather than greatly reduced. the Jurassic. Probably later in the Mesozoic My conclusions are basically the same as there was a secondary migration of some si- in 1973. While there are extinctions at the phonate groups into the epifaunal realm to end of the Permian (such as most of the bel- account for the epifaunal siphonate forms lerophontids), many Paleozoic families sur- found today. Prodiozoptyxis, to illustrate, has vived into the Triassic, finally becoming ex- by far the deepest siphon of any known Pa- tinct at the end of the Triassic when a major leozoic genus and even the trochid Eucy- extinction occurred in the Mollusca. I would closcala has an incipient siphon and varices amend my view that most of the extinction suggesting an infaunal habitat. took place at the end ofthe Middle Permian. The presence of diverse faunas in China right at the boundary of the Triassic means ADDENDUM that the extinction that took place at the close Several taxa of importance were discov- of the Permian was a sudden event. This is ered in processing bulk loaned material for reinforced by the presence at many localities return: (1) A low-spired trochiform species in China ofan iridium "spike" in a boundary with a single spiral rib near the suture, a spiral clay, recently discovered by a Beijing re- rib on the periphery, and a concave outer search group. The extinction of gastropods whorl face was recognized as a species of was not, however, as profound at the Permo- Glyptospira Chronic, 1952, which is a mi- Triassic boundary as it was at the Triassic- crodomatid trochid, figure 58. (2) A new Jurassic boundary when most ofthe families species ofphymatopleura Girty, 1939, which were replaced by the cosmopolitan families is distinguished by having dominant spiral that dominate today's seas. Thus again we ornament with subdued collabral ornament, have the quandry of many groups that sur- figure 59, and Worthenia deKoninck, 1883, vived the meteoritic holocaust that the Al- a low-spired form quite distinct from other varez theory pictures. described species, figure 60; both of these Some of the new species described herein pleurotomarians were reported in Part 1 of have convergent counterparts in the Western this series (Batten, 1972). (3) A new species Hemisphere Pennsylvanian of Lower Car- of Mourlonia deKoninck, 1883 (Pleuroto- boniferous. Delpey (1941) alluded to this mariina), figure 61, has a shape much like M. 38 AMERICAN MUSEUM NOVITATES NO. 2829

~~~~~~~~~~~~6 1 60 FIGS. 60-61. 60. Worthenia sp., AMNH 42920, oblique side view, x 17. 61. Mourlonia sp., AMNH 42922, side view, X 10. talboti (Dickens), 1963 (reported in Part 1, by the lower margin of the selenizone and a Batten, 1972, p. 14) but with the collabral lower spiral thread that marks the boundary ornament greatly subdued and with strong of the base. The base is convex with numer- spiral costae on the upper whorl surface and ous evenly developed spiral threads. The col- base. (4) A new species of Glabrocingulum, labral elements are very faint. The umbilicus a flat, tabulate form with a strikingly wide occupies about one-third the area ofthe base umbilicus, is so unique that it is described and the spiral threads continue along the sides. below: The umbilical sutures are sharply defined. There is a slight thickening ofthe columellar Glabrocingulum (Glabrocingulum) lip. umbilicatus, new species DISCUSSION: The fine spiral threads which Figures 62a, b dominate the ornament pattern are similar DIAGNOSIS: Very low-spired trochiform to those found in G. (Stenozone) brennensis shells with a very wide umbilicus revealing (Reed) but the more important features such internal sutures and ornament to the early as the presence of an alveozone, subsutural whorls. Whorls embrace at the lower margin trough, and well-defined selenizone margins of the selenizone. Sutural trough is narrow clearly indicate its affinities with G. (Glabro- and deep. The upper whorl face is convexo- cingulum). Many species have an umbilicus. concave with collabral nodes on the upper For example, several undescribed species convex portion. Fourteen equally developed from West Texas and Greece have a modest spiral threads intersect the collabral nodes umbilical opening, but none are broad enough with weakly to reveal the earliest whorls as in this species; formed interference develop- also, no other known species has dominant ment. The slightly concave outer whorl face spiral ornament. One specimen. (here the same as the alveozone) is marked MEASUREMENTS: Holotype AMNH 42921, H 10.1 mm, W 6.0 mm, SpAng 1200 ETYMOLOGY: Umbilicus, Lat. navel.

LITERATURE CITED Batten, R. L. 1966. The lower Carboniferous gastropod fau- na from the Hotwells Limestone of Compton Martin, Somerset. Pts. 1 and 6 2 ..... 2. Palaeont. Soc. Monographs, Publ. 509 and 513, 109 pp., 10 pls. 62b 1972. Permian gastropods and chitons from FIG. 62. Glabrocingulum (Glabrocingulum) Perak, Malaysia. Part 1. Chitons, bel- umbilicatus, holotype, AMNH 42921; (a) side lerophontids, euomphalids and pleu- view, x 3; (b) oblique basal view, x 3.5. rotomarians. Bull. Amer. Mus. Nat. 1985 BATTEN: PERMIAN GASTROPODS 39

Hist., vol. 147, art. 2, pp. 1-44, figs. 1- Hoare, R. D., and M. T. Sturgeon 52. 1978. The Pennsylvanian genera Cyclozyga 1973. The vicissitudes of the gastropods dur- and Helminthozyga and the classifica- ing the interval of Guadalupian-Ladi- tion of the Pseudozygopleuridae. Jour. nian time. In Logan, A (ed.), The Perm- Paleont., vol. 52, no. 4, pp. 850-858, 2 ian and Triassic systems and their pls. mutual boundary, Can. Soc. Pet. Geol., 1981. Pennsylvanian pseudozygopleurid gas- Sp. Pap., pp. 596-607, 5 figs. tropods from the Appalachian Basin. 1979. Permian gastropods from Perak, Ma- Jour. Paleont., vol. 55, no. 6, 3 pls. laysia. Part 2. The trochids, patellids and Horny, R. neritids. Amer. Mus. Novitates, no. 1955. Palaeozygopleuridae nov. fam. (Gastro- 2685, pp. 1-26, figs. 1-33. poda), Ze st'redoceskeho siluru. Ustired 1984. The calcitic wall in the Paleozoic fam- Ustavu Geol. Sborn., odd. Paleont., vol. ilies Euomphalidae and Platyceratidae, 21, pp. 17-143, pls. 2-11. (Archeogastropoda). Jour. Paleont., vol. Houbrick, R. 58, no. 5, pp. 1185-1192, 2 figs. 1979. Classification and systematic relation- Bayer, F. L., and G. McGhee ships of the abyssochrysidae, a relict 1984. Iterative evolution in the middle Juras- family ofbathyal snails (Prosobranchia: sic ammonite faunas of wesern Ger- Gastropoda). Smithson. Contr. to Zool., many. Lethaia, vol. 54, no. 2, pp. 544- no. 290, 21 pp., 11 figs. 566. 1981. Anatomy, biology and systematics of Cossmann, M. Campanile symbolicum (Gastropoda: 1909. Essais de paleoconchologie comparee. Prosobranchia). Malacologia, vol. 21 Livr. 8, pp. 1-287, pls. 8. (1), pp. 263-289. Cox, L. R., and J. B. Knight Huddleston, W. 1960. Suborders of Archeogastropods. Jour. 1889. British Jurassic Gasteropoda, Part 1, no. Malac. Soc. London, vol. 33, pp. 17- 3. Gasteropoda of the Inferior Oolite, 18. pp. 137-192, pls. 7-11. Delpey, G. Jones, C. R., D. J. Gobbett, and T. Kobayashi 1941a. Les Gasteropodes Permiens de Cam- 1966. Summary offossil record in Malaya and bodge. Jour. de Conch., vol. 84, pp. 255- Singapore, 1900-1965. Geology and 278 and 346-369. Palaeontology ofSoutheast Asia, vol. 2, 1941b. Histoire du genre Camapanile. Ann. de pp. 309-359. Paleont., vol. 24, pp. 3-25. Kittl, E. 1942. Les Gasteropodes Permiens de Cam- 1893. Die Gastropoden des Schichten von St. bodge. Ibid., vol. 85, pp. 50-83. Cassian der Sudalpinen Triassic. Ann. Gemmellaro, G. G. der K. K. Naturhistorischen Hofmu- 1889. La fauna dei calcari con Fusulina della seums, Bd. 7, pp. 35-97, pls. 5-9. valle del fiume Sosio nella provincia de 1894. Die Gastropoden des Schichten von St. Palermo. Fasc. 2, Nautiloidea-Gastro- Cassian der Suidalpinen Triassic. Ann. poda, pp. 97-182, pls. 11-19. der K. K. Naturhistorischen Hofmu- Grabau, A. W. seums, Bd. 9, pp. 143-275, pls. 4-12. 1931. The Permian ofMongolia. Nat. Hist. of Knight, J. B. Mongolia, Amer. Mus. Nat Hist., vol. 1930. The gastropods of the St. Louis, Mis- 4, pp. 1-665, 35 pls. souri Pennsylvanian outlier: the pseu- Haas, 0. dozygopleurinae. Jour. Paleont., vol. 4 1953. Mesozoic invertebrate faunas of Peru. (suppl. 1), 89 pp., 5 pls. Bull. Amer. Mus. Nat. Hist., vol. 101, 1934. Ibid., vol. 8, no. 4, pp. 433-447, pls. pp. 1-321, 18 pls. 56-57. Harper, J. A. Knight, J. B., R. L. Batten, and E. L. Yochelson 1981. The use-misuse of Ianthinopsis Meek 1960. Part I. Mollusca. In Moore, R. C. (ed.), and Worthen, 1866 (Mollusca: Gastro- Treatise on invertebrate paleontology. poda). Jour. Paleont., vol. 59, pp. 180- Geol. Soc. Amer., Univ. Kansas Press, 185. pp. I1-I351, figs. 89-216. Hoare, R. D. Koken, E. 1980. New Pennsylvanian gastropods from 1892. Uber die Gastropoden der Rothen Ohio. Jour. Paleont., vol. 54, no. 5, pp. Schlernschichten nebst bemerkungen 1035-1040, 1 pl. uber verbreitung und herkunft einiger 40 AMERICAN MUSEUM NOVITATES NO. 2829

triassischen Gattungen. Neus. Jahrb. collections of the South African Mu- Min., Bd. 2, pp. 25-36. seum: families Abysochysidea, Ocory- Koninck, L. G. de thidae, Haliotidae and Tonnidae. Ann. 1883. Faune du calcaire Carbonifere de laBel- South Afr. Mus., vol. 25, no. 1, pp. 77- gique. Gasteropodes. Ann. Mus. Roy. 83, 4 pls. d'Hist. Nat. Belg., vol. 8, pp. 1-240, pls. Waagen, W. 22-54. 1880. Productus limestone fossils. Palaeont. Laube, G. C. Indica, Mem. Geol. Surv. India, ser. 13, 1869. Die Fauna der Schichten von St. Cas- vol. 1, pt. 2, pp. 73-183, pls. 7-16. sian. K. Akad. Wiss., Denkschr., Abt. Wang, G. G. 2, Bd. 28, pp. 29-94, pls. 21-28. 1978. Gastropods. In Atlas offossils ofsouth- Mansuy, H. west China, Guizhou Volume, Pt. 2. 1912. Etude geologie du Yunnan oriental. 2e Carboniferous to Recent, Guizhou partie. Paleont. Indochina, Service Working Team (eds.), pp. 394-413, pls. Geologique. Mem. 1, fasc. 2, pp. 1-146, 108-129. 25 pls. Wang, H. J., and Y. Xi 1914a. Faunas des calcaire a Productus de 1980. Late Permian and early Triassic gastro- l'Indo-chine. 2e ser.: Indo-china. Ibid., pods of western Guizhou. In Stratigra- fasc. 2, pp. 1-190, 10 pls. phy and Paleontology of Upper Perm- 1914b. Nouvelle contributions a la Paleonto- ian coal-bearing formations in western logie du Yun-nan. Ibid., Mem. 1, vol. Guizhou, Nanking Instit. of Geol. and 3, fasc. 2, pp. 1-12, 1 pl. Paleont. (eds.). Beijing, China Science Pan, Yin Tang Press. 1983. The character of the geologic and geo- Wenz, W. graphic distribution ofgastropods in the 1938. Gastropoda, Allgemeiner Teil, und Yangtze district. Acta Paleont. Sinica, Prosobranchia. In Schindewolf, 0. (ed.), vol. 22, pp. 1441-1447. Handbuch der Palaozoologie, Bd. 6, Teil Pchelintsev, V. F. 1, pp. 1-240. 1968. Mesozoic Murchisoniata of the Cri- Winters, S. S. mean Highlands. Internatl. Geol. Rev., 1963. Supai Formation (Permian) of eastern vol. 10, no. 11, 46 pp., 8 pls. Arizona. Geol. Soc. Amer., Mem. 89, Signor, P. W. pp. 1-99, 9 pls. 1982. Resolution of life habits using multiple Yin, H. F. morphological criteria. Paleobiol., vol. 1983. Uppermost Permian Pectinacea from 8, no. 4, pp. 378-388. South China. Riv. Ital. Paleont., vol. 88, Taylor, D. W., and N. F. Sohl no. 3, pp. 337-386. 1962. An outline of gastropod classification. Yochelson, E. L. Malacol., vol. 1, no. 1, pp. 7-32. 1956. Permiangastropodsofthesouthwestern Thiele, J. United States, pt. 1. Bull. Amer. Mus. 1931. Handbuch der Systematischen Weich- Nat. Hist., vol. 10, art. 3, pp. 173-260, tierkunde, Bd. 1, 778 pp. Jena, Gustav pls. 9-24. Fischer. Tomlin, J. R. 1927. Reports on the marine Mollusca in the

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