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Neogene to recent Naticidae (Mollusca : Gastropoda) of the eastern Pacific

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Neogene to recent Naticidae (Mollusca : Gastropoda) of the eastern Pacific

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Content NEOGENE TO R E C E N T NATICIDAE (M OLLUSCA GASTROPODA) O F THE E A ST E R N PACIFIC by Louie N ick M a rin c o v ic h , Jr. A D iss e r ta tio n P r e se n te d to the FA C U L T Y O F THE GRADUATE SCHOOL UNIVERSITY OF SO UTH ER N CALIFO RNIA In P a r tia l F u lfillm e n t of the R eq u irem en ts fo r the D e g r e e DOCTOR O F PHILOSO PH Y (G eo lo g ica l S c ie n c e s ) June 1973 INFORMATION TO USERS This material wm produced from a microfilm copy of the original document. While the moat advanced technological meant to photograph and reproduce this document have baen used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The si(pi or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing paga(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus causa a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper loft hand comer of a large dieet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may bo ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Soma pages may have indistinct print. Filmed as received. Xorox Untvorslty Microfilms MO North Zoofe Hoad Ann A fter. MtcMgan 401 O S p- 73-30,021* MARINCOVICH, Jr., Louie Nick, 1943- NEOGENE TO RECENT NAT1CIDAE (NOLLUSCA: GASTROPODA) OF THE EASTERN PACIFIC. University of Southern California, Ph.D., 1973 Paleontology University Microfilms, A X E R O X Company, Ann Arbor, Michigan i THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. UNIVERSITY O F SOUTHERN CALIFORNIA THE GRADUATE SCHOOL UNIVERSITY PARK LO S A N O E L U . C A LIFO R N IA * 0 0 0 7 This dissertation, written by Louie Nick Marincovich, Jr. under the direction of A i s . . . . Dissertation C om mittee, and approved by all its members, has been presented to and accepted by The Graduate School, in partial fulfillment of requirements of the degree of D O C T O R O F P H I L O S O P H Y \J Dtmm Date June.1973 CONTENTS Page ABSTRACT.............................. xi INTRODUCTION........................... 1 ACKNOWLEDGMENTS ......................... 4 METHODS AND FORMAT ...................... 6 TAXONOMIC PROCEDURES.................... 11 BIOGEOGRAPHY AND PALEOBIOGEOGRAPHY....... 15 BIOSTRATIGRAPHY AND PHYLOGENETIC RELATIONS . 35 SHELL-OPERCULAR RELATIONS .............. 46 BORING................................ 48 SEXUAL DIMORPHISM....................... 52 SYSTEMATIC ACCOUNT...................... 54 Family Naticldae Gray....... . ......... 54 Subfamily Ampullospirinae Cox ......... 57 Genus Amauronala Morch ............ 58 Amflyrongip Islandlea (Gmelin) .... 59 Subfamily Folinicinae Finlay & Marwlck • 66 Genus Pollnlcea Montfort ........... 68 Subgenus Pollnlcea Montfort, aenau atrlcto....................... 70 ro lln lw (£aUol£fift) ufesc (Valenciennes).............. 70 li Pose Pollnlcea (Pollnlcea) oanaaaenala (Recluz)...................... 78 Pollnlcea (Pollnlcea) Ui^mWV9 (Philippi).................... 80 Pollnlcea (ffilloLfifift) OUA (Broderip & Sower by)..................... 83 Pollnlcea (Pollnlcea) rayjdvg (Souleyet).................... 88 Pollnlcea (Pollnlcea) Hfagsjatufl (Griffith & Pldgeon)........... 90 Subsenua Euaoire Agassiz ........... 93 Pollnlcea (Biaolra) aaHanoi Dali . . 94 Pollnlcea (Euaoira) dlabloenala (Clark)....................... 100 Pollnlcea (Euaoira) lewlail (Gould) . 102 Pollnlcea (Euaplra) draconla (Dali) . no Pollnlcea (Euaplra) oallldua (Broderip & Sowerby) ........... 113 Pollnlcea (Euaplra) vlctorianua Clark & Arnold................ 120 Pollnlcea (Euaoira) aaulanua Dali . . 124 Pollnlcea (Euaoira) Dali......................... 126 Pollnlcea (R»pUa) IttgrUWg Dali . 128 111 Page Pollnlcea (Euaoira) part paw? Dali . 129 Pollnlcea aookenala Clark & Arnold . 131 Pollnlcea olvmoldll Reagan ....... 133 Subgenua Hvoterlta Woodring ......... 134 Pollnlcea (Hvoterlta) teHffflldSg (Gray).................... 135 Subgenua Mammilla Schumacher ....... 139 Pollnlcea (Mammilla) sfipcas (Philippi)................. 140 Genua Neverita Rlsao............. 143 Subgenus Neverita Rlsao, aenau atrlcto 144 Neverita (Neverita) nana (Moller) . . 144 Neverita (Neverita) klrkenals (Clark) 148 Neverita (Neverita) polltlana (Dali) 152 Neverita (Neverita) new species . . . 153 Subgenua Gloaaaulax Pllsbry ......... 156 Neverita (Gloaaaulax) anderaonl (Clark)...................... 156 Neverita (Gloaaaulax) reclualana (Deahayea)................. 164 Neverita (Gloaaaulax) lameaae Moore . 182 Genua Callnatlclna Burch & Campbell • • . 184 Callnaticlna oldrovdll (Dali).... 185 Genua Bui bus Brown............... 187 lv Page Buibus fragllls (Leach) ............. 189 Genua Chorlatea Carpenter............. 194 Cfrgrlatgg carpenterl Dali ........... 195 Chorlstea new species .............. 197 Subfamily Sininae We n z................ 199 Genua Slnum RBding ........... 200 Slnum cvnba (Menke) ....... • • • * 201 Sinua gray! (Deshayes) ............. 206 Slnum acopuloaua (Conrad)......... 208 Sinua debile (Gould)......... 216 Sinua noveali Dali................ 219 Sinua sanctllohannia (Pilabry & Lowe) 221 Sinua perrini (Arnold)............. 223 Genua Eunatlcina Fischer........... 224 Eunatlclna InaculPta (Carpenter ... 226 Subfamily Naticinae Gray.............. 229 Genus Natlca Scopoli................ 230 Subgenus Natica Scopoli, sensu atrlcto........................ 233 Hfltlfia (tiailaa) sl&£kl Etherlngton . 234 Natica (Natlca) inexoectana Olsson . 237 (Natlca) McLean . . 238 (Natlca) juikMl Addlcott . . 240 Page Subgenus Natlcarlus Dumerll ....... 242 Natlca (Natlcarlus) chemnltzll Pfeiffer, nomeo Inquirendum . . . 243 Natlca (Natlcarlus) unlfasclata Lamarck...................... 248 Natlca (Natlcarlus) othello Dali . 252 Natica (Natlcarlus) Wislren?!? Hertlein U Strong............. 255 Natlca (Natlcarlus) collma Strong & Hertlein.................. 258 Natlca (Natlcarlus) schethra Dali . 261 Natica (Natlcarlus) brunneollnea McLean...................... 264 Natlca (Natlcarlus) aravl Philippi 267 Natlca (Natlcarlus) oosuncula Hanna & Hertlein............. 270 Natlca (Natlcarlus) teftlandl Hanna & Hertlein.................. 273 Subgenus Lunala Berry ............. 275 Natlca (Lunaia) lunarla (Berry) . . 276 Subgenus Glvpheplthema Rehder .... 278 Natlca (Glvpheplthema) ldlopoma Pllsbry & Lowe.............. 279 vl Page Subgenus s»isthan.iax Morch . 283 Natlca (Stlanaulax) broderlplana Reluz.............. 284 Natlca (Stlanaulax) elenae Reluz 289 Subgenus Tectonatlca Sacco .... 294 Natlca (Tectonatlca) lanthostona Deshayea ........... 295 Natlca (Tectonatlca) sateooensla (Addlcott)........ 300 Subgenus Crvotonatlca Dali .... 301 Natlca (Cryptonatlea) clausa Broderip & Sowerby. 302 Natlca (Crvptonatlca) ore&onensls (Conrad).......... 316 Natlca Inezana Conrad ............. 320 Natlca saxea Conrad, noaen dublun . • 322 Rejected and Indeteralnate Taxa...... 324 REFERENCES CITED................. 347 vll LIST OF FIGURES Figure Page 1. Distribution of naticld species in eastern Pacific latitudes and molluscan provinces. Unconfirmed range data shown as dotted lines................................ 16 2. Stratigraphic occurrence of eastern Pacific Natlcidae. All Pleistocene species are extant. Data based on museum specimens are shown in solid lines, probable oc currences as dashed lines, and less reli able data as dotted lines............... 36 3. Phylogenetic relations of Neogene Natlcidae of western North America................ 42 4. Comparison of height to diameter ratios of Atlantic and Pacific specimens of AnffWr*?Pg< f f is land lea (Gmelln). Based on specimens from several institutions • • • • 64 5. Comparison of height to diameter ratios of Neverita recluslana (Deshayes) and Neverita alta (Arnold).Based on Recent LACM specimens.......... 175 viii LIST OF TABLES Table Page 1. Homologous tropical eastern Pacific and Caribbean natlclds .................. 27 2* Geographic occurrence of Miocene Natlcidae in western North America ... 31 3. Geographic occurrence of Pliocene Natlcidae in western North America ... 33 Ix LIST OF PLATES Plate Page 1. Fossil and Recent specimens ................. 326 2. Fossil and Recent specimens ................. 334 3. Fossil and Recent specimens..................340 x ABSTRACT The Miocene through Recent species of the marine gastropod family Naticidae in the eastern Pacific have been reviewed. Recent species have been studied from the Arctic Ocean to northern Peru, and fossil species have been studied only In western North America. Fifty-seven species are included in this study* of which 30 have fossil records and 13 are extinct. In order of their taxonomic value* the most signi ficant features of naticids are umbilical morphology* shell form and sculpture* and opercular sculpture. Radular dentition Is generally useful only at the generic level for separating taxa. Using these criteria* the family in this region is divided into nine genera and 13 sub-genera. Some genera, subgenera and species of Recent naticids are restricted to particular marine hydro- climates * so that they may be used by paleontologists to Interpret past environments• Several Neogene species evolved or became extinct in response to climatic fluctuations of the Late Tertiary. Neogene naticids were most abundant and diverse in the Middle Miocene* during a short-lived warm interval, and the Neogene species of Oregon and Washington existed in a xl cooler hydrocllmate than those of central and southern California* A number of species in Neogene sediments of each region are useful as zonal fossils* The modem eastern Pacific naticld fauna is largely derived from endemic Neogene ancestors. One liv ing species Is a migrant from the Indo-Pacific region and three arrived from the North Atlantic. There are at least seven pairs of homologous species between the tropi cal eastern Pacific and Caribbean faunas. A phylogeny for eastern Pacific naticids is proposed and discussed. There is an apparent functional relationship be tween the shells and opercula of Natlca (Stl gnaaulax) broderlplana Recluz and N. (S.) elenae Recluz, which has not previously been described in this family. xll INTRODUCTION The molluscan family Natlcidae la comprised of a large and similar group of marine gastropods of world wide distribution, living in habitats from the intertidal to the deep sea* Naticids first occurred in the early Jurassic (Carriker & Yochelson, 1968i Sohl, 1969a), but are most abundant in Cenozolc sediments and Recent seas* Because this family Is morphologically conservative, yet contains a large number of species, it is beset with a number of difficult taxonomic and systematic problems* The present study is a taxonomic review of the Recent Natlcidae from the Arctic Ocean to northern Peru, and also of the Miocene through Pleistocene species of western America. Recent clrcumboreal species are included, but the Recent and late Cenozolc species of western South America have largely been excluded for lack of available specimens* Naticids are routinely and abundantly present in the rich Cenozolc molluscan faunas of western North America, However, their utility as zonal fossils has been limited by lack of comparative studies and poorly documented stratlgraphlc occurrences. A major objective of the present study has been to enhance the usefulness 1 2 of naticids In stratlgraphic correlations, in addition to simplifying the taxonomy and classification of fossil and Recent species* Neither the Recent nor late Cenozolc naticids of the eastern Pacific have ever been reviewed critically. Prior to this century, worldwide Recent accounts of the broadly defined genus Natica were compiled by Philippi <1849-1853), Reeve (1855), Sowerby (1883), and Tryon (1886). Monographs of Slaaretus fsinunl were done by Reeve (1864), Sowerby (1882), Welnkauff (1883), and Tryon (1886). These works are helpful for providing illustra tions of many previously unfigured species, but they are largely uncritical accounts and did not clarify or advance study of the taxonomy and systematics of the family. In the case of Tryon (1886), his ill-considered lumping of distinct species only confused later workers. The living tropical eastern Pacific naticids have recently been surveyed by Keen (1958, 1971) as part of an extensive faunal summary of the Panamic molluscan province, although critical evaluation of the family was not within the scope of that work. Dali (1921) listed the nominal Recent species from the extra-tropical northeastern Pacific, Keen & Bentson (1944) did the same for Tertiary species of California, and Weaver (1942) gave an illustrated ac count of Tertiary naticids of Oregon and Washington, all as part of largerfaunal accounts. Dali (1909b) made some 3 useful generic consents In his account of Pliocene species of Oregon, and Grant & Gale (1931) did the sane for genera and species from the Pliocene and Pleistocene of California. Natlcidae of the cool-water Peruvian molluscan province) to the south of the region considered here, are very poorly known. These species were Included In this study during Its early stages, but the obscure literature and almost complete lack of specimens from that area dis couraged the effort. Extensive collecting In Chile and Peru Is necessary before the Recent and fossil naticids there can be treated. The Recent nominal species of the Peruvian province are listed by Dali (1909a), and the Chilean Cenozolc species are cataloged by Philippi (1837). Works dealing with naticids from elsewhere than the eastern Pacific are mentioned at appropriate places In the text. ACKNOWLEDGMENTS I would like to thank Dr. William H. Easton of the University of Southern California for guidance and en couragement * for many lengthy discussions of difficult problems* and for evaluation of my manuscript. Dr. James H. McLean of the Natural History Museum of Los Angeles County allowed me access to his voluminous private notes and photographs» sorely tested my taxonomic ideas at times* and reviewed the final work* The manuscript was also evaluated and improved by criticism from Dr. Orville L. Bandy of the University of Southern California. Colleagues who provided access to collections in clude Dr. R. Tucker Abbott of the Delaware Museum of Natural History* Dr. Warren 0. Addicott of the United States Geological Survey* Dr. J. Wyatt Durham and Mr. Joseph Peck of the University of California at Berkeley* Dr. William K. Emerson of the American Museum of Natural History* Dr. A. Myra Keen of Stanford University* Dr. Robert Robertson of the Academy of Natural Sciences of Philadelphia* Dr. Peter U. Rodda of the California Academy of Sciences* Dr. Joseph Rosewater and Mr. Frederick Col lier of the United States National Museum* and Dr. Ruth D. 4 5 Turner and Dr. Kenneth J. Boas of Harvard University. Thanks are also due to many people who helped in small but significant ways* but who are too numerous to mention. This study was supported in large part by a grant from the Edwin C. Pauley Foundation. Travel expenses were partly financed by a grant from the Theodore Roosevelt Memorial Fund of the American Museum of Natural History* and completion costs were provided by the Sea Grant Program at the University of Southern California. METHODS AND FORMAT The mass of data and observations accumulated dur ing this study required a systematic approach to research efforts and note keeping. The initial step was to survey the principal Miocene through Recent literature of the eastern Pacific, searching for relevant species-level names. The original description of each taxon and all major published commentaries and critical citations for each species were photocopied and entered into loose-leaf notebooks. In this way, the Important literature on each species was kept at hand for evaluation in its original form. An attempt was made to locate the type specimens for each available taxon. I was successful In doing this for all of the fossils, but in many cases I have not seen type specimens of Recent species described by European workers. In nearly all cases where I located type material, the specimens were borrowed and kept with me during this study, making comparIsons between them con venient . Trips were made to the principal American in stitutions housing mollusk collections, during which all eastern Pacific naticids were examined, and in many cases were photographed and measured. Extensive notes were made 6 7 for especially interesting specimens* and a large number of specimens was borrowed for the duration of this study. Slightly fewer than 20*000 Recent and fossil specimens were examined and recorded in my notes. Areal and bathymetric limits given here are based on specimens I have seen* unless otherwise indicated* al though an effort was made to locate specimens upon which published range limits or unusual occurrences were based. Recorded stratlgraphlc occurrences are a combination of literature citations and my own observations (indicated by initials of the institution housing the specimens). The list of institutions visited for this study is given below* along with abbreviations used in the texti Academy of Natural Sciences* Philadelphia* Pennsyl vania - ANSP Allan Hancock Foundation* University of Southern California* Los Angeles* California (gastropod collection on permanent loan to Natural History Museum of Los Angeles County) - AHF American Museum of Natural History* New York* New York - AWH British Museum (Natural History)* London* Eng land - BM(NH) California Academy of Sciences* S«n Francisco* California - CAS 8 Delaware Museum of Natural History, Greenville, Delaware - DMNH Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts - MCZ Natural History Museum of Los Angeles County, Los Angeles, California - LACM San Diego Natural History Museum, San Diego, Cali fornia - SDNHM Stanford University, Stanford, California - SU United States Geological Survey, Menlo Parle, Cali fornia - USGS United States National Museum of Natural History, Washington, D. C. - USNM University of California at Berkeley, California - UCB University of California at Davis, California - UCD University of Southern California, Los Angeles, California - USC Specimens in the private collections of Dr. S. Stillman Berry and Dr. Donald Shasky, both in Redlands, California, were also examined. Information in the "Systematic Account” Is present ed in a fairly standard format. The headings and con ventions used for each species are as followsi 1. Species-level names are presented in their original combinations) with a chronological list of authors who used them* The synonymies are Intended to be as thorough as possible) but for commonly cited and wide* spread species such as Natica (Crvptonatlca) clausa Broderip & Sowerby, or Slnum scopuloaum (Conrad)• many minor citations are omitted. An attempt was made to list all works containing taxonomic innovations* Standard taxonomic conventions are employed* In some casesi sup plementary comments are Inserted in brackets after a name citation. 2. The description of each species is in a standard format* with variations to suit particular morphologic features* Average sizes are based on measure ments with calipers of a large number of specimens) and the collection number is given for the largest known specimen of each taxon* Because natlcid radulae do not generally bear specific characters* they are described under generic headings except in a few cases. The oper- cula of a few Recent species and nearly all of the fossils are unknown* but are assumed to be chltinous or calcareous according to shell form. The number of specimens examined la given. 3* Records of geographic occurrence and ecology are based on specimens I have seen* unless otherwise stated. The major range end-points are given from north to south* and located in degrees of latitude along coast- 10 lines* or in degrees of latitude and longitude for is lands, 4. Stratlgraphlc occurrences are based on litera ture citations and specimens 1 have seen* the latter indicated by institutional abbreviations. Where the published ages of formations were in question* current age estimates were provided by Dr. Warren 0. Addlcott of the U. S. Geological Survey* 5. Type localities are given in modem ortho graphy if their Identity is certain* but many are listed as unknown pending examination of type material in European Institutions. Specific names are listed in their original form. 6. The location of type material is given for all certainly synonymous species* if it is known. The prob able location of many collections presumably containing the type material in question is often taken from Dance (1966). Specific names are listed in their original form. 7. A nomenclature 1 commentary is given for most species. 8. A discussion is given for each species* to elucidate Important morphologic variations* Justify the allocation of species-level taxa, or to explain aspects of each species not suitable to the outline format of the other headings. TAXONOMIC PROCEDURES The biological species concept, as described by Mayr (1963, 1969), should be the guiding principle for a taxonomic monograph of this sort. Under this concept, a species forms a reproductive community, an ecological unit, and a genetic unit, and is a group of interbreeding natural populations that is reproductlvely isolated from other such groups. The reproductive Isolation of a biological species thus results in a discontinuity of the genotype of the daughter species from that of the parent species. This in turn alters the morphology and other aspects of the phenotype, to produce a visually distinct species. Reproductive Isolation can be achieved without corresponding morphological change, and strong morpho logical differences may arise without reproductive Isola tion, however. In addition, the occurrence of hybridiza tion (breakdown of Isolating mechanisms) or geographic Isolates can cause taxonomic difficulty. Thus, the morphology of a supposed species might not be a reliable guide to Its status as a biological species. Fortunately, the experience of zoologists is that morphology is generally a reliable guide to biological 11 12 speclatlon (Mayr, 1969), and that assumption is made here. Despite close similarity between some species, the great majority of naticlds show a predictable range of varia tion. Some species-level taxa, however, show such a be wildering array of intergrading morphologies that it is a matter of choice whether to treat them as one or more species or subspecies. This is the case with species such a« (Glossaulax) reclusiana (Deshayes), Polinicee (Pollnlces) uber (Valenciennes) and associated species, and Natlea (Crvotonatlea) clausa firoderlp & Sowerby. For such species, other than simple morphological criteria are used to distinguish species, including variation within samples, ecology and paleoecology, areal distribution, and stratlgraphlc occurrence (Imbrle, 1957). This situa tion is more difficult to deal with in fossils. In in stances where the definition of discrete phenotypes is an Intractable problem, as in forms of f l . (G.) recluslana. it seems best to rely on a polytypic species concept, In which geographically representative forms once separated as monotyplc species are united under one name (Mayr, 1969*38). As Mayr notes, this leads to simplification of the system and restores to the species a definite bio logical meaning and homogeneity, because the awarding of species rank to local populations destroys the biological significance of the species category. Although we are still left with the problem of where to draw the line be 13 tween species, It is easier to deal with a few specimens that cannot be confidently assigned to a species than to deal with an unwieldy array of names based on local variations. The relative taxonomic and systematic Importance given to the hard parts of Natlcidae has varied greatly among workers. Although the shell Is usually considered to be of primary Importance, opercular sculpture also has been used as the basis for erecting genera. Radular dentition Is of some utility In distinguishing genera (Odhner, 1913i Thiele, 1931i Azuma, 1961i Oyama, 1969), but it is of almost no use In separating species, based on my own studies. In order of their taxonomic value, the significant features of naticlds are their umbilical areas, shell sculpture and form, and opercular sculpture. The operculum is of major Importance when making the basic distinction between Natlcinae and Follniclnae, but numerous species occur in which the calcareous opercula are indistinguishable. Shell sculpture Is uncommon In the family, and most often occurs as radial grooves that are nearly Identical among species. Sculpture is thus usual ly of only generic Importance. On the other hand, the shape of the umbilical callus and the degree to which it conceals the umbilicus is of critical generic and specific value. Callus morphology is relatively consistent at the species level or varies through a predictable range. En 14 vironmental effects on callus shape are not documented* but are possible In variable species such as Neverlta (Glossaulax) recluslana or Natlea (Crypt onatlea) clausa. Shell color Is described for each Recent species treated here* but It Is not emphasized. It seems best to des- crlbe natlcld taxa emphasizing shell morphology, because this is usually all there is to work with among fossils* Relying on features common to fossil and Recent species permits meaningful comparisons between living and extinct species. The generic treatment used here Is conservative* because the family is in need of a thorough worldwide revision. The subgenera used here have often been used as genera* but 1 feel that this is premature and also tends to conceal relationships between similar genus- level taxa. BIOGEOGRAPHY AND PALEOBIOGEOGRAPHY Figure 1 lists the Recent eastern Pacific naticids and graphically gives their distributions in degrees of latitude. This clearly shows that although the family is represented in all marine climatic zones* its greatest diversity occurs in the tropics. Only five species live in Arctic latitudes* whereas nine are well-established in the warm-temperate Californian province and 23 are found at the equator. The sharp drop in species diversity at 28°N» the boundary between tropical and warm-temperate hydroclimates* indicates the essentially tropical nature of the family, more so than does the decrease in species with the more severe climatic change from the tropics to the Arctic. The persistence of this essentially tropical family in high latitudes emphasizes its highly adaptive nature. Among naticids, the Naticinae in particular are a tropical group. The subgenera Natlea. Natlcarius. Lunais. Glvpheplthema and stianauia* are found exclusively in tropical waters, except for a short northward exten sion of Glvphepfrtlttf into the Californian province. Ex cept for Lunaia. which is not known elsewhere* these sub- 15 FIGURE 1 Distribution of naticid species In eastern Pacific latitudes and molluscan provinces* Unconfimed range data shown as dotted lines* 16 = * X fe* * > - 1 * o CO CO I I 1 1 111 ]) ] ilium 1 i I I i I i i N u b (KlW titlwlw Oll i m m m & & M H 1 1 1 jj i I 1 Iili 1 I ii i ' i i i 1 ! 1 f 1 ! i i li ill X to CL. 19 genera seen to be distinctly tropical groups In other regions also. These groups contain species with ranges extending to the southern limit of tropical water In northern Peru, or nearly so, but not Into the cool- temperate waters of the Peruvian province farther south. For the most part these species have colorful, relatively thin shells and highly sculptured opercula, and are of moderate size. These relations also apply to the Neogene naticids of the eastern Pacific, so that distinctly warm- water fossil assemblages have a preponderance of Natlclnae species and cool-water assemblages do not. The subgenera Cryptonatlea and Tectonatlca. each represented by one species In this study, are exceptions among the Natlclnae In living only In cool waters. Natlea (Tectonatlca) tenthostoma Deshayes lives only In the western Pacific today, but apparently evolved and lived In the eastern Pacific from the Early Pliocene to at least the Early Pleistocene. Its southernmost extent was In northern California, in the Middle Pliocene to Pleistocene, and It occurs sparsely In Pleistocene deposits of the Ber ing Sea area, Tectonatlca seems to be a largely tropical group elsewhere, although the generic name has not been used by many workers. Woodrlng (1957*88-89) mentions four Tectonatlca species from the Caribbean, so the group Is more abundant In the tropics, but further study may show It to be represented more extensively In middle latitudes. 20 Although Watlea (Crvptonatlca) clausa ranges to southern California, it Is definitely a cold-water species characteristic of Aleutian and Arctic waters. It occurs in southern latitudes by living at progressively greater depths in bottom water of Arctic temperature. The optimal hydroclimate of this species Is Indicated not only by its shift in depth range, but also by Its size range. Arctic specimens average about 30 ram in height and 28 mm in diameter, whereas those from southern California are about 13 ram in height and 11.5 mm in diameter, with the same number of whorls per specimen. The steady southward de crease in size parallels the increasing depth habitat and slight southward increase In bottom-water temperature, Indicating the essentially Arctic nature of the species. The Middle Miocene to Early Pliocene species f j * (£.) oreaonensls (Conrad), the only other Crvptonatlca In this area, lived in the cooler-water hydrocllmate of Oregon and Washington, but not in the subtropical Miocene marine basins of southern California. (Crvptonatlca) filfiuift occurs in cool-water fossil assemblages throughout the western coast of North America. Its earliest occurrence is in Lower Miocene beds in Alaska, then in Middle Miocene beds of Oregon, and in Upper Miocene beds of northern California and Washington. In Middle Pliocene strata it occurs extensively through out central California, and reached southern California 21 In the Late Pliocene* Paleontological evidence thus re veals the northern origin of £. (£«) clausa and £. (C.) oreaonenais and Indicates that the subgenus has long been established in cold hydroclimates. Fossil specimens of Crvptonatlca and Tectonatlca species have smaller and thinner shells than do Recent specimens. The largest fossil of £ 1 . (£•) clausa, from the Pliocene of northern Washington, is 47 mm high and 46 mm in diameter, and has a slightly thinner shell than an equivalent Recent specimen. The largest Recent specimen, from Alaska, measures 61 mm high and 54 mm in diameter. Similarly, the largest Recent specimen of £. (£•) lantho- stoma is 57.4 mm X 50.0 mm, whereas the largest fossil (Lower Pliocene of Oregon) measures 34.2 mm X 34.6 mm. The Middle Miocene to Early Pliocene species (£.) oreaonensia attained 37,1 mm X 29.6 mm, but averaged about 24 mm X 19 mm. The northern Natlclnae are thus distinct from the southern species, not only in thermal tolerance but also in general shell morphology. The Recent Crvptonatlca and Tectonatlca species are larger and more globose, with pro portionately thicker shells, more plainly ornamented oper- cula and more drab colors than their southern counterparts. These distinctions have existed since at least the Early Miocene and have been increasingly emphasized through time, Judging by the size differences between fossil and 22 Recent specimens of the sane species. The eastern Pacific Pollclnlnae as a group occur in a greater variety of narlne cllnatic zones than do the Natlclnae* although there is still sone segregation by subgenus. Species In the subgenus Pollnlces. £.£.» occur only in tropical waters * and some of then range throughout the eastern Pacific tropics. This subgenus seems to be essentially restricted to the tropics on a worldwide scale in Recent and Neogene faunas. Pollnlces (Pollnlces) irniflirgnnlIt (Recluz) does occur south of the tropics* in the cool-water Peruvian nolluscan province* but Its tax onomic relationships with several other species there are unclear. A taxononlc review of this species-complex night restrict the range of £. (£.) somewhat. The subgenus Eusolra shows a greater climatic range than Pollnlces. £.&• * but it is divided into a southern tropical group and a northern warm-temperate to Arctic group* which do not overlap in distribution. The southern group of four consists of obscurely known deep-water species represented by little more than primary type material. These may be related to the group which Hedley (1916151) commented on when he proposed Frisinatiosi "There is an Antarctic natlcold group which • • • amounts to about a dozen rather featureless species* all small* mostly uniform olive buff In colour* four whorls* a slightly raised spire* a caducous epidermis* comparatively 23 thin, unsculptured, except for Incremental striae, with out umbilical funicle or a callus pad at the Insertion of the right lip. Operculum corneous pauclsplral." On the other hand, the northern Eusplra group consists of three species that are all relatively common and range Into shallow water. Pollnlces (Eusplra) draconls (Dali) and £• (£•) lewis11 (Gould) are large forms (£. (£.) lewis11 attains 166 mm X 131 mm) that are closely related to each other and have geologic histories that can be confidently traced back to Late Miocene ancestors In California, They do not seem to be closely related to the other northern Eusplra. £. (£.) pallldus (Broderlp & Sowerby), or to the group of southern Euspiras, Pollnlces (Eusplra) pallldus first appeared during the Middle Miocene in California and possibly in Alaska and has no unequivocal Neogene ances tors, although It may be related to £. (£.) vlctorianus Clark & Arnold. It may also have been derived from the Oligocene species £. rectus Tegland, 1933, which occurs In Washington. The subgenera Mammilla and Hvoterlta are each represented by one tropical species In this region. Hvoterlta Includes only two species, the type-species £. (£.) hellcoldes (Gray) of the eastern Pacific, and the Miocene species £. (£.) nereldls (Maury, 1922) from the Dominican Republic and Costa Rica (Woodrlng, 1957). Mam- aUli Is represented only by £. (£.) caorae (Philippi) In 24 the eastern PacLfic, but contains a number of species in the western Pacific* the only certain Indo-Pacific fauna1 element among the eastern Pacific Naticldae. It is re ported as a Pleistocene fossil in southwestern Mexico (Palmer & Hertlein, 1936)* The subgenus Neverita. g.g., is essentially a northern group, with one obscure and one well-known species in this region. The species £ J . (fj.) nana (Moller) occurs as far south as Baja California, but does so by liv ing at progressively greater depths southward, to maintain itself in Arctic bottom water. The subgenus Glossaulax is represented only by J J * (£•) recluslana (Deshayes), which occurs in hydroclimates from cool-temperate to tropical. It is found throughout Neogene to Recent deposits in the eastern Pacific as a bewildering variety of intergrading forms and is very closely related or identical to Miocene species I have seen in collections from the Caribbean area, Central America and western South America south to Chile. It is also very similar to Oligocene species in California, but elucidating the origins of this species- complex must await future work. As elsewhere, sinnw species here are confined main ly to the tropics, although the most abundant species, £• leonulftaua (Conrad) lives largely in a warm-temperate en vironment and even ranges slightly into cool-temperate waters. It seems to have lived in an equally broad range 25 of environments from the Late Oligocene onward, during which it was a persistent faunal element. None of the living species of Panamic Slnums Is known as a fossil north of its present range* although £• deblle (Gould) has been erroneously reported from the Pleistocene of southern California. There are five naticids with circumpolar distribu tions » as seen in Figure 1. Of these* Natica (Crvptona tlca) clausa and Pollnlces (Eusplra) MlUdUfl occur In Miocene deposits in the eastern Pacific and in Plio- Pleistocene deposits in the northern Atlantic* suggesting a Pacific origin for both. Amauronala islandica (Gmelin), Buibus fra&llls (Leach) and Neverlta (HsXSEltfi) DfiOfi occur in Pleistocene beds of Britain but are unknown as fossils in the eastern Pacific* suggesting an Atlantic origin for these species. Each of these migrations between the Pacific and Atlantic apparently took place during the episodic Late Pliocene to Recent openings of Bering Strait* as documented by Hopkins (1967a* 1967b) and Durham & Mac- Neil (1967). The earliest opening of Bering Strait took place in about Late Miocene time* followed by closure and reopening during the Late Pliocene. The Bering seaway may have been intermittently open and closed since the Late Pliocene (Durham & MacNell* 1967i346). The naticld species noted here apparently migrated between oceans after the second major opening of Bering Strait* during 26 the Late Pliocene or later. There are thus three species at most that migrated into the eastern Pacific from the northern Atlantic. Expectably, a number of living eastern Pacific and Caribbean naticids is closely related* as shown in Table 1. Other species-pairs may be recognized once the Carib bean naticid fauna is thoroughly evaluated and compared to that of the eastern Pacific in greater detail. During most of Tertiary time* essentially a single molluscan fauna lived in the Central America region* and the present- day Isthmus was a series of Islands separated by east-west seaways. This arrangement persisted until the end of Miocene time (Emerson, 1967) or perhaps later (Woodrlng, 1966i Olsson, 1972), when tectonic uplift produced a con tinuous land connection between North and South America that has lasted until today. So, the modem tropical eastern Pacific and Caribbean molluscan faunas in large part have a common origin in the Central American Miocene fauna, and this is reflected in the number of homologous naticid species between the two modem faunas. The Ter tiary molluscan faunas of Panama and adjacent areas have been extensively treated by Woodrlng (1957, 1959, 1964, 1970). Pollnlces (Eusplra) lewlsil and HflYtElt* (filflg- saulax) recluslana are similar to £. (£.) heros (Say, 1822) and f i . (Neverits) duplicate (Say, 1822), re- Table 1, Honologous tropical eastern Pacific and Caribbean naticids Tropical eastern Pacific species Natlea (Stianaulax) broderlplana Nat lea (Stlftnaulax) elenae Natlea (Naticarius) cheanltzii Natlea (Glvpheplthena) idlopona Pollnlces (Pollnlces) otis (Pollnlces) uber 51BUS dfibllfi Honoloaous Caribbean species____ Nat lea (StlRiiaulax?) cavennenals Natlea (Naticarius) canrena Natlea (Naticarius?) narocfalensis Natlea (Glypheplthena) florldlana Pollnlces (Pollnlces) hepaticua Pollnlces (Pollnlces) lacteus Slnun persoectlvua 28 apectlvely, of the American east coast. The first pair of similar species do not have an obvious common ancestor in the Central American Miocene fauna, and they may have developed Independently on separate coastlines in response to similar environmental stresses. This seems likely» be cause the ancestry of £. (£.) lewis 11 can seemingly be traced to an early Miocene species endemic to western North America, £. (£.) gallanol Dali, The other two species, f j . (G.) recluslana and g. (£.) duplicate, are definitely related and represented by ancestors in the Central American Miocene. Woodring (1957) described £. (G,) recluslana xena from the Middle Miocene of Panama, noting that it was probably a migrant from the Pacific. He also mentions a series of naticids related to f t , (G.) recluslana that persisted through the Late Miocene. It is certainly from this stock that the present-day £ £ , (£},) duplicate of the American east coast evolved. As noted above, elucidating the relationships between the many species and intergradlng forms of the Oligocene to Recent £. (G.) recluslana lineage would require more study. Comparisons between the temperate faunas of the eastern and western Pacific are difficult to make, because the Japanese literature on the family is scattered and the naticids there are in need of revision. The tropical eastern Pacific naticid fauna is thus largely derived from a more cosmopolitan Central American 29 Miocene fauna, owing only one species to migration from the Indo-Pacific region. This agrees well with the general trend noted by fliers on (1967) for the eastern Pacific mollusk fauna to have a large Caribbean fauna1 element and a minor Indo-Pacific one. As noted above, there are relatively few naticids that were introduced across the Arctic region from the North Atlantic. Thus, the tropical eastern Pacific naticids have a more cosmo politan ancestry, and the warm-temperate to cool- temperate species are more provincial. As will be shown below, the lineages of many temperate-water naticids can be traced through endemic Pliocene or Miocene ancestors. The Tertiary molluscan faunas of California, Oregon and Washington are rich and fairly well documented, and cam be used to delineate climatic fluctuations (Addicott. 1969). In turn, the warm- and cool-temperate Neogene naticids of this region have a better documented history than those from elsewhere in the eastern Pacific, and can be used to supplement paleocllmatlc and paleobiogeographic evidence derived from more general faunal studies. Miocene and Pliocene faunas of western North America display clear differences between the warm-water naticids of central and southern California and the cooler-water species of the Tertiary embayments of coastal Oregon and Washington, although the boundaries of hydrocllmatlc zones cannot be drawn from this evidence alone. As seen 30 In Table 2, four Miocene species are characteristic of southern and central California (with a fifth, NeverIta (Glossaulax) recluslana. after the Middle Miocene), another three are restricted to Oregon and Washington, and eight Miocene species are common to both regions. Of the four southern species, Slnum oerrlnl (Arnold) and Natlea (Naticarius) posuncula Hanna & Hertleln have generic affinities more characteristic of Recent warra- water faunas. Among the northern naticids, surprisingly, two of the three species are in the subgenus Natlea. £•£•, which is restricted to the tropics of the modem eastern Pacific. The third species is in the subgenus Crvpton atlca. the only living species of which has Arctic af finities. The eight Miocene species common to both northern and southern areas are included in six different genera and subgenera, some of which, such as Crvptonatlca and Naticarius. are not found together in Recent seas. The Miocene faunas of these two areas are thus more cos mopolitan at the generic level than are their derivative modem faunas. This condition may reflect the fact that climatic gradients were less pronounced during most of Tertiary time than they are today in the same region. Be cause of the generally warmer climate throughout the eastern Pacific during the Miocene, molluscan faunas were somewhat less provincial due to climatic factors, although the development of separate basins tended to partially Table 2. Geographic occurrence of Miocene Natlcldae In western North America Species restricted to southern Species restricted to Oregon and and central California_______________________Washington___________________ Natlca (Naticarius) posuncula Natlea (Natlca) clarkl Pollnlces (Eusplra) dlabloensls Natlca (NatIp*) vfikeal Never!ta (Glossaulax) andersonl Natlca (Crvptonatlca) oregonensls Slnum perrlnl Species common to both of the above regions Natlca (Naticarius) teglandl Neverlta (Neverlta) klrkensls Natlca (Crvptonatlca) clausa Neverlta (Glossaulax) jamesae Pollnlces (Eusplra) gallanol Pollnlces (Eusplra) vlctorlanus Neverlta (Glossaulax) recluslana Slnum scopulosum 32 isolate faunas. Species could thus range into a wider variety of hydrocllmates than is possible today with the more sharply defined climatic boundaries between faunal provinces. For example* there is no faunal break in the Miocene of this region comparable to that between the Panamic and Californian molluscan provinces of today (Figure 1). As seen in Figure 2* numerous naticids became ex tinct at the end of the Middle Miocene* in response to the increasing rate of climatic deterioration that culminated in the Pleistocene glacial stages. In addition* some species that survived into the Late Miocene* such as Neverlta (Glossaulax) recluslana. did so only in Cali fornia basins and became extinct in the cooler waters of Oregon and Washington. Climatic cooling is reflected in the Pliocene naticid faunas of these northern and southern areas (Table 3), in which only one of the provinclally typical species* Natlca (Crvptonatica) oreaonensls (Con rad) of Oregon and Washington* is a survivor from the Miocene. In addition* Natlca (Tectonatlca) sataopensis Ad- dlcott of the northern fauna is the only Pliocene species in a subgenus characteristic of Recent warm-water faunas. All of the other species* including those common to both areas* belong to generic taxa more typical of extra- tropical hydrocllmates. For example* several of the nine species given in Table 3 belong to the subgenera Eusplra■ Table 3* Geographic occurrence of Pliocene Naticidae In western North America Species restricted to southern Species restricted to Oregon and and central California______________________ Washington___________________ Pollnlces (Eusplra) draconls Natlca (Tectonatlca) satsopensls Neverlta (Glossaulax) recluslana Natlca (Crvptonatlca) oregonensls Species common to both of the above regions Natlca (Crvptonatlca) clausa Pollnlces (Eusplra) lewlsll Natlca (Tectonatlca) lanthostoaa Pollnlces (Eusplra) pallldus Pollnlces (Eusplra) gallanol u > u > 34 Crvptonatlca or Tectonatlca. which are suggestive of warn-temperate to Arctic conditions today. it is not possible to describe differences ac curately in the Pleistocene naticid faunas between the northern and southern areas discussed here. Although several naticids are common in Pleistocene formations of the southern region, they are relatively rare to the north, because most northern Pleistocene molluscan as semblages represent rocky-shore habitats in which naticids did not live. BIOSTRATIGRAPHY AND PHYLOGENETIC RELATIONS Faunal changes in response to climatic fluctuations during the Late Tertiary have made some naticids useful as zonal fossils. As shown in Figure 2* Middle Miocene time is most easily characterized by its naticid assemblage* be cause four species lived at that time that did not exist before or survive afterward. In addition* there are an additional seven species whose stratigraphlc occurrences either begin or end In Middle Miocene rocks. This time segment can thus be defined by a combination of range zones and c oncurrent - range zones. The Late Miocene Is also characterized by the presence of two species peculiar to it* and by the overlapping stratigraphlc oc currences of two other species. Similarly* the Early Pliocene contained one species peculiar to it* the first occurrences of two species and the final occurrence of another one. The first appearance of one species and the last appearance of another species overlap in Middle Pliocene strata. Some intervals of Late Tertiary time can thus be recognized* with varying degrees of con fidence* by the naticids that occur in them. The Middle Miocene is moat easily recognized because of the relative- 35 FIGURE 2 Stratlgraphlc occurrence of eastern Pacific Natlcldae. All Pleistocene species are extant. Data based on museum specimens are shown as solid lines, probable occurrences as dashed lines, and less reliable data as dotted lines. 36 SPECIES PLE I S T OCE NE UPPER | LOWER PLIOCENE UPPER MIDDLE LOWER MIOCENE UPPER | MIDDLE | LOWER Notico (Natico) clarki Natica (Notica) vokesi Natico (Naticarios) chemnitzii Natica (Ncticarius) grayi Natica (Naticarius) posuncula Notica (Naticarius) teglandi Natica (Stigmoulax) broderipiana Notico (Cryptonatica) clausa Natica (Cryptonatica) oregonensis Natica (Tectonatica) janthostoma Notica (Tectonatica) satsopensis Bulbus frog it is Eunoticina insculpta Potinices (Polinices) bifascialus Polinices (Polinices) intemeratw Polinices (Polinices) uber Polinices (Euspira] diabloensis Polinices (Euspira) draconis S P E C I E S P L E I S T O C E N E UPPER | LOWER P L I O C E N E M I O C E N E UPPER | MIDDLE | LOWER UPPER | MIDDLE LOWER Polinices (Euspira) g a U a n o i Polinices (Euspira) lewisii Polinices (Euspira) pallidus Polinices (Euspira) victorianus Polinices (Mammilla) caproe Polinices (Hypterita) helicoides Neverita (Neverita) kirkensis Neverito (Glossaulax) andersoni Neverita (Glossaulax) j a m e s a e Neverita (Glossaulax) reclusiana S i n u m c y m b a S i n u m perrini S i n u m s c c p u l o s u m 39 ly large number of species chat are found In It, includ ing common ones such as Natica (Natlcarius) posuncula Hanna & Hertleln. The Late Miocene is less easily recog nized in terms of practical field work, because the species restricted to it, Polinices (Euspira) diabloensis (Clark) and Neverita (Neverita) kirkensls (Clark) do not occur commonly. Lower Pliocene strata are most confident ly recognized in Oregon and Washington because the species with concurrent-range zones are most restricted to that region. The overlapping species, Natica (Cryptonatica) Qgg&qnanglg and a. (Tectonatica) lanthoatoma. also are fairly abundant In rocks there. Conversely, for Middle Pliocene beds, only one of the species that defines a zone, Polinices (Euspira) aaHanoi Dali, is rare. For strata deposited at other times, such as during the Early Miocene, or the Late Pliocene through Late Pleistocene, naticlds are less useful as zonal fossils, but may still be helpful in making more general correlations. All of the species In Figure 2 that existed during the Pleisto cene are still extant. The changes in abundance of Neogene natlcid species shown here agree well with the Late Tertiary climatic trends and attendant faunal changes described by Addlcott (1969) for the marginal northeastern Pacific. Addlcott shows a steadily warming climate during the Early Miocene, peaking In the Middle Miocene, then followed by a sharp cool ins trend through the Late Miocene and Pliocene. The Middle Miocene thermal maximum is reflected by the natl- cldsi with an Increase In number of species from six to 14 between the Early and Middle Miocene (Figure 2), and by an Influx of warm-water subgenera into higher latitudes. Of the species first appearing In the Middle Miocene of western North America, four belong to genera (sinum1 ) or subgenera (Natica, Natlcarius) that are characteristic of warm hydroclimates. The warm-water Miocene species Neverita (Glossaulax) andersonl (Clark), which occurred only in southern and central California during the Early Miocene, is found in Middle Miocene deposits in northern Oregont although rarely, as further evidence for a warm ing trend at that time. At the close of the Middle Miocene nearly all of these warm-water taxa became extinct in western North America. Natica (Tectonatica) satsopenais Addlcott in the Lower Pliocene deposits of Washington is the only species Introduced after the Middle Miocene that belongs to a warm-water generic taxon, and this species is associated with a cool-water fauna. Except for J J . (J.) satsopenais. the other three species first appearing in the Pliocene belong to the largely extra-tropical subgenera EUBPlrf And Cryptonatica. By the appearance of several species In the Middle Miocene only, it is clear that optimum conditions for the warm-water natlcids existed only briefly in this region, 41 as indicated by the Tertiary climatic curves of Addicott (1969). The utility of many species as zonal fossils thus results from a warming peak that briefly reversed the general trend of climatic deterioration along the north eastern Pacific margin during the Cenozoic. Species intro duced after the Middle Miocene have tended to persist for longer periods, with most of them surviving to the present. The most likely phylogenetic relations of western North American Neogene naticlds are given in Figure 3. Appraisal of ancestral relationships is based on strati- graphic occurrences and shell morphology, with the aid of opercular features in the few cases where they are known. In this scheme, five lineages are proposed, with the realization that future work might demand a different ar rangement. The first lineage shown, of species related to Natica (Cryptonatica) clausa, has fairly clear-cut rela tionships. The only real uncertainty is the inclusion of £ J . (Tectonatica) satsopensls in this lineage. It is not clearly related to this group of northern species, but has even fewer similarities to other species. The lineage arising from Natica (Natlcarius) tec- landl Hanna & Hertleln was short-lived, but apparently produced a number of the distinctively warm-water species that so strongly characterize the Middle Miocene. The kinship of the Middle Miocene daughter species to U. (fl.) t.Klandl 1. not established beyond reaeanable doubt> and FIGURE 3 Phylogenetic relations of Neogene Natlcldae of western North America. 42 M I O C E N E EARLY | MIDDLE | LATE P L I O C E N E EARLY 1 MIDDLE I LATE NATICA OREGONENSIS /---------------- / f______NATICA CLAUSA NATICA PCSUNCULA L NATICA SATSOPENSIS v . NATICA VOKE3I NATICA CLARKI NEVERITA JAMESAC NEVERITA KIRKENS1S i NEVERITA ANDERSONI ■ \ \ * \ NEVERITA RECLUSIANA 1 V POLINICES RALIANOI \ \ \ V POLINICES LEWIS I I I \ V POLINICES DIABLOENSIS V POLINICES DRACONIS POLINICES VICTORIANUS > SINUH SCOPULOSUH * A \ V SINUH PERRINI 43 44 the relationship of (Natica) clarki Etherington to this lineage Is even less certain. The Pollniclnae lineage apparently arising from Neverita (Glossaulax) andersonl (Clark) is generally well established but does have some uncertainties. The rela tionships between (G.) andersonl. (G.) lamesae Moore, and £ J . (G.) recluslana (Deshayes) are well-founded, but the branching-off from this stock of Polinices (Eusolra) gaHanoi Dali is not. The association between (G.) andersonl and £. (£.) gaHanoi is made because they both have a prominently grooved umbilical callus, and be cause some of the many growth forms of (G.) andersonl and { £ . (G.) recluslana are quite similar to typical speci mens of £. (£.) gaHanoi. An alternate possibility would be a Paleogene ancestor for the £. (£.) gaHanoi group, with as yet undiscovered connecting specimens in the Early Miocene. This idea does seem unlikely because no similar or comparably large species are known in Oligocene de posits, but must await study of the Paleogene naticids for confirmation or rejection. The relationships of £. (£.) gallanol. £. (£.) lewisll (Gould) and £. (£.) draconls (Dali) are unequivocal, whereas the position of £. (£.) dlab1oens1s (Clark) in this group Is uncertain. The simple lineage involving £• (£.) victorlamia Clark & Arnold and £. (£.) oallldus (firoderlp & Sowerby) is uncertain because of a lack of interconnecting forms. 45 although the species occur together. The final proposed lineage, involving Sinum «enmiln«u» (Conrad) and £. Per rin i (Arnold), is obvious because these are the only two Neogene Slnums in this region. However, the extremely elongate shell of £• perrlnl is so radically different from that of £. scoouloaum that the species is possibly a migrant from another area, Just as Polinices (Mammilla) caprae (Philippi) in the Recent eastern Pacific fauna came from the Indo-Pacific region during the Pleistocene. The proposed phylogenetic scheme accepts origins for some subgenera, such as Eusolra and Natica. in more than one lineage. In fact, this is perhaps an expectable result of evolution in a family as morphologically con servative as the Naticldae. The limited number of shell characters observed in the family has resulted in the repetition with time of certain features thought to be of generic significance, such as a slender umbilical callus or radial wrinkles at the suture. Because Neogene naticfcb of this region are all of modem aspect, it has not been possible to trace the origins of given shell characters to more primitive groups such as the Ampullosplrinae, which were important in the Paleogene. Study of the Paleogene natlcld fauna of western North America has been neglected, so that it is not known from which of its stocks the Neogene species evolved. SHELL--OPERCULAR RELATIONS Two of the Recent species under consideration* Natica (Si-lgnaulax) broderlplana Recluz and ft. ( 5 .) elenae Recluz* show an apparent functional relationship between the sculpture of the operculum and the shape of the umbilical callus and umbilicus. Morphologic details are given fully in the discussion of the former species* but basically the massive central spiral ridge of the operculum fits exactly into the curved umbilical opening and also seats Itself in the notch excavated by the umbilical sulcus in the inner lip. This relationship is also seen in the Caribbean Miocene species f t . (£.) k u p - plana (Toula), and the Caribbean Recent species f t . caven- nensla Recluz* and probably occurs in species elsewhere as well. The advantage of this feature is not apparent. The function of the interlocking shell and operculum is probably to maintain a constant relative position between them* perhaps to have the operculum in the best position for rapid closure* or perhaps to provide a position for better leverage when the animal is extended from the shell and maneuvering in search of prey. This apparent function al relationship is one which could be easily understood by 46 47 observation of living individuals. None of the other eastern Faclflc fossil or Recent species shows such a relationship. Partially calcified opercula are known in at least two species. Clarke (1961*363) described the operculum of the abyssal south Atlantic species (Ker- auelenatlca) arises (von Martens* 1878) as "homy with a prominent * thin calcareous layer over the central part." Eunatlcina insculpta (Carpenter) in the present study has the same kind of operculum* of which 1 have seen two. The two species Involved are properly placed in their respec tive genera, Judging by shell characters. The type- specles of * - A* Is land lea (Gael in), is treated here * and I have seen no evidence of opercular calcifica tion among hundreds of specimens. The type species of Eunatlcina. £. papilla (Gmelin, 1791) of the Indo-Pacific region, is said to have a chitlnous operculum (Souverbie & Montrouzier* 1874i198, pi. 7, fig. 8t Oyama, 1969*79-80, fig. 6), but 1 have not seen a specimen of it. It would be quite interesting to determine how commonly such partly calcified opercula occur in these genera, and to learn whether this is a constant feature through the life of an individual. BORING Use of Che radula in boring the shells of prey species is ubiquitous in the Natlcidae* also occurs com monly in the Muricidae and Cymatlidae* and is known in several other gastropod taxa (Carriker ££ al« * 1963i Carrlker & . Yochelson* 1968t Sohl, 1969a), Sohl (1969a) notes 37 genera of living Natlcidae, 34 of Cymatlidae* and 183 of Muricacea (comprised mostly of Muricidae), so that the murlcaceans might have the largest number of bor ing species in modem seas. However* the naticlds are noted for their rapacious feeding habits and some species attain large size* so that they can be a major factor in the dynamics of a molluscan community. Of the gastropods with boring habits* the naticlds have the longest geologic history* The first undoubted natlcaceans appeared In the Early Jurassic (Carrlker & Yochelson* 1968i Sohl* 1969a)* although Cossmann (1925) thought they started in the Trlassic and was quoted by many later workers. The first murlcaceans occurred during the Early Cretaceous and the first cymatlids appeared in the Late Cretaceous (Sohl* 1969a). In upper Cenozoic strata of the Pacific Coast* all three groups occur commonly* but the naticlds are found in the greatest numbers. In some Neogene deposits 48 49 of this region, naticlds often make up an appreciable volume of the total mollusk specimens there. In such as semblages a high percentage of prey bivalves have been bored. Naticlds are also very commonly cannibalistic, as it Is common to find drilled naticld shells In Recent and fossil deposits. Naticlds characteristically drill a spherical parabolic hole in their prey, In contrast to the cylindri cal hole of cymatiids, and this variance is due to dif ferences In anatomy of the boring organs (Carrlker & Yochelson, 1968). Ihe actual boring by naticlds and cymatiids is a combination of rasping with the radula and chemical softening of the prey shell by a non-acidic fluid secreted by an accessory boring organ (Carrlker, ££ al., 1963). After clearing away surface debris from the prey shell with Its radula, an attacking snail presses its ac cessory boring organ to the victim and the secretions loosen surface crystals of calclte rather than entirely dissolving them (Carrlker, 1961). The radula is then applied to the weakened prey shell and the attacker rasps off and Ingests flakes of calclte. The radula apparently has little effect on a prey shell that has not first been chemically weakened (Carrlker, 1961). Much more detailed accounts of the anatomy, histology and function of naticld boring organs, and references to numerous other works, are given by Carrlker (1961), Carrlker & Yochelson (1968) and 50 Sohl <1969a). The distribution of raurlcld boreholes on prey bi valves is apparently random, whereas the borings of naticld are more localized, and this difference is related to the anatomy of the attacking snails (Carrlker & Yochel son, 1968). Muriclds have relatively small bodies and hold on to the outer surface of the prey when drilling, whereas the naticlds have much more voluminous bodies and entirely envelop the prey. Thus wrapped around the prey shell, the naticid is restricted in its choice of drill site by the way in which Its body fits around the victim. Reyment (1966) notes that Recent naticid borings are usually localized on a small portion of the prey valves, and Sohl (1969a) describes the same relationship for Late Miocene species. Limitations in the way the naticid grasps the prey might explain the preferential boring of one prey valve (Fretter & Graham, 1962), and Sohl (1969a) notes that "valve-preference for borings is not so pro nounced in equivalved pelecypods.” The influence of naticld predation on other mol- lusks and on community dynamics Is an obvious field for future work by biologists. For example, the deep In fauna 1 position of thyaslrld bivalves may be a response to attack by naticlds and other borers (F. R. Bernard, per sonal communication, 1973). Naticlds are obviously a major environmental stress on certain bivalve species* but their influence as an aspect of selective pressures on bivalves has yet to be investigated. SEXUAL DIMORPHISM Sexual dimorphism Is well-known among many groups of gastropods (Sohl, 1969b), and when It occurs among prosobranchs the adult females tend to have larger and broader shells than the males (Cox, I960)* Sohl (1969b) noted this trend for two American Late Cretaceous natl- clds, and also mentioned a similar relationship for the Recent western Atlantic species Neverita (Neverita) duplicate (Say)* Bernard (1968) showed that sexual di morphism In Polinices (Euspira) lewis11 (Gould) Is ex pressed as a difference in total weight to shell-welght ratios, with a mean ratio for males of 0*472 and for females of 0*401* However, he did not correlate sex dif ferences with shell form in his sample of 1875 specimens. As explained more fully in the discussion of this species, £. (£.) lewis11 does have two distinct morphologies in fossil and Recent populations, and these are probably related to sex differences. A simple examination of a large living population would determine this. Some past researchers have made brief reference to male and female forms of Neverita (Glossaulax) recluslana (Deshayes). If they exist, these forms were not noted during the present study* Given the broad range of 52 53 morphologies shown by Recent and fossil specimens of this species* It would probably be very difficult to detect sexual dimorphism on the basis of statistical analysis alone. An Investigation of Recent specimens* if it could relate sexual differences to some of the morphologies in this complex of forms* might be very useful in understand ing reasons for the great variation in this species. Addlcott (1970) first described two forms of the Miocene species Polinices (Euspira) vlctorlanus Clark & Arnold* which were also observed in this study. These forms were not ascribed by Addlcott to sexual dimorphism* although they could be. As Sohl (1969bi97) cautions* it is difficult to determine sexual dimorphism from shell characters alone* and such determinations are risky unless correlated with soft-part anatomy. SYSTEMATIC ACCOUNT Family Natlcidae Gray, 1834 Diagnosisi Shell globose, ovate-conic or auriform, rarely elongate) spire low to moderately elevated) whorls commonly convex, usually smooth) some species with radial grooves at suture or (mainly Slnlnae) with spiral costellae) umbilicus open or closed, commonly with a funiclej aperture moderately large, usually entire, oval to semilunari outer lip usually sharp, oblique, columel- lar lip usually with a callus, especially posteriorly) operculum chitinous or calcareous, rarely partly calcified, paucispiral and with eccentric nucleus, may be rudimentary. Discussioni Differences in shell features may Indicate evolutionary trends in the Natlcidae. Of the four sub families treated here, the Ampullosplrlnae occurs earliest in the geologic record and was a major group in the Mesozoic and Paleogene, although it contains only a few living species. If this group Is considered to be primi tive, then narrow or closed umbilici, channeled sutures, and chitinous opercula presumably are primitive features. Assuming that the most diverse naticld subfamilies are the most advanced ones would place the eastern Pacific Ampullosplrlnae as the most primitive subfamily and the 54 55 Natlclnae as Che most advanced subfamily, with the Polinicinae and Sininae in intermediate positions. Using this assumption, the Polinicinae would be more advanced than the Ampul1ospirinae because they mostly have dis tinctly open umbilici and non-channeled sutures, although they all have chitinous opercula. In the Polinicinae, the formation of distinct umbilical calli and weak funlcles perhaps presages the elaboration of these features in the Natlclnae. The Sininae would be the next most advanced group, because although they have features similar to those of the Polinicinae, their extremely depressed shells are more highly modified than those of the Polinicinae. Natlclnae would presumably be the most advanced of the eastern Pacific naticid subfamilies. In this group, the umbilical features are most elaborate, shell sculpture and bold color patterns are common, and calcareous opercula are ubiquitous. Species rarely occur in the Ampullospirl- nae and Sininae with partly calcified opercula, but all of the Natlclnae have such opercula, many of which are elaborately sculptured. If the development of shell sculpture and calcified opercula, and the elaboration of umbilical morphology are advanced features, then the Natlclnae are clearly the most advanced living naticlds. Such speculation can only be tested by more extensive study of Mesozoic and Paleogene naticld faunas, to deter mine the order of appearance of specific shell features in 56 Che geologic record. The Natlcidae range from Jurassic to Recent* with about 300 living species worldwide. A key to the sub families used here followsi la. Outer surface of operculum entirely calcareousi shells mostly smooth and umblllcate, with a funicle separated from parietal callus by a sulcusi imperforate shells with a semicircular callus . ........... Natlclnae lb. Outer surface of operculum chiti nous or with central portion thinly calcareousi shells smooth or with spiral costellaei mostly umbllicate, lacking distinct funlcle ....................... 2 2a. Sculpture of distinct spiral costellae ...................... Sininae 2b. Shells lacking distinct spiral sculpture...................... 3 3a. Suture often channeledi umbilicus closed to very narrowly openi umbilical callus thini umbilical area commonly bounded by low basal ridgei operculum chitinous to partly calcareous .............. Ampullosplrlnae 57 3b. Suture not channeled In most general umbilicus usually dis tinctly open, may be closedi umbilical callus usually thicken ed posteriorly, may be massivei operculum chitinous ........... Polinicinae Subfamily Ampullospirinae Cox, 1930 Ampullosplrinae Cox, 1930i305. Diagnosisi Shells globose to ovate, thin- to thick- shelledi spire usually low, may be elevatedi suture often channeled) whorls smooth or with weak spiral costellae, especially near suturej umbilicus narrowly open or closedi umbilical callus thini umbilical area sometimes bound by low basal ridge% operculum entirely chitinous or partly calcareous in living species. DiscussIoni This subfamily is represented by a single genus and species in the eastern Pacific, which is con fined to Arctic waters. The Ampullosplrinae are poorly known in Recent faunas worldwide, but were a major group in Paleogene seas, comprising most of the Natlcidae at that time. Genus Anauropsis Morch, 1857 58 Amanr^pffMorch in Rink, 1857i81. Type-specleai Katies hellcoldea Johnston, 1835 (- Nerlta lslandlca Gmelln, 1788), by subsequent designation (Dali, 1909bt89). Recent, clrcumboreal. Figured herein. PisanosIsi Shells small to medium in size, elongate, thlm whorls laterally compressed, sculpture with minute spiral costellaei suture deeply channeled. Umbilicus extremely narrow and slit-like. Inner lip slightly thickened, simple, lacking distinct umbilical callus. Parietal callus very thin. Operculum chitinous, entirely filling aperture, Radula typically naticid, with trlcuspate rachidian, one multlcuspate lateral, one blcuspate inner marginal, and one monocuspate outer marginal tooth per half-row. Discussioni AfflftVrWftlff characterized by its deeply channeled suture, slit-like umbilicus, and thin shell. In Recent seas, the genus occurs only in northern and southern polar and sub-polar regions, although in earlier geologic ages it extended into temperate climates. 59 Aaauropsls UHndl?? (Gmelln, 1788) Plate 1, figure 1 Nerlta islandlea Gmelln, I788i3675. Mamma^(^»guyon«t«) Islandlea (Gmelln)i Morch In Rink, Natica lalandlca (Gmelln)i Jeffreys, 1867i214t Jeffreys, 1865T2157pT. 78, fig. 1. Aiy*yrftpftifl lalandlca (Gmelln)i Sara, 1878t156, pi. 21, fig, 17, pi. V, fig. 10, pi. XVIII, fig. 10| Frlele & Grieg, 1901i111, 68 j Odhner, 1913*9, 44-46, pi. 4, figs. 29-35i Bousfleld, 1960(16, pi, 2, fig. 20* Ha be & Ito, 1965i31, pi. 8, fig. 7i MacPherson, 1971(54- 55, pi. 3, fig. 11. Bulbus (Amaulopsls) lslandlcus (Gmelln)i Wenz, 194111035, fig. 29o5f Kotaka, 1962*134, pi. 33, fig. 16. Natica hellcoldes Johnston, I835i69| Lov&i, 1847 d49» Mlddendorff, 1849i416-419, pi. 7, figs. 8, 9* Forbes & Hanley, 1851, vol. 3i339-341, vol. 4i pi. 100, fig. 6| not Philippi, 1851i93-95, pi. 13, figs. 13, 14 oj not raiiLppi, idoliyj-yo, pi. u, rigs, u, f ?-Bulbus~ l i Reeve, 1855ipi. 30, fig. 142i Danielssen, 1861t31* Sowerby, 1883t91, pi. 2, fig. 12* Tryon, 1886(53, pi. 22, fig. 31i Gould, 1870(348-349, fig. 617. Ain«yf^pa1 f hellcoldea (Johnston)t Dali, 1885(525 [as A* hellcoldea "Mlddendorff-J. Natica suturalIs Gray, 1839i136, pi. 37, fig. 4i Philippi, 1851(11(1, pi. 15, fig. 17. Natica canalIculata Gould, 1839 d97| Gould, 1841(235-236, flg.161 • not Natica canalIculata Deshayes, 1832, vol. 2i170-171, Atlas, pi. 21, figs. 9, 10 fossil, France . Natica Rouldll C. B. Adams, 1847(21 [new name for canaliculate Gould, 1839, not Deshayes, 1832]t Clench & Turner, 1950(288, pi. 41, fig. 12. ? Natica gfiuldU Philippi, 1845(77-78, 60 Natlea cornea Moller, 1842ai80i Moller, I842b|7. Natica cxulans Loven MS--Gould, 1841i235i Jeffreys* 1867i 216 fas synonym of 1$ . lalandlca^. Amauropsis purpurea Dali* 187lil24, pi. 15* fig. 16j Dali* 1574i251i Dali, 1885i525i Dell, 1902i551, pi. 38, fig. 9i Dali, 1921i166i Oldroyd, 1927i736* Keen* 1937*29i Burch, i946i33i MacPherson, 1971i55, pi. 3, fig. 10, PgflfiElPtlWH Color.--Shell exterior and callus whitei interior white, sometimes with pale violet stain. Perlostracum thin, medium yellowish white to brownish grey, pale olive brown or rust brown, the colors often axially streakedj earlier whorls often darker. Size.— Average* height 28 mm, diameter 22 mmi largest speclment height 48.7 mm, diameter 41.9 mm [USNM 111301, Cape Menshikof, AlaskaJ. Shell Form.--Shell elongate* spire very elevateds body whorl moderately inflatedi whorls laterally compress ed i shoulder tabulate, roundedi shell very thlnf whorls about 5 apex always eroded i suture deeply channeled. Spiral sculpture of minute* low, irregularly spaced costellae and microscopic, wavy, closely spaced llnea- tionsi axial sculpture of Incremental growth lines. Parietal callus very thin* not thickening into posterior aperture1 angles lacking anterior lobe. Umbilicus open, extremely narrow and siit-like* usually concealed by perlostracum of inner lip margin. Inner lip slightly 61 thickened, simple, lacking umbilical callus. Basal lip thickened. Operculum.-- Chit inous. filling aperture. Specimens Examinedi 362. Geographic Occurrence and Ecologyi Clrcumboreal, In the eastern Pacific, south to Cape Menshikof, Bristol Bay, Alaska (57°31t N), apparently unknown in the Aleutians) in the western Pacific, south to the Sea of Okhotsk) in the western Atlantic, south to Chesapeake Bay) in the eastern Atlantic, south to northern England and Ireland. Known on soft bottoms in depths between 9 and 1267 meters, but primarily in 30 to 60 meters depth) apparently occurs in shallower depths northward, but depth data are poor for verified live specimens. Stratiaraphic Occurrencei Reported by Frlele & Grieg (1901) "as a fossil in Scandinavia, the British Isles, Canada, Siberia, northern Russia and Spltzbergen,N but exact ages of the fossil deposits were not given. Type Localitiesi Nerita islandlca - Unknown (Gmelin, 1788). Natlea hellcoldes - Berwick Bay, Scotland (Johnston, 1835). Natlca suturalis - "North Sea" (Gray, 1839). canaliculate - Unknown (Gould, 1839). 62 Natlea aouldil - Maine, U.S.A. (Philippi, 1845). Natlea cornea - Unknown (Mtiller, 1842). Amaurooaia purpurea - Saint Michael Island, Norton Sound, Alaska (Dali, 1871). Type Materiali Nerlta islandlca - Unknown, presumably in Linnaean Society of London (Dance, 1966). Natlea hellcoides - Unknown (Johnston, 1835). Natlea suturalls - Unknown, presumably in BM(NH) (Dance, 1966). Natlea canaliculata - Holotype, MCZ 151080 Qlnad- vertently designated lectotype by Clench A Turner, 19503. Natica aouldll - Unknown, presumably in BM(NH) or Humboldt Museum, East Berlin, German Federal Republic (Dance, 1966). Natica cornea - Unknown, presumably in BM(NH) or Zoo logical Museum, Copenhagen, Denmark (Dance, 1966). Ananrnna1m purpurea - Holotype, USNM 108988. NniMnfllatural f!wi— nrarvi The seven named taxa included here under A, islandlca have been considered synonymous in various combinations by earlier workers. Only purpurea has been consistently regarded as a separate species in recent years, and it is supposed to be darker in color, less elongate, and to have a thicker shell than £. Island- 63 lea. Pacific specimens have been assigned to £. purpurea and Atlantic specimens to Islandlca. Shell color and thickness vary with specimen size, younger shells having lighter color and a thinner shell than older specimens. Pacific and Atlantic populations show the same range of color and shell thickness. Figure 4 shows that the ranges of shell size and height/diameter ratios are the same for Pacific and Atlantic individuals. For the speci mens measuredi the height/diameter ratio for Pacific specimens ranges from 1.15 to 1.38, and the same ratio for Atlantic specimens ranges from 1.17 to 1.14. Amanron«i« purpurea is thus not separable from islandlca as indi viduals or populations, on the basis of shell dimensions, color or shell thickness. Discussioni There is no apparent size gradation with latitude in this species, as there is in Natica (Crvp- tanatlea) clausa Broderip & Sowerby, although islandlca is not distributed throughout an equally broad range of latitude. The largest known individual of £. islandlca was found at Cape Menshlkof, Alaska, which is also the southern range end-point for the species* Odhner (1913) reported that the umbilici of Atlantic specimens may be closed, but I have not seen this. The perlostracum is usually thickened within the umbilicus, which may conceal the slit-like umbilical opening. FIGURE 4 Comparison of height to diameter ratios of EgyttU* raslualw (Deshayes) and Neverlta alta (Arnold). Based on Recent LACM specimens. 64 HEIGHT I N MM 65 50- Amauroptit Ulondica + Atlantic spacimans • Pacific ipccimtni 40- 4 4 30- 4 ■ 4 ♦ * % ♦ 20H %. V . ■ * : 4 / . 10H ** + + 4+ 4 1------------------------ 1 ------------------------ r 10 20 30 40 DIAMETER IN MM 66 Although the color range of Pacific and Atlantic specimens Is Identical, more southerly specimens from each ocean tend to be lighter in color. Subfamily Pollnicinae Finlay & Marwlck, 1937 Polinicinae Finlay A Marwlck, 1937*47. Diagnosisi Usually globose, sometimes very largei spire low to moderately elevatedt whorls smooth and usually umblllcate, with funlcle rudimentary or absenti parietal callus sometimes very thickt operculum chltinous, thin. Discussioni This Is the largest and most diverse among the subfamilies of eastern Pacific Natlcldae. A key to the genera and subgenera used here followst la. Shell very thin, umbilicus open, dis tinct umbilical callus lacking • 2 lb. Shell moderately thick, umbilicus open or closed, with a distinct umbilical callus ... ....... 3 2a. Base distinctly flattened, final whorl greatly inflated ....... 2b. Base not flattened, umbilicus Callnatlcina slit-like 67 2c. Base not flattened* umbilicus dis tinctly open* not slit-like .... Chorlstes 3a. Umbilicus open.............(Pollnlces. s.l.) 4 3b. Umbilicus closed* except rarely....................(Neverlta* s.l.) 6 4a. Umbilical callus large to massive* shell ovate ....... PolinIces. s.s. 4b, Umbilical callus not large or massive* usually slender* shell globose to very depressed.............5 5a. Shell elongate pyriform* thin* umbilical callus slender ............. Mammiila 5b. Shell globose* umbilical callus slender to thickened (especial ly posteriorly) .................... Eusolra 5c, Shell strongly depressed! um bilicus broadly open* with umbilical callus as a thin lobe suspended anteriorly from pillar-like funlcle ............. Hvoterlta 6a. Umbilical callus without a transverse groove ........... Meverlta. £•&• 6b. Umbilical callus with a transverse groove ........... Glosaaulax 68 Genus Pollnicea Montfort, 1810 Pollnlces Montfort, 1810t223-224, pi. 56. Type-speciesi Pollnlces albus Montfort, 1810, by original designation. Recent, ?West Indies. Diagnosisi Shells small to very large, globose to ovatei whorls inflated to subdued, smooth except for incremental growth lines and microscopic spiral costellaei suture slightly to deeply impressed. Umbilicus narrowly to broadly open, funlcle lacking to very weak. Umbilical callus slender to broad and massivet parietal callus thin to thick. Operculum chltinous, entirely filling aperture. Radula typically naticld, with trlcuspate rachidlan, one multicuspate lateral, one blcuspate inner marginal, and one monocuspate outer marginal tooth per half-row. Discussioni Pollnlces is characterized by Its globose form, chltinous operculum, and open umbilicus. In the original indication of Pollnlces. Montfort (1810) described the genus as having a solid operculum ("opercule solide”) like that of "natlce canrene" (Natica canrena Linnaeus, 1758), a well-known Recent Caribbean species. This has apparently been overlooked by other workers, and is probably a result of Montfort never having seen an operculum of £. albus and assuming it to be the same as those of other natlcids he was familiar with. All 69 subsequent workers have considered species of Pollnlces to have chltinous opercula. The status of the type-species Is unsettled. In the Indication for Pollnlces. Montfort (1810) mentions Nerlta mamllla Linnaeus* 1758* but It Is not clear whether he considered £. albus and £• mamllla to be synonymous or only meant mamllla as another example of Pollnlces. The Identity of £. albus Is also In question* and numerous authors have equated It with £f . mamllla. Woodring (1957i 89) places £. albus together with Natica mamlllarls (Lamarck* 1767) In the synonymy of Natica brunnea Link* 1807, a Recent Caribbean species. However, Cemohorsky (1971i191) believes £. brunnea to be a synonym of Albula heoatlea Rodlng* 1798* and in any case to be a tan or orange-brown species* whereas £. albus was described (and named) as white. Cemohorsky further believes £• albus to be most probably synonymous with Nerlta mamllla. the second species mentioned by Montfort (1810), although mamllla Itself is a dubious taxon that may be an earlier name for £. lacteus (Guildlng, 1834) of the West Indies, or for £. tumldus (Swainson* 1840) or £. pvrlformla (Re- cluz, 1855) or the Indo-Paclflc. Linnaeus' (1758) type locality for g. fAlU1 V1 of "Barbados" places the species In the Caribbean, but still does not Identify it. 70 Subgenus Pollnlces Montfort, 1810, sensu strlcto Diagnosist Shell small to medium In size, commonly ovate but may be globosei whorls moderately inflated to subduedi suture slightly Impressedi shell thickness average or greater* Umbilical callus nearly always broad and mas sive, rarely slender, and often nearly closing umbilicus. Parietal callus moderate to heavy. Discussioni This is comprised of a large number of species, essentially Including most of the tropical Pollniclnae worldwide. Pollnlces (Pollnlces) uber (Valenciennes, 1832) Plate 1, figures 2, 3 Natica uber Valenciennes, 1832i266i Orbigny, 1840ivol. 5, p. 401, vol. 9, pi. 55, figs. 12-14 (plate 1835)t M. E. Gray, 1850ivol. 4, p. 82t Troschel, 1852i157i Philippi, 1852(60-61, pi. 10, fig. 1 (plate 1850) fas uber "Humboldt"3i Reeve, 1855ipi. 13, figs. 54a,b* Carpenter, 1857i292i Sowerby, 1883t87, pi. 4, fig. 51t Tryon, 1886i48, pi. 17, fig, 61 Cnot fig. 66, ?« £. panamaenaia (Reclus, 1844)]» Hanna, 1926(451-452. Polinlces uber (Valenciennes), Carpenter, 1857i452-453i Steams, 1894(195-196 C«® "Polynices"}1 Baker, 1902( 41t Dali, 1909(235i Jordan, 1924(156j Valentine & Meade, 1961(25, 28i Jordan, 1936(114i Hertleln & Strong, 1955ti40i Parker, 1964(153, pi. 5, fig. 5i Stanton, 1966(23. Polinlces uber uber M. Smith, 1944i12, fig. 123 nomen nudumifas P. uber uber "Valenciennes"J. Polinlces (Polinlces) uber (Valenciennes), Grant & Gale, 1931(799, text-rig7l2i Keen. 1958(323, fig. 272 Cin part, also £. oanamaensls (Recluz, 1844)3i Keen, 71 1971.480, fig. 882. Watlea vlrftlnea R&luz, 1850.388-389, pi. 12, fig. 6. Watlea ovuib Msnke, 1851.165-166) Carpenter, 1857.237, "Natica uberlna Orbigny", Recluz, 1844.210 fas of Valen ciennes i not W, uberlna Orbigny, 1842, ? • » £. lacteus (Gullding, 1854), Recent Caribbean}) Philippi, 1853. 142-143, pi. 19, fig. 16 (plate 1851) fnot N. uberlna Orbigny, 1842, 7 - £. lacteus (Gullding, 1834), Re cent, Caribbean}. Description. Color.--Shell exterior white, with pale buff to yellow band below suturej first 1-2 nuclear whorls tinted pale browni Interior white. Perlostracum thin, usually pale yellowish white, but ranges to medium yellowish brown, and may have Irregular rust brown stains. Size.--Average. height 20 mm, diameter 16 rami largest specimen, height 35.7 mm, diameter 31.4 ram £lACM 70-15, Venado Island, Panama}. Shell Form.— Shell globose to elongate, spire moderately to greatly elevated) body whorl not greatly Inflated) whorls range from slightly tabulate In globose forms to slightly concave in elongate formsi shell thick ness average) nuclear whorls 3%, with closely set spiral costellae separated by narrower Interspaces) postnuclear whorls 3%) suture slightly Impressed. Spiral sculpture of microscopic, closely spaced, very weak costellaei axial sculpture of Incremental growth lines. Parietal callus very heavy, thickly filling posterior apertural 72 anglei anterior lobe weak. Umbilicus narrowly open, often slit-like. Umbilical callus narrowed nearly to a point anteriorly, evenly expands to posterior end of umbilicus where it merges smoothly with parietal callusf shallow groove traverses callus at Juncture of umbilical and parietal callii anterior inner lip thickened. Operculum.— Chltinous, filling aperture. Specimens Examined! 2000. Geographic Occurrence and Ecology* Cedros Island, western Baja California, Mexico (28°10'N), throughout the Gulf of California, and south to the Galapagos Islands and Paita, Peru (5°S). Found alive on sand bottoms, intertldally and in depths to at least 100 meters. Stratiaraphlc Occurrencei Ranges from Upper Miocene (?) to Recent. Upper Miocenei Castaic Formation, near Castalc, Los Angeles County, California (Stanton, 1966). Pliocene (?)t Imperial Formation, Imperial County, Cali fornia (Hanna, 1926). Pleistocenei Punta San Telmo (UCB), Islas Coronados (UCB), Magdalena Bay (Jordan, 1926), and El Molege (CAS), Baja California, and Isla Marla Madre (CAS), Mexico. Type Localitiesi Natica uber - "Habitat ad portum Cumanensem" (Valen ciennes, 1832). Hertleln & Strong (1955) thought this to be in Venezuela probably Cumana, 10°28*N , and to be an error. Accepting this species as an eastern Pacific one, the type locality is unknown. Natica vtrainee - Realejos (R&luz, 1850) CProbably in Nicaragua at 13°N]. Natica ovum - Mazatlan, Mexico (Menke, 1851). Type Materiali Natica uber - Unknown, may be in Museum d'Hlstoire Naturelle, Paris (Dance, 1966). Natica vlrainea - Unknown, presumably in Museum d*Hlstoire Naturelle, Geneva, Museum d*Hlstoire Naturelle, Paris, or BM(NH) (Dance, 1966). Natica ovum - Unknown, sold to a dealer (Zilch, 1958i53). Discussioni The £. (£.) uber species-group is one of the more difficult to deal with taxonomlcally among the tropi cal eastern Pacific natlclds, and includes the species £. (£•) UbSEt £• (£•) intemeratus (Philippi, 1853), £. (£.) panamaensls (Recluz, 1844), and £. (£.) otls (Broderlp & Sowerby, 1829), in addition to cool-water Chilean and Peruvian species not treated here. These species exhibit overlapping variations In form and umbilical morphology, although they can be identified with confidence in most cases. These four species range essentially throughout the 74 entire tropical eastern Pacific (Panamic zoological province), but some of them reach their greatest develop ment in different parts of the tropics. The largest speci mens of £. (£.) uber (height above 20 mm) generally occur in the Gulf of California and southward, the largest speci mens of £. (£.) lntemeratus (height above 20 mm) are found between Panama and Peru, and the largest individuals of £. (£.) (height above 45 mm) also occur south of Panama. Large specimens of £. (£.) otls are not localized in this way. The two most closely related species, £. (£.) uber and £. (£.) nanama<maiw. have separate areas of she11- size maxima, which reinforces the separation of these species made on shell characters. When typically developed, the shell of £. (£.) uber is elongate and high-spired, with an evenly tapered umbilical callus that reduces the umbilicus nearly to a slit. Large populations are often found at one time and consist mostly of smaller, elongate Individuals, the large, globose specimens being comparatively rare. The largest specimens £* (£•) uber are globose rather than slender. Indivi duals with umbilical calll more slender than usual for most of their anterior lengths are difficult to separate from specimens of £. (£•) panamaansla. Individuals of £. (£.) uber and £. (£.) " ar« commonly found to gether, and there is no difference in habitat, based on museum label data, to account for observed differences in 75 callus morphology. Large specimens of £. (£.) uber are not known along the outer coast of Baja California, al though the species ranges north to Cedros Island, so there might be environmental controls on shell size, and possibly also callus form. The most characteristic feature of £. (£.) panamaensis is its umbilical callus, which is slender for most of its length then abruptly broadens at its posterior end to match the width of the parietal callus. If the slender portion of this callus is thicker than usual, with a swol len appearance, then the specimen may be mistaken for a £• (£•) uber. Such errors are especially likely when specimens are small and have an elevated spire, which is more characteristic of £. (£.) uber but occurs in £. (£.) paf l aif l aenylp. Individuals of £. (£.) uber have ratios of body-whorl height to spire height of about l.Sil to 2.Oil, whereas individuals of £. (£.) panamaensis have ratios of about 2.5tl to 9.0tl, so there is a difference in relative elevation of the spires, although the difference may be minor in individual cases. Some past workers have stressed that £. (£.) panamaen sis has a strongly shouldered shell with a flattened peri phery, giving a distinctly squared appearance* However, this is not a common condition and seems to be best developed in cool-water populations of the Peruvian zoo logical province, which are not treated here. Such strong- 76 ly shouldered specimens from the Peruvian province may belong to other species* and a number of names have been used there for shells similar to £» (£. ) 1 have seen specimens from as far south as Isla Chiloe, Chile (42o30'S), that are very much like tropical indivi duals of £. (£.) nqpAmflPinal «. Peruvian and Chilean natl- cids related to the £. (£.) uber species-group need fur ther study to define their relationships with the tropical species. Polinlces (Polinlces) lntemeratus has the most con sistent morphology of any member of the £. uber species- group. The shells are consistently globose* with ratios of body-whorl height to spire height of 3il to 4*1, and the arcuate umbilical opening and distinct rlblet within the umbilical channel are characteristic features. The most common variation is for the parietal callus to be more massive than usual* which restricts the umbilical channel to a smaller arc of curvature. This is accompanied by a thickening of the anterior inner lip and a more severe narrowing of the posterior umbilical callus. In some specimens* the median expansion of the umbilical callus is reduced* and the callus assumes the straighter form seen on some £. (£.) uber specimens. However* the rlblet in the umbilical channel of £. (£.) is a distinguish ing feature in these Instances. Specimens of £. (£.) lntemeratus are never as elongate as those of £. (£.) uber 77 or as large as those of £. (£.) panamaensis. Pollnlces (Pollnlces) otls Is also relatively easy to identify because of its consistently elongate shape. In flated anterior portion of the body whorl, and commonly occurring brown color in the umbilical area* It is closest to £. (£.) lntemeratus In form, but its more elongate shell with a higher spire, broader umbilicus lacking a sharp rlblet, and greater size are distinctive. The northern range-limit for this species-group is well established at Cedros Island, Baja, California. Museum specimens labeled as £. (£.) uber from southern California have proven to be Polinlces (Euspira) pallldus (Broderlp & Sowerby, 1829). The southern range-limits of these species, mostly in northern Peru, are less certain* A relatively small number of cool-water Peruvian and Chilean specimens is available for study, and these large ly resemble £. (£.) panamaensis. although they sometimes differ in details of umbilical morphology, shell form and shell thickness. The earliest geologic occurrence of £. (£.) uber needs confirmation, given the difficulty of identifying members of this species-group. It is possible that £. (£.) uber or related forms existed in western North America during the Miocene, because there are similar and perhaps related species described from the Neogene deposits of Ecuador and Peru (Hanna & Israelsky, 19251 Olsson, 1932, 78 1964). Although there are distinct differences In protoconch sculpture between some members of this species-group* these can be seen in very few specimens. Even on living specimens* the protoconch is almost always worn and its delicate sculpture missing. Pollnlces (Polinlces) panamaensis (Recluz, 1844) Plate 1* figures 4, 5 Natica f Recluz* 18441208-209» Reeve* 18551pi. 5, figs. lla*b. Natica panamensis Csic} Recluz* Philippi* 1852(45-46* pi. 7* fig. J (plate 1850)t Sowerby, 1883i86, pi. 2* fig. 18i Tryon* 1886i48* pi. 17* fig. 60. Pollnlces panamensis Calc} (Recluz), M. Smith, 1944*12. Polig^ges^(golinice|)^panamaensis (Recluz)* Keen, 1971* J2ea£riBLlafli Color.--Shell exterior and Interior white, with pale buff to yellow band below suture. First 1-2 protoconch whorls medium brown. Periostracum thin* usually pale yellowish white, but ranges to medium yellowish brown* may have Irregular brown stains. Size.--Average* height 40 mm, diameter 33 mm* larg est specimen* height 62.8 mm* diameter 50.0 am C(LACM A.375* Islas de las Tr^s Marlas* Mexicoj. Shall Form.— Shell globose to elongate* spire 79 moderately to strongly elevatedi body whorl may be in flated in larger specimens} shoulder slightly flattenedf may be slightly concavet shell thickness average} nuclear whorls 3%, apparently smooth} postnuclear whorls 4} suture slightly to moderately impressed. Spiral sculpture of microscopic, closely spaced, very weak costellaei axial sculpture of incremental growth lines. Parietal callus very heavy, thickly filling posterior apertural anglei anterior lobe weak. Umbilicus narrowly to moderately open, rarely slit-like. Umbilical callus comes to a point anteriorly, narrowly expands toward posterior end of umbilicus, where it abruptly broadens to span the width of the umbilicus and merge with the parietal callus} shallow groove traversing callus at Juncture of umbilical and parietal calli Is usually absent} anterior inner lip thickened. Operculum.--Chltinous, filling aperture. Specimens Examinedi 750. Geographic Occurrence and Ecologyi Cedros Island, western Baja California, Mexico (28°10*N), throughout the Gulf of / _ California, and south to Bahia Independencia, Peru (14 14*S). Not known in the Galapagos Islands, but it may range farther south in Peru and Chile. Found living on sand bottoms intertidally and to depths of 139 meters. 80 Tvt>e Localityi Panama (Recluz, 1844). Type Material! BM(NH) (Keen, 1971, fig. 880). Nomenclature! Commentaryi Natica amlculata Philippi, 1849, has sometimes been considered a Junior synonym of £. (£.) nanamaftnwi«. but its type figure clearly places it as a synonym of £. (£.) ravldua (Souleyet, 1852). Discussioni The discussion of this species is included with that of the closely related species £. .(£*) uber (Valenciennes, 1832). fQttniCgfl (Pollnlces) lntemeratus (Philippi, 1853) Plate 1, figure 6 Natica interne rata Philippi, 18531233-2341 Philippi, 1853< l29, pi. 18, fig. 10 (plate 185l)i Sowerby, 1883t87, pi. 4, fig, 44t Tryon, 1886i46, pi. 18, fig, 83, pi. 19, fig. 93. Polinlces uber var. lntemeratus (Philippi), Dali, 1908i334 Cas £. y. var. Intemerata1. Polinlces lntemeratus (Philippi), Palmer & Hertleln, 1936i 78, pi. 19, fig. 3 Cas £, Intemerata1. Polinlces uber lntemeratus (Philippi), N. Smith, 1944i12 Cas £. y. intemerata1. Polinlces (Pollnlces) lntemeratus (Philippi), Keen, 1958i j22, fIgT Zo9) Parker* 19o4i153, pi. 6, fig. 8 [>s £. (P.) IntemerataJi Keen, I97li478, fig. 877. Natica alabaster Reeve, 1855ipl. 9, figs. 33a,b.t Carpenter, 1857i292. 81 Natlca^r*^y^mn Reeve* 1855tpi. 12, figs. 47a,b( Keen* Descriptioni Color.--Shell exterior and interior white* with pale buff to yellow band below suture* First 1-2 protoconch whorls medium brown. Periostracum thin* usually pale yellowish white* but ranges to medium yellowish brown* may have irregular rust brown stains* Size.--Averagei height 18 mm* diameter 14.5 mmj largest speciment height 29.5 mm* diameter 24.3 mm £AHF 381-35, Bahia Independencia* Peru}. Shell Form.--Shell globose to slightly elongate, spire moderately elevatedi body whorl not greatly elevatedi shoulder slightly flattened* may be slightly concave on final adult whorl * shell thickness average t nuclear whorls 3, with short, weakly developed radial wrinkles ex tending from suturei postnuclear whorls 3^j suture very slightly impressed. Spiral sculpture of microscopic, closely spaced* very weak costellaei axial sculpture of incremental growth lines. Parietal callus very heavy, thickly filling posterior apertural anglei anterior lobe weak. Umbilicus moderately open, arcuate, never slit- like. Umbilical callus comes to a point anteriorly* ex pands gradually to its midpoint, then is pinched to form a rounded sulcus before expanding to meet parietal callusi shallow groove traverses callus where it merges with 82 parietal callust anterior Inner lip thickened. A low* sharp ridge runs along mldllne of umbilical channel and Is expressed on columella as a low fold. Operculum.--Chltinous. filling aperture. Specimens Examined! 429. Geographic Occurrence and Ecologyi Cedros Island, western Baja California, Mexico (28o10'N), throughout the Gulf of California, and south to the Galapagos Islands and Bahfa Independencla, Peru (14°14'S). Living on sand bottoms In depths of 9 to 160 meters. Stratlgraphlc Occurrencei Pliocene ^Pleistocene?Ji Carmen Island, Gulf of California, Mexico (CAS). Type Localitiesi Natica intemerata - Gulf of California, Mexico (Philippi, 1853). Natica alabaster - Mazatlan, Mexico (Reeve, 1855). Natica raoulum - Palta, Peru (Reeve, 1855). Type Materiali Natica Intemerata - Unknown, presumably in BM(NH) (Dance, 1966), Natica giafraatgr - Unknown, presumably in BM(NH), or National Museum of Wales, Cardiff, Wales (Dance, 1966). 83 Natica raouiun - Sane as above. Discussioni The discussion of this species is included with that of the closely related species £. (£.) uber (Valenciennest 1832). PolLnlces (Polinlces) otla (Broderip f i t Sowerby, 1829) Plate 1, figures 7, 8 Natica otls Broderip & Sowerby, 1829i372i Gray* 1839i136t Philippi, 1852i57, pi. 9, fig. 4 (plate 1850)i Tryon, 1886(43-44, pi* 12, fig. 2, pi. 17, figs. 7-72 Cnot pi. 19, fig. 91, - Natica perspicua Recluz, 1850, Rec«it, PhllippinesjT Pollnlces otla (Broderip & Sowerby), Carpenter, 1864i624t Steams, 18941196 Cas "Polvnlces"11 Dali, 190912351 M. Smith, 1944*12, fig. 134 [as "Broderip" only]. Lunatla otls (Broderip f i t Sowerby), Steams, 1894*409. Pollnlces (Pollnlces) otls (Broderip f i t Sowerby), Keen, l558i323, fig. 271i Keen, 1971.478, fig. 879. Pollnlces otls var. fusca Carpenter, 1864*523, 624 fnomen nudunli Carpenter, 1872*9, 110i Steams, 1894»19o Las "Polvnlces**7* Palmer, 1963| 376| Tryon, 1886*44 Cas Junior synonym of Pollnlces otls]. Natica fusca Sowerby, 1883*89, pi. 8, fig. 104 described]. Runs subfuscs Dali, 1919*35, Keen, 1958*323 as Junior synonym of Pollnlces otls . Pollnlces subfusca (Dali), M. Smith, 1944*12, fig. 129. Hfttlga 1843.2ll( Adams, 1852.207| KilipplT l” "* 131-132, pi. 18, fig. 131 Tryon, 1886*44 Lea Junior synonym of Polinlces otlsl. Natica aalapaaoaa Recluz, 1844*213i Philippi, 1853*131, pi. 18, fig. 12 (plate 1851)i Reeve, 1855, pi. 19, figs. 86a,bt Carpenter, 1857*i85i Sowerby, 1883*89, 84 pi. 7, fig. 95i Tryon, 1886i44, pi. 17, fig. 71 fas Junior synonym of Pollnlces otls]. Lunatla aalapagosa (Recluz), Carpenter, 18571360 Qetnenda- tIon of name)i Steams, 18941406. Pollnlces (Polinlces) aalapaaosus (Recluz), Keen, 1971t 478, fig. 879 [photo of syntype). Natica unlmaculata Reeve, 1855, pi. 19, figs. 85a,bi Carpenter, 185712931 Sowerby, 1883183, pi. 8, fig. 105i Tryon, 1886*46-47, pi. 19, fig. 96. "Natica persplcua Recluz" Cnot "Natica" persplcua Recluz, 1859ij79-380, pi. 14, figs. 1,2, Recent, Philippine islands)) Reeve, 1855) pi. 4, fig. I2i Carpenter, 1857t292) Sowerby, 1883i87, pi. 6, fig. 70. not Natlea persplcua Pictet & Roux, 1849t51, pi. 18, figs. 4a,b Cfossil, Switzerland). Descriptioni Color.--Shell white with a pale yellow to orange- brown band below suturei umbilicus white, may be partly or completely dark brownj umbilical callus rarely all white, usually with an anterior spot of medium to dark brown. Interior white to brown. Nuclear whorls light to dark brown. Periostracum thin, medium yellowish white. Size.--Average slzei height 35 mm, diameter 30 mmi largest speciment height 44.1 mm, diameter 40.7 mm (USNM 4124, Cabo San Lucas, Baja California, Mexico • Shell Form.— Spire moderately elevated, shell elongate) shoulder flattened) shell thick, nuclear whorls 3%, with weak radial wrinkles on shoulder) postnuclear whorls 3) 85 suture very slightly inpressed. Spiral sculpture of minute, weakly developed costellae% axial sculpture of in cremental growth lines. Parietal callus massive, thickly filling posterior apertural angle, with a weakly to strongly developed transverse groovei anterior lobe broad, overhangs umbilicus. Umbilicus broadly open, with distinct posterior sulcusi channel broad. Umbilical callus flattened, slender anteriorly, broadens to greatest width at its midpoint, then Is abruptly constricted before ex panding again to meet parietal callusi funicle broad, distinct. Operculum.--Chltinous■ filling aperture. Specimens Examinedi 172. Geographic Occurrence and Ecologyi The Gulf of California, Mexico, to Santa Elena, Ecuador (2°11'S). Workers begin ning with Tryon (1886) have reported this species from the Galapagos Islands, but I have seen no specimens from there. Such records are probably based on Polinlces n. sp., dis cussed herein. Most specimens of £• (£.) otls are from the Gulf of California and Panama, rarely from elsewhere. Occurs most commonly offshore in depths of 20 to 297 meters, rarely intertldally on mud flats. Type Localitiesi &U££ at1a - Mazatlan, Mexico (Broderip & Sowerby, 86 1829). Natlea fuaca - Mazatlan, Mexico (Sowerby, 1883). Ruma subfuaca - Panama (Dali, 1919). Natlea salanaonensis - "Salango, West Colombia [Ecuador}i found In sandy mud” (Recluz, 1843). Natlea aallapajtosa - Galapagos Islands. Ecuador, "found In coral sand at Albemarle Island" (Recluz, 1843). Natlea unlmaculata - Mazatlan, Mexico (Reeve, 1855). Type Materiali Natlea otls - Unknown, presumably In BM(NH) (Dance, 1966). Natlea fusca - Unknown, presumably In fiM(NH) (Dance, 1966). Ruma subfuaca - USNM 46544, holotype. Natlea salanaonens1s - Unknown, presumably In Museum d'Hlstolre Naturelie, Laboratolre de Malacologle, Paris (Dance, 1966), Natlea aallapaaosa - As above* Natlea unlmaoulata - Unknown, presumably In BM(NH) (Dance, 1966). Nomenclature1 Commentaryi Colors of the aperture, callus and umbilicus of £. (£•) otls range continuously from brown to white In various combinations, some of which have served as the basis for species names. Two of the more recently 87 used names are £. Ralapaaosus. described as having a brown umbilicus and aperture but a white callus* and £. uni- macula tus . with a brown spot on the callus and white or brown aperture and umbilicus. As noted above* Mazatlan, Mexico* Is the type locality for £. <£.) otls and two of its synonymous species. Pollnlces otls var. fusca Carpenter* 1864* was pro posed without description and succeeding authors listed it only as a name until Sowerby (1883) validated it as Natlea fusca. although he attributed the species to Carpenter. Dali (1919) proposed Ruma subfusca as a replacement for Carpenter's nomen nudum, apparently unaware of Sowerby*s previous validation of Natlca fusca. The syntypes of £. Ralapaaoaus (figured in Keen* 1971) are considered to be specimens of £. (£.) otls. How- ever* £. (£.) otls is not known to occur in the Galapagos* so the locality data with the syntypes is probably in error. All of the literature illustrations of £. ftalaoa- r o s u s known to me show specimens of £. (£*) otls. Galapagos records might be based on specimens of £. (£.) interneratus (Philippi), because this species is found there and is often raisldentifled in collections as £. (£*) otls. Discussioni Among eastern Pacific natlcids, £. (£.) otls is most similar to £. (£.) lntemeratust they have widely 88 overlapping geographic ranges, and they have often been confused in collections* These species are closely related but do not intergrade, and £. (£.) otis is distinguished by its more elongate shell with a higher spire, broader umbilicus lacking a sharp spiral ridge, and greater size* Specimens of £. (£.) inteaeratus are also never stained with brown on their calli. Except for colored specimens, early Juveniles of £. (£.) otis are often indistinguishable from those of £. (£.) Inteaeratus. £. (£.) uber and £. (£.) panaaaensls* The problem of distinguishing these species is discussed further under £. uber. Natica ravida Souleyet, 1852(576,582, pi. 35, figs. 12-15 f as MNatice Jaune-roux” in plate captionli Reeve, 855ipl* 16, figs. 68a,bt Sowerby, 1883(89, pi. 5, fig. 53i Tryon, 1886(43, pi. 15, fig. 38. Pollnlces ravidua (Eydoux & Souleyet) Calc), Dali, 1909ai 235i Burch, 1946(26. Natica aaiculata Philippi, 1849i155i Philippi, 1851(98, pi, W7 tig. 4. Pollnlces (Pollnlces) ravlduB (Souleyet, 1852) Plate 1, figures 9, 10 ravldus (Souleyet), Keen, 1971(480, Type Localitiesi Natica ravida - Paita, Peru (Souleyet, 1852). 89 Natica amlculata - Palta, Peru (Philippi, 1849). Type Materiali Natica ravida - BM(NH), 3 syntypes, illustrated by Keen (1971). Natica amlculata - Unknown, presumably In BM(NH) (Dance, 1966). Geographic Occurrence and Ecologyt Santa Elena, Ecuador (2°10*S) to Isla Lobos de Tierra, Peru (5o20'S). Two of the seven specimens seen by me are from Palta, Peru (5°S), which is the type locality. Keen (1971) extends the range north to Panama, but I have seen no specimens from there. No ecological data accompanies any of the museum speci mens, but one specimen was found dead on the beach at Palta, which is perhaps evidence for a near-shore habitat. Specimens Examinedi 7. Descriptioni Color.-«Shell whitei thin periostracum light orange- brown to chocolate browni callus and interior white. Form.--Average alzei height 31 mm, diameter 27 mmi largest specimeni height 44.9 mm, diameter 39.6 mm fUSNM 538002, Isla Lobos de Tierra, Peru]i spire low to moderate ly elevatedt shell thick, of 2k nuclear and 4 postnuclear whorls, clearly set off from each other* suture distinct but not impressedi shoulder flattened, usually slightly 90 concavei spiral sculpture of minute, indistinct costellaei axial sculpture of incremental growth lines. Parietal callus massive, thickly filling posterior apertural anglei margin slightly concave, with a broad anterior lobe that overhangs umbilicus) a very weakly Impressed groove partly traverses the parietal callus at its margin, below center of the callus. Umbilicus broadly open, exposing earlier whorlst inner (central) portion of channel abruptly deepened, indenting inner lip. Umbilical callus narrow anteriorly, broadly expanded posteriorly to meet parietal callus and conceal upper part of umbilicus* Operculum.--Chltlnous■ filling aperture. Dlscussloni This species is easily distinguished from other natlclds by the combination of its massive form and simple, broadly open umbilicus in which earlier whorls are exposed. Pollnlces (fflHolSfifl) blfasciatus (Griffith f i t Pidgeon, 1834) Plate 1, figure 11 Natica blfasclata Griffith f i t Pidgeon, 1834, vol. 12i598, pi. 1, fig. 2. Natica blfasclata "Gray" [error for "Griffith & Pidgeon"}i Philippi, 1852159, pi. 9, fig. 7 (plate 1850)) Reeve, 1855ipi. 10, figs. 40a,b| Carpenter, 1857i292i Sowerby, I883i93, pi. 4, fig. 45* Try on, I886i44, pi. 18, fig. 75| Hanna & Hertlein, 1927*143, 146i M. Smith, 1944il93, pi. 4, fig. 45. 91 Polynlces blfaacUta -Gray," Steams, 1894*195. Pollnlces blfasclatus "Gray," Durham, 1950*127, pi. 34, fig. T. Pollnlces (Pollnlces) blfasclatus (Griffith & Pidgeon), Grant & Gale, 1931*800 fas £. (£.) blfasclatus “Gray“31 Keen, 1958*322, fig. 266 Cas £. (P.) blfasclatus “GrayM3i Keen, 1971*478, fig. 873, pi. 14, fig. 4. Description* Color.--Shell tan to medium brown, with 4 narrow, spiral white bands* one at suture, two near middle of whorl, and one low on basei Irregular chocolate brown band may occur between suture and uppermost white band) umbili cal callus and parietal callus chocolate brown, may be partly white* callus filling at posterior apertural angle white. Nuclear whorls white or reddish brown. Interior white, with margin of aperture sometimes brown. Perio- stracum thin, pale yellowish white. Size.--Average* height 35 mm, diameter 29 mm* larg est specimen* height 60.7 mm, diameter 46.7 mm fCAS 34874, San Felipe, Baja California, Mexico}. Shell Form.--Shell elongate, spire moderately elevat ed* body whorl not strongly inflated* shell thick* nuclear whorls 3%, smooth* postnuclear whorls 4* sutures very weakly Impressed. Spiral sculpture of minute, weakly developed, obscure costellae* axial sculpture of Incre mental growth lines. Parietal callus massive, heavily filling posterior apertural angle, with a shallow trana- 92 verse groove at Its midpointi anterior lobe thick, not clearly set off from remainder of callus. Umbilicus open, relatively large to small, depending on callus develop ment! channel broad, sometimes bounded anteriorly by low ridge. Umbilical callus narrow anteriorly, gradually broaden to meet parietal callus and conceal posterior end of umbilicus* Basal lip thickened. Operculum.--Chitlnous. filling aperture. Specimens Examinedi 427. * Geographic Occurrence and Ecologyi Bahia Magdalena, western Baja California, Mexico (24°30*N), throughout the Gulf of California and south to Panama Bay, Panama (8°N). Not common except in the Gulf of California. This is chiefly an Intertidal species, but is known rarely in depths to 60 meters. Stratigraphlc Occurrencei Lower Pliocene (?)i San Marcos Fm., San Marcos Island, Gulf of California (Dur ham, 1950). Upper Pliocenei Carmen Island, Gulf of California (Durham, 1950). Pliocenei Coronados Islands, Gulf of California (Hanna & Hertleln, 1927). Pleistocene t Santa Elena Bay, eastern Baja California (CAS)t Carmen Is land, Coronado Island, Santa Inez Bay, and Punta San Telmo, Gulf of California (Durham, 1950). 93 Type Localityi Unknown (Griffith & Pidgeon* 1834). Type Material! Unknown* presumably in BM(NH) if included with J. £. Gray collection (Dance. 1966). Nomenclatural Commentaryi Most workers have attributed this species to J. E. Gray, because the original citation in Griffith & Pidgeon (1834) is "Natica blfasclata. Gray." However, there is no other evidence of Gray having des cribed the species, although it nay have been a manuscript name of his. Keen (1971) first attributed the species nane to Griffith & Pidgeon. Discussioni The Pliocene specimens 1 have seen of £. (£.) blfasclatus from the Gulf of California islands are well preserved and may actually be of Pleistocene age. Subgenua Eusolra Agassiz* 1838 Eusplra Agassiz in Sowerby* 1838i14. Type-species« Annullaria slaaretlna Lamarck, 1804, by subsequent designation (Harris, 1897>264). Paleogene of France and England. Lunatla Gray* 1847 il49. Type-speciesi Natica ampullaria Lamarck* by monotypy. Paleogene of France. 94 Diagnosis« Shell small to very large, globosei whorls moderately to greatly Inflated) suture weakly to deeply Impressed. Umbilical callus usually slender, may be slightly broad, never closing umbilicus. Parietal callus thin to moderately thick. Discussioni Euspira is characterized by a globose shell, open umbilicus and slender umbilical callus that becomes thicker posteriorly. Dali (1908i333) has commonly been cited as designator of the type-species, but 1 find that Harris' (1897) cita tion is earlier. The type-species designated by Harris (1897) is one of several fossils included by Agassiz (in Sowerby, 1838) in Euspira■ The subgenus is primarily found in temperate waters, but ranges into the Arctic. Lunatla has been used by numerous workers, but is clearly a Junior synonym of Euspira. Both of these sub genera, however, are based on Paleogene type-species, and it remains to be seen whether Euspira is being accurately applied to Recent species. Pollnlces (Euspira) ga Hanoi Dali, 1909 Plate 1, figures 12, 13, 14 Pollnlces (Euspira) ga Hanoi Dali, 1909(88-89, pi. 5, figs. 12, 13| Etherington, 1931i94-95, pi. 12, figs. 1,5, 8,20i Grant & Gale, 1931i805i Clark, 1932*825) Weaver, 1942*344, pi. 70, figs. 11,14) Weaver, 1945i 57. 95 Pollnlces aallanol Dali, Arnold & Hannibal, I9l3i576, 584, 591, 592i Weaver, 1916i2l6, 218 fas "Polvnlces**1. Natica «allanol (Dali), Howe, 1922isp. list. Natica (Euspira) aaHanoi (Dali), ?Lutz, 1951•380, 391, pi. 18, fig. 3 Lfigure unrecognizable]. Natica (Neverlta) orbicularis Nomland, 1916»207, pi. 10, figs. 4a,b| Keen & Bentson, 1944*177. Natica orbicularis Nomland, Moody, 1916*44, pi. 2, fig* 6* Nomland, 1917bi213, 221i Keen & Bentson, 1944*177. Pollnicea (Euspira) orbicularis (Nomland), Grant A Gale, 1931*805 raa P. C?E.) orbicularis 1* Soper & Grant, 1932*1063. Pollnlces orbicularis (Nomland), Adegoke, 1969»i66. Neverlta (Glossaulax) n. sp.?, Addlcott, 1970*70, pi. 6, figs. 1, 9. Description* Size,--Average * height 43 mm, diameter 42 mmj larg est specimen* height 57.4 mm, diameter 53*3 ram (incom plete) [SU NP220, Sylvia Creek, Montesano, Washington, Pliocene7]. Shell Form.--Shell globose, spire low to moderately elevated, rarely hlghi body whorl moderately inflated, may be slightly flattened at periphery! shoulder distinctly flattened at suture, may be slightly concavef shell thick ness averagei nuclear whorls 2, sculptured with low, closely spaced axial costellaet postnuclear whorls 4* suture moderately to slightly impressed. Shell smooth ex cept for minute incremental growth lines. Parietal callus thin, thickens into posterior apertural anglei anterior 96 lobe moderately developed, merges smoothly with umbilical callus. Umbilicus broadly open, occupied to varying degree by umbilical callus. Umbilical callus confined to posterior half of inner lip, clearly divided into two lobes by sharply incised transverse groove* lobes flatten ed or rounded with swollen appearance* anterior lobe usually more massive, but posterior lobe may be more elongatei funlcle low, inconspicuous. Anterior inner lip and basal lip slightly thickened. Operculum.— Unknown. Specimens Examined* 208. Stratiaraphic Occurrence* Ranges from Lower Miocene (?) to Middle Pliocene. Middle Ollaocene to Lower Miocene* Poul Creek Formation, Yakataga area, Alaska (Clark, 1932). Lower Miocene (?)i Sooke Formation, southern Vancouver Island, British Columbia (Arnold & Hannibal, 1913). Lower Miocene to Middle Pliocenei Yakataga Formation, Yakataga area, Alaska (Clark, 1932). Middle Miocene* Astoria For mation, northwestern Washington (Arnold & Hannibal, 1913), Montesano (SU), Grays Harbor County (UCB), Washington, and Lincoln County (UCB), Newport (CAS), and Coos Bay (UCB), Oregon* Monterey Formation, Briones Valley quadrangle, California (UCB)* Sobrante Formation, Pacheco syncline, California (Lutz, 1951)* Temblor Formation, Caliente quadrangle, California (UCB)* upper Olcese Sand and lower Round Mountain Silt, Kern River area, California (Addicott, 1970i USGS)» Upper Miocene to lower Pliocenei Monteaano Formation, Montesano (CASr SU), Grays Harbor County (UCB), Washington. Lower Pliocenei Empire Formation, Coos Bay (CASi Weaver, 1945), and Coos Bay-Cape Blanco area (Arnold & Hannibal, 1913), southwestern Oregon. Middle Pliocenei Merced Formation, San Francisco peninsula and Eel River Valley, California (Arnold A Hannibal, 1913)j Etchegoin Formation, San Miguel quadrangle, Monterey County (SU), Coallnga area (Nomland, 1916), California. Type Localities! Pollnlces (Euspira) aallanol - Coos Bay, Oregon. Empire Formation, lower Pliocene (Dali, 1909). Natica (Neverlta) orbicularis - UCB 2679, middle of north line, SEfc NE^, section 7, T22S, R15E, Fresno County, California. Jacalltos Formation, lower Pliocene (Nomland, 1916« Keen & Bentson, 1944), Tvoe Material! Pollnlces (Euspira) aaHanoi - Holotype, USNM 153916. Natica (Neverlta) orbicularis - Holotype, UCB 12058. Nomenclature! . pollnlces (Euspira) orbicularis is made a Junior synonym of P. (£.) aallanol here for the first time, based on a comparison of type specimens. The types of £. (£.) aallanol are poorly preserved, but some 98 clearly show remnants of the diagnostic bilobed umbilical callus that is displayed well on the holotype of £. (£.) orbicularis. Although the operculum of this species is unknown, its shell morphology unequivocally allies it with a number of living Follnicinae species, such as £. (£•) lewis 11 (Gould, 1847) and flgYSrUS (Glossaulax) recluslana (Deshayea, 1839). The inclusion of this species in the calcareous-operculate genus Natica by some past workers is therefore in error. Discussiont Recognition of this species has been difficult because the type specimens are poorly preserved, with critical umbilical features missing from roost types. How ever, unmistakable remnants of the bilobed umbilical callus on some specimens, plus details of shell shape, size and stratlgraphlc occurrence, leave no doubt as to the identity of Dali's species. The umbilical callus varies in prominence and the degree to which it occupies the umbilicus. Juveniles have flattened calll, with a fairly weak transverse groove, the posterior callus lobe is sometimes very small, whereas the umbilical callus of an adult has a deeply Incised trans verse groove and a swollen appearance to the lobes. The compact, bilobed umbilical callus is very similar to that of ft. (G.) lamesae (Moore, 1963), an 99 early to middle Miocene species. £ 4 . (G.) lamesae Is easily distinguished by Its greatly flattened base, wider umbilicus, lower spire and more Inflated body whorl, but the two species occur throughout much the same region and may have a common ancestor. The posterior umbilical callus lobe of £, (£.) «allanol Is never detached from the underlying whorl as seen In some specimens of £ £ , (G.) Jamesae. Individuals with elevated spires are rare, and when they occur with a prominent umbilical callus they resemble the alta form of £ £ . (G.) recluslana. The resemblance may be quite close, but £. (£.) aallanol specimens have thinner shells, less elongated posterior lobes of the umbilical callus, and more flattened bases than specimens of £* , (G,) recluslana. Policlnes (£.) aallanol Is probably an off shoot of the vigorous £ J . (G.) recluslana stock that has produced such a varied array of species and forms from the Miocene or earlier to the present day. Pollnlces CEuspira) aallanol Is also similar In form to £. (£.) lewis11 and may be ancestrally related to it. It differs from £. (£.) lewlsll by its smaller size, lack of a spiral flexure on the shoulder, more abruptly tapered anterior margin of the umbilical callus, and more deeply Incised callus groove* However, the general aspects of shell form and umbilical details are quite similar, and £, (£.) aallanol is the most likely ancestor for the large £. 100 (£.) IfigLftU. Pollnlces (Euspira) dlabloensls (Clark, 1915) Plate 1, figure 15 Natica (Euspira) diabloenais Clark, 1915bt486-487, pi. 68, fig• 7* Keen & Bentson, 19441176. Natica diabloenais Clark. Nomland, 19l7bi301. Pollnicea (Pollnicea) diabloenais (Clark), Addicott & Vedder7l963i66. Euaolra diabloenais (Clark), Adegoke, 1969i169, Description* Size.--Averagei height 45 mm, diameter 37 r a r o j larg est specimeni height 48.8 mm (incomplete), diameter 50.4 mm (incomplete) CSU 39471, Temblor Formation?, Santa Clara County, California^. Shell Form.--Shell elongate, spire elevated* body whorl not greatly inflated* whorls evenly rounded except for slight flattening near suture* shell thickness average* whorls 5^* suture moderately impressed. Shell smooth except for minute incremental growth lines. Parietal callus thick, solidly filling posterior apertural anglei anterior lobe very weak, slightly projecting above umbilicus. Umbilicus narrowly open, lacking a distinct sulcus* channel narrow, only slightly tapering anteriorly* umbilicus bounded by a broad cord-like angulation on base. Umbilical callus narrow, not clearly set off from inner 101 lipI funlcle Indistinct. Anterior inner lip and basal lip thickened. O p e r c u l u m . — Unknown. Specimens Examinedi 13. Stratlaraphlc Occurrencei Ranges from Middle (?) to Upper Miocene. Mlddle Miocene (?)* Temblor Formation (?), Calaveras Valley* Santa Clara County, San Jose quadrangle, California (SU). Upper Miocenei Santa Margarita Forma tion, north Coalinga area (Clark, 1915bi Nomland, 1917b), Domengine Ranch quadrangle (UCB; Adegoke, 1969), Cali fornia i upper San Pablo Formation, central California (Clark, 1915b)t Neroly Formation, Tassajara quadrangle, California (UCB). Discussioni The holotype is by far the best-preserved specimen, and no juveniles are known, Pollnlces (Euspira) dlabloensls differs from the Miocene species £. (£.) vlctorlanus Clark & Arnold, 1923, by its large size, more elevated spire, more strongly impressed suture, thinner parietal callus, and thinner inner lip-umblllcal callus. It differs from £. (£,) lewlsll (Gould, 1847) by its smaller size, lack of a shoulder flexure, narrower umbilicus, much weaker anterior lobe of the parietal callus, and lack of a transverse groove on the callus. Similarly, £. (£.) diabloenais differs from £. (£,) gaHanoi Dali, 102 1909, the probable ancestor of £. lewisli. by Its higher spire, narrower umbilicus, and Its much narrower and un grooved umbilical callus. The single Middle Miocene occurrence of £, (£.) dlabloensls should be questioned until additional speci mens of the same age are found. Until then, it is best to consider this as an exclusively Late Miocene species. Pollnlces (Euspira) lewlsll (Gould, 1847) Plate 1, figures 16, 17 Natica lewlsll Gould, 1847*239, Gould, 1852i211-212i Gould, lo56ipl. 15, fig. 253t Gould, 1862*49, 244* Try on, 1886*35, pi. 9, fig. 70, pi. 13, fig, 11, not pi. 13, fig. 12 L~ £• relnlanus (Dunker, 1877), Re cent, JapanJt Martin, 1916*230, 239, 257 Cas "cf,"]* Weaver, 1916*i72, 176 £as "cf."]t Johnson, 1964*101. Lunatla lewisli (Gould), Carpenter, 1864*661t Keep, 1888* 45, fig. 25i Arnold, 1907*25, pi, 54, fig. 3| Arnold & Anderson, 1907*144, pi. 21, fig. 3, pi. 44, fig. 1* Arnold, 1908*353-354t Arnold, 1910*31, 150, pi. 22, fig. it Arnold & Anderson, 1910*109, 110, 127, 133, pi. 44, fig. It Jordan, 1924*i50t Woodrlna et al.. 1940*86, pi. 15, fig. 8, pi. 31, fig. 5* Keen & Bentson, 1944*168i Woodrlna et al.. 1946*72) Wood- ring & Bramlette, 1950*73, pi. 12, fig. It Winterer & Durham, 1962*296) Stanton, 1966*23i Adegoke, 1969* 166-167. Polvnlces (Lunatla) lewisli (Gould), Arnold, 1903*315, pi. io7 ttg. 14. Pollnlces (Euspira) lewlajLl (Gould), Dali, 1921* 1,651 Oldroya, 1954*ioz-io3, pi. 36, fig. It Oldroyd, 1927*7291 Grant & Gale, 1931*804-805, figs. 15a,bt Soper & Grant, 1932*1063) Willett, 1937*400) Burch, 1946*29) Glen, 1959*184 £as -cf. £, (£.) lewlsii"]* Faustman, 1964*136. 103 Euspira lewlsll (Gould), Clark, 1931isp. list* Valentine, 1956i2o0* Valentine, 1957*300 [>s "£* lewis1"It Valentine, 1961*335, 336, 338 fa* "£• lewlsl"! * Valentine & Meade, 1961*17,19,23,27 fa* "£. lewis1*1» Valentine, 1962*96. Pollnlces lewlsll (Gould), Packard, 1918*325, pi. 38, fig. 1 [as "£. lewlsl"3 * Jordan, 1936*2461 Arnold & Hanni bal, 1913*598* Keen, 1937*44* DeLong, 1941*241* Smith & Gordon, 1948*199* MacGlnltle & MacGlnltle, 1949*370, 376, fig. 238* Kanakoff & Emerson, 1959*30* Addlcott & Emerson, 1959*16* Emerson & Addlcott, 1953*440* Durham & Addlcott, 1965*12* McLean, 1969* 37, fig. 19.1* Wright, 1972*690. Natica herculea Mlddendorff, 1849*424-425, pi. 7, figs. 5- 7* Philippi, 1853*148-149* Gould, 1862*244* Carpenter, 1864*531* Sowerby, 1883*77, pi. 1, fig. 3. Natica herculea forma normalls Mlddendorff, 1849*424. Natica herculea forma elatlor Mlddendorff, 1849*424. Natica herculea var. lewlsll (Gould), Sowerby, 1883*pi. 1, fig. 3. Natica alaIda Gould, 1848*73, Gould, 1852*214* Gould, l85oipi, 15, figs. 256, 256a* Gould, 1862*50, 244* Carpenter, 1864*531* Johnson, 1964*38-39. Euspira alalda (Gould), Dali, 1919*352. Pollnlces (Euspira) alaldus (Gould), Dali* 1921*164 [As "£• (£7) ala1daH1 * Oldroyd, 1924*l62* Oldroyd, 1927*730* Bure h, 1946 * 29. Pollnlces alaldus (Gould), Keen, 1937*44, Description* Color.--Shell exterior white. Perlostracum thin, light to medium yellowish brown, often stained rust brown, especially on shoulder, becoming thicker, darker and scaly within umbilicus. Interior white or clouded with pale brown* umbilical callus mottled with pale brown and white, 104 parietal callus usually white. Size.--Averagei height 90 mm, diameter 75 mmi largest specimeni height 166.0 mm! diameter 131.0 mm [Santa Monica Bay, Californiai LACM records]]. Shell Form.--Shell globose to elongate, spire moderately to strongly elevatedt body whorl Inflated} adult whorls with a weakly to strongly concave shoulder profile, sometimes forming a tabulation below suturei shell average or greater In thickness) nuclear whorls 2* f , smooth) postnuclear whorls 7. Spiral sculpture of minute, weakly developed, closely spaced costellae, which may be absent) axial sculpture of Incremental growth lines that may form Irregular rugae below suture and In umbilicus. Parietal callus average to thick, moderately to heavily filling posterior apertural angle) anterior lobe distinct. Umbilicus open, relatively broad to narrow, not exposing many earlier whorls) sculptured with Irregular axial wrinkles and weak spiral costellae) often with broad angulation along basal margin of umbilicus. Umbilical callus narrow, broadens posteriorly, divided Into two lobes by weakly to strongly developed transverse groove) posterior lobe may be swollen and partly conceal posterior of umbilicus) funicle lacking) anterior Inner lip thickened. Operculum.--ChitInous. filling aperture. Specimens Examinedi 225. 105 Geographic Occurrence and Ecologyi Northern Vancouver Island* British Columbia, Canada (5l°N), to Isla San Geronimo, western Baja California, Mexico (29°47'N). Liv ing intertidally, especially in embayments, and offshore in depths to at least 200 meters, on sandy bottoms. Willett (1937) reported specimens in southeastern Alaska, without citing a specific locality. Stratlgraphlc Occurrence! Ranges from Upper Miocene (7) and Lower Pliocene to Recent. Upper Miocenei Castaic Formation, near Castaic, Los Angeles County, California (Stanton, 1966). Lower Pliocene» Empire Formation, Max- field Creek, Washington (SU)i Jacalitos Formation, Coallnga area, California (Arnold & Anderson, 1910, as N?")l Pancho Rico Formation, Salinas Valley, California (Dur ham & Addlcott, 1965). Middle Pliocenet Etchegoin Forma tion, Coallnga area (CASf Arnold & Anderson, 1910i Martin, 1916i Adegoke, 1969) and Kettleman Hills (SU| Woodring, et al.. 1940), California. Middle to Upper Pliocenei Rio Dell Formation, Scotia quadrangle, Humboldt County, Cali fornia (Faustman, 1964). Middle Pliocene to Pleistocenei Merced Formation, San Francisco South quadrangle (UCBf Martin, 1916i Glen, 1964), Bolinas Bay (Martin, 1916), San Mateo County (UCB), near Santa Rosa (CAS), and Santa Cruz quadrangle (Arnold, 1908), California. Upper Pliocenei 106 Pico FormatIoni southeastern Ventura basin (Winterer & Durham, 1962), Anaheim quadrangle (UCB), and Piro quad rangle (UCB), Califomiai Pico Formation, Sand Canyon, Los Angeles County, California (LACM)t San Joaquin Formation, Kettleman Hills (USGSi Woodrlng ££ a^., 1940) and La Clma quadrangle (UCB), Californiai San Diego Formation, San Diego County, California (LACM)t Fernando Formation, central Los Angeles basin (LACMj Soper 6 Grant, 1932) and Newport Bay (LACM), California. Upper Pliocene to Plelsto- cenei Saugus Formation, Pirn quadrangle, California (UCB). Pliocenei Purlsima Formation, Halfmoon Bay (Martin, 1916), Ano Nuevo quadrangle (SU), San Juan Bautista quadrangle (UCB), San Mateo Mountains (SU), Capitol quadrangle (UCB), Santa Cruz quadrangle (Arnold, 1908i UCB), and Soquel quadrangle (USGS), Californiai middle Wildcat Formation, Humboldt County, California (Martin, 1916)i Foxen Mud stone and Careaga Sandstone, Santa Marla basin, California (Woodrlng & Bramlette, 1950)j Port Orford Formation, Cape Blanco quadrangle, southwestern Oregon (USGS)f Imperial Formation, southwestern San Diego County, California (UCB)i Coallnga quadrangle (UCB)i San Benito quadrangle (UCB)i La Clma quadrangle (UCB). Lower Pleistocenei San Pedro Formation, Deadman's Island, San Pedro, California (UCBi CAS)i San Pedro Sand Member (LACMiCAS) and Timms Point Silt Member (LACMi Clark, 1931) of San Pedro Formation, San Pedro, Califomlaj "San Pedro Formation,** near Camulos, 107 Ventura County, California (CAS)i Elk River Formation, Cape Blanco quadrangle (USGS) and Curry County (UCB), Oregon* Upper Pleistocenei Palos Verdes Sand, San Pedro quadrangle (LACMj UCB) and Torrance quadrangle (UCB), Los Angeles County, and Newport Bay (UCB), Orange County, Cali fornia) Bay Point Formation 7, San Ysidro quadrangle, San Diego County, California (UCB)i Bahia San Quintin (UCB) Jordan, 1924) north of Punta Cabras (UCB), and Magdalena Bay (Jordan, 1924), Baja California, Mexico* Type Localitiesi Natica lewisil - Puget Sound, Washington, Gould, 1847). Natica herculea - "Kolonle Ross in Neukalifornlen” (Mlddendorff, 1849). Natica herculea forma normalis - Same as above. Natica herculea forma elatlor - Same as above. Natica alalda - Rio Negro, Argentina (Gould, 1848) Q>robably Classet, Oregon]. Type Materiali Natica lewlsll - Holotype, probably USNM 3903 (John son, 1964). Natica herculea - Academy of Sciences, Leningrad, U.S.S.R. (Dance, 1966). Natica herculea forma normalls - Same as above. Natica herculea forma elatlor - Same as above. Natica alalda - Unknown (Johnson, 1964). 108 Nomenclatural Commentaryi Tryon (1886) considered the Japanese species £. relnlanus (Dunker, 1877) a Junior synonym of £. (£.) lewisli. As noted and illustrated in Kuroda fit Al* (I97lil2l, pi. 18, fig. 2), £. relnlanus is smaller than £. (£.) lewisli (height 31.8 mm, diameter 30.0 mm) and has a thicker shell with the umbilical callus abruptly pinched-off anteriorly. The two species thus are distinct, although the range of morphologic variation of £. relnlanus has not been described. Discussioni Pollnlces (Euspira) lewisli is the largest known Recent natlcld, and nearly equally large fossil specimens are known through the Late Pliocene. One speci men from the Pico Formation fUCB 4164] has a height of 63 ram and a diameter of 61 mm, whereas the largest from the Saugus Formation [UCB 7090] is 102 mm X 80 mm, from the Purlsima Formation is 90 mm X 84 mm CUCB 1779], and from the upper Pleistocene of BahCa San Quintin is 111 mm X 97 mm. The probable ancestor of £. (£.) lewisli was the Miocene and Pliocene species £. (£.) aallanol Dali, 1909, of which the largest known specimen has a height of 57.4 mm and a diameter of 53.3 mm (broken). Bernard (1968) showed that sexual dimorphism in £. (£.) lewisli is expressed as a difference in total weight to shell weight ratios. For males, this mean ratio is 109 0.472, whereas it is 0.401 for females, based on a sample of 1875 specimens. Probable sexual dimorphism Is also ex pressed In shell differences. There are two easily separable forms of £. (£.) lewlsll. one with a more globose, thinner shell, a narrower umbilical callus and a more open umbilicus, and the other with a more elongate, thicker shell, a broader umbilical callus, and narrower umbilicus. The latter form also has a strong concave flexure at the shoulder, whereas that of the first form is never as well developed. The possible correlation of these shell differences with primary sex characters was not investigated In my study, but it is an obvious topic for future work. Pollnlces (Euspira) lewlsll and £. (£.) draconls are similar species, as noted in the discussion of the latter species, but £. (£.) lewlsll Is distinguished by Its higher spire, non-tabulate and less Inflated whorls, and thicker shell. Pollnlces (Euspira) draconls apparently evolved as a middle Pliocene offshoot of £. (£.) lewlsll. and the two species may still be found together, although £. (£.) draconls is generally more of an offshore form. As noted above, the type locality originally stated for Natica alaIda Gould, 1848, of Rio Negro, Argentina, is probably in error. In a citation for f j . alaIda in 1852, Gould gave a locality of Classet, Oregon, and apparently never again referred to a la Ida as a South American 110 species. Dali (1919*352) noted that U. S. Exploring Ex pedition material, from which Gould described £. aIkIda, often has incorrect locality data. Pollnlces (Eusolra) draconls (Dall, 1903) Plate 1, figures 18, 19 Lunatla draconls Dall. 1903*174-175. Pollnlces (Euspira) draconls (Dall), Dall, 1921*165, pi. 14Tfigs. 4, 6* Oldroyd, 1927*730-731, pi. 99, figs. 3, 6i Burch, 1946*30. Pollnlces draconls (Dall), Packard, 1918*324-325, pi. 38, figs. 2a,bt Oldroyd, 1924*163, pi. 36, figs. 2a,b» Keen, 1937*44* Smith & Gordon, 1948*198* MacGlnltle & MacGlnltle, 1949*330, 375, figs. 170, 232i Rodda, 1957*2483i Kanakoff & Emerson, 1959*30* McLean, 1969* 37, fig. 19.2. Description* Color.--Shell exterior white to buff. Perlostracum thin, light to medium yellowish brown, becoming thicker, darker and scaly within umbilicus. Inner lip and umbilical callus medium to chocolate browni filling of posterior apertural angle white* interior cloudy white and pale brown. Size.--Average* height 55 mm, diameter 52 mm* larg est specimen* height 78.0 mm, diameter 69.5 mm. Shell Form.--Shell globose, spire low* body whorl inflated) shoulder distinctly tabulate* shell relatively thini nuclear whorls 2%, smoothi postnuclear whorls 4%* Ill suture deeply impressed. Spiral sculpture of minute* wavy* weakly developed costellaet axial sculpture of in cremental growth lines. Parietal callus thin, moderately filling posterior apertural anglei anterior lobe distinct. Umbilicus broadly open, exposing earlier whorlst sculptur ed with irregular axial wrinkles and weak spiral costel- laej with distinct angulation along basal margin of umbilicus. Inner lip thickened, broadens posteriorly, not forming a distinct umbilical callusi basal portion greatly thickenedi funicle lacking. Operculum.— Chitinous, filling aperture. Specimens Examined! 375. Geographic Occurrence and Ecology* Redding Rock, Cali fornia (41°22*N), south to Cedros Island, western Baja California, Mexico (28°10'N). Usually found living on sandy bottoms offshore, in depths from 15 to at least 400 meters, also in bays. Stratiaraohlc Occurrencei Ranges from Middle Pliocene to Recent. Middle Pliocenei £tchegoin Formation, La Clma quadrangle, central California (UCBi identification questionable). Upper Pliocenei San Diego Formation, San Diego County, California (LACM)i Fernando Formation, central Los Angeles basin, California (LACM). Pliocene i Purlslma Formation, Ano Nuevo quadrangle, San Mateo County, 112 California (SU)i Eeel River Formation, Fort Orford, Oregon (Arnold & Hannibal, 1913)* formation unknown, Chalome area, central California (USGS). Lower Pleistocenei San Pedro Formation, Deadraan's Island, San Pedro, California (UCB)i Anchor Silt, northwestern Los Angeles basin, California (Rodda, 1957). Upper Pleistocenei Palos Verdes Sand, Newport Bay, California (Kanakoff & Emerson, 1959). Tvoe Localityi Four localities in California were men tioned by Dali (1903)t Drake's Bay, the Farollon Islands, Monterey, and Avalon, Catalina Island. The label with the holotype indicates the type locality as Monterey Bay. Type Material! Holotype, USNM 172859. Discussioni The umbilicus is sometimes partially restrict ed by the inner lip, so that not all of the earlier whorls are visible within it, which has led people to confuse this species with £. (£.) lewlall (Gould, 1847). The combination of low spire, tabulate whorls and thin shell distinguishes £. (£.) draconls from £. (£.) lewlall in al most every case. In occasional instances where there is still some doubt about identification, the presence of the shoulder flexure on £. (£.) lewlall is a dependable criter ion for separating adults of the two species. Juveniles of the two species are so similar that it is sometimes not possible to identify them with certainty. 113 The oldest known fossils, from the Middle Pliocene Etchegoin Formation* are poorly preserved* so that their Identification Is questionable, whereas the Upper Pliocene and later fossils are well preserved. Pollnlces (Eusplra) draconls thus apparently evolved In central or southern California during the Middle to Late Pliocene* and was un doubtedly an offshoot of £. (£.) lewlsll. which first ap peared In the Miocene. From Its first appearance In the fossil record, £. (£.) draconls has not been as abundant as £. (£. ) lewlsll. although Its thinner shell may have diminished Its chances for preservation in fossil as semblages . Dali (1921) reports a northern range limit of Port Althorp, southern Alaska (58°11*N), but the specimen upon which this is apparently based, USNM 207378, is a Juvenile of £• (£•) oalIldus (Broderip & Sowerby, 1829). Pollnlces (Eusplra) palIldus (Broderip & Sowerby, 1829) Plate 1, figures 20, 21, 22 Natlea pallida Broderip & Sowerby, 1829i372i Gray, 1839t 13o7 pi. 34, fig. 15* Mtddendorff, 1849i421-422i Mlddendorff, 1851i210-212i Philippi, 1851i96-97, pi. 14, fig. 2) Carpenter, 1864i523t Sowerby, 1883*92, pi. 9, fig. 137| Tryon, 1886i37, pi. 9, figs. 76-78, pi. 13, fig. 15, pi. 14, figs. 26-28. Natlea pallida forma normalla Mlddendorff, 1851i210. 114 Lunatla pallida (Broderip & Sowerby), Carpenter, 1864i661i Dali, 1874i251i Odhner, 1913*8, 31-40, pi. 3, figs. 15, 19-37, pi. 4, figs. 1-8, pi. 5, figs. 16-18* Thorson, 1951i22-23j Okutanl, 1964t393, pi. 1, fig. 19, pi. 5, fig, 8i Okutanl, 1966i16, pi, 2, fig. 6j MacPherson, 1971i58-59, pi. 3, fig. 8. Pollnlces (Eusolra) palIldus (Broderip f i t Sowerby), Dali, 19211164, pi. l4, fig, 5 Os "£■ (£•) oallIda"11 Oldroyd, 19271728, pi. 97, fig. 9 tas ’ *£.(£.) pallidaM3* Oldroyd, 1924i162, pi. 3, fig. 2 Cas "£. (£.) pallida"!! Burch, 1946*29. Eusplra pallida (Broderip f i t Sowerby), Kuroda & Habe, 1952«57* Kotaka, 19621135, pi. 33, figs. 19-20. Pollnlces pallldus (Broderip f i t Sowerby), Pllsbry, 1895* 72* Keen, 19*7i49* MacGinltie, 1959*91, pi. i2, fig. 10* Clarke, 1963*97. Eunatica pallida (Broderip f i t Sowerby), Habe f i t Ito, 1965* 30, pi, 37 fig. 3. Natlea (Lunatla) pallida (Broderip f i t Sowerby), Frlele f i t Grieg, 1901*69. Natlea aroenlandlca Moller, 1842*80, Moller I842*7i Jeifreys, lSo7*216| Jeffreys, 1869*215, pi. 78, fig. 2* Sowerby, 1883*96, pi. 9, fig. 140* Watson, i886* 447. Lunatla aroenlandica (Moller), Gould, 1870*341-342, fig. 611 * Sars/ 1878■158, pi. 21, fig. 15, pi. V, fig. 13* Carpenter in Dawson, 1872*392* MacNeil, 1957*103, pi. 12, fig. 21 O s "aff."J* Bousfield, 1960*17, pi. 2, fig. 23. Natlea (Lunatla) groenlandlca Moller, Frlele f i t Grieg, 1901765. Pollnlces aroenlandlcus aroenlandlca"!* Sm (Moller), Burch, 1946*29 £as "£• Smith f i t Gordon, 1948*198. (^yng^la) Rroenlandlea (Moller), Morch in Rink, Eusplra nonterona Dali, 1919*352-353. Pollnlces (Eusplra) aotiterona (Dali), Dali, 1921*164* Oldroyd, 1527 *727* BurcE, 1946*29. 115 Pollnlces aonteronus (Dali), Keen, 1937*44 i&s ________ >rc _ monterona"1i MacGinltie. 1959*91-92, pi. 12, fig. 9. Natlea caurlna Gould. 1847t239, Gould, 1852i212-213i Gould, 1856ipi. 15, fig. 254i Carpenter, 1857i209, 213i Gould, 1862t50, 244* Johnson, 1964i53, pi. 16, fig. 13. Pollnlces (Eusplra) caurinus (Gould), Dali, 1921i164 fas "P. (E.) caurlna"It Oldrovd. 1924*162* Oldroyd, 1927i 729i Burch,194#i29• (Gould), Keen, 1937t44i Smith & Gordon, Lunatla caurlna (Gould), Woodring, 1938i22, pi. 5, fig. 19 fas *cf, j * Keen & Bentson, 1944*168 as "cf." i Adegoke, 1969*167 Cas cf.]. Eusplra canonlca Dali, 1919*353. Pollnlces (Eusplra) canonicus (Dali), Dali, 1921*165 Cas "P. (E.) canonlca"1 * Oldroyd, 1927*727* Keen, 1937* 44* Burch, 1946*29. ? Natlea borealis Gray, 1839*136, pi. 37, fig. 2* Philippi, 1851 *109-110, pi. 15, fig. 16. Description* Color.--Shell exterior and interior white. Perio- stracum thin, ranging from pale yellowish white to olive brown and rust brown, usually paler on base. Size.--Average* height 25 mm, diameter 20 mm* larg est specimen* height 47.6 mm, diameter 41*8 ram [CAS 39460, Bristol Bay, Alaska3. Shell Form.--Shell somewhat elongate, spire moderately elevated* body whorl not greatly inflated* whorls usually slightly flattened below suture* shell thin* whorls about 6 (apex eroded). Spiral sculpture of minute, wavy, very 116 weak costellaei axial sculpture of incremental growth lines that are heavier Just below suture and on base. Parietal callus thin, usually transparent at its center, moderately filling posterior apertural anglet anterior lobe distinct. Umbilicus narrowly open in adults, with a shallow sulcus and narrow channel, may be closed in Juveniles. Umbilical callus narrow with a gentle central swelling, expanded posteriorly to bridge umbilicus and meet parietal callus, may plug umbilicusi funicle incon spicuous. Anterior inner lip and basal lip thickened. Operculum.--Chitlnous. filling aperture. Specimens Examinedi 1500. Geographic Occurrence and Ecology> Circumboreal. In the eastern Pacific, south to the U.S.-Mexican border (32°32*N)i in the western Pacific, south to Japani in the western At lantic, south to Cape Hatteras, North Carolina) In the eastern Atlantic, south to the North Sea and Ireland. Not common in the eastern Pacific south of Monterey, Cali fornia (36°36'N). Known on soft bottoms in depths from 15 to 2498 metersi reported intertldally alive at San Juan Islands, Washington [UCD, no number], but this needs con formation. Strat iaraphlc Occurrencei Ranges from Middle Miocene to Recent. Middle Miocenei Kern River area, California (CAS). 117 Middle Miocene to Pliocenei Yakataga Formation, Yakataga area, Alaska (UCfi). Middle Pliocenei Etchegoin Formation Coallnga area, California (Adegoke, 1969, as cl< ). Upper Pliocenei upper part Rio Dell Formation, Centerville, Humboldt County, California (CAS). Pliocenei Port Orford quadrangle, Oregon (UCB)f Tugidak Island, Trinity Islands, Alaska (USGS). Pliocene to Pleistocenei Tjomes sequence, Serrloes and Mactra groups, Iceland (USGS). Lower Pleistocenei Elk River Formation, Cape Blanco quadrangle (USGS) and Curry County (UCB), Oregonj Santa Barbara Formation, Santa Barbara, California (UCB). Type Localitiesi Natica pallida - Icy Cape ^Arctic coast of AlaskaJ (Broderip A Sowerby, 1829). $9 Natica aroenlandlca - Greenland (Moller, 1842). Eusplra monterona - Station 1199, Captains Bay, Un- alaska, Alaska, 75 fathoms, mud and gravel (Dali, 1919). Natica caurlna - Straits of Juan de Fuca, Washington (Gould, 1847). Eusplra canonlca - USFC station 2923, off San Diego, California, 822 fathoms, green mud (Dali, 1919). Type Materiali flfttlCt PttUUi - Unknown, presumably in BM(NH) (Dance, 1966). 118 Natica aroenlandlca - Unknown, presumably in BM(NH) or Zoological Museum, Copenhagen, Denmark (Dance, 1966), Eusolra monterona - Holotype, USNM 220856. Natica caurlna - Lectotype, MCZ 169081. Eusolra canonlca - Holotype, USNM 209411. Nomenclature! Commentaryi In his original description of Natica aroen land lea. Moller (1842) cited the name as " J J . groenlandlea &ec,N and the species was attributed to Beck, a Danish zoologist, by many subsequent authors. However, there seems to be no evidence that Beck authored this species, although he may have used aroenlandlea as a manuscript name. All adults of £. (£.) palIldus have an open umbilicus, although It Is reduced to a slit In some Individuals. Because Juveniles have broader umbilical calll than adults, their umbilici are generally narrower, although usually still open. However, about a fourth of Recent Juveniles have a closed umbilicus, and a Recent population will show all intergradations from closed to open umbilici. In Juveniles with closed or nearly closed umbilici, broadening of the umbilical callus is accompanied by thickening of the anterior Inner lip and a thicker filling of parietal callus in the posterior apertural angle. The anterior Inner lip Is often thicker and more elevated than 119 the umbilical callus in both Juveniles and adults. Fossils of £. X£*) oallldua are much smaller than Re cent specimens and do not have completely closed umbilici. The largest fossil specimen is 21.3 mm in height and 17,7 mm in diameter £UCB b-7380. Elk River Formation, Cape Blanco, Oregon, Lover PleistoceneJ and is only slightly larger than average for a Pliocene or Pleistocene specimen. Because no Upper Pleistocene fossil £. palIldus are known in the eastern Pacific, it is not known whether the present large size of specimens is a Late Pleistocene or Recent development. Closure of the umbilicus by the umbilical callus, seen in some Recent Juveniles, is not known among fossils. Thickness of the umbilical callus and its closure of the umbilicus varies among fossils, but to a lesser degree than among modem specimens. Fossil speci mens also tend to have thicker parietal calll and higher spires than Recent specimens. The origin of £. (£.) oallldua in the fossil record is not known, and there is no likely eastern Pacific Neogene species from which it might have evolved. However, £. rectus Tegland, 1933, from the Ollgocene Blakeley Forma tion of Washington, Is similar to fossil £. (£.) palIldus in size and morphology and may prove to be its ancestor when Paleogene natlcids become better known. The lack of this species in upper Pleistocene de posits may be due to circumstances of preservation. Al- 120 most all upper Pleistocene marine deposits In western North America represent shallow-water, rocky-shore environ ments, in which species such as £. (£.) palIldus are un likely to have lived. Lower Pleistocene and older de posits of this region preserve a greater variety of marine habitats, including those in which £. (£.) palIldus lived. The occurrence of £. (£.) oalIldus in CAS collections from the Kern River Miocene deposits of southern Cali fornia needs confirmation, because Addlcott (1970) does not Include this species in his comprehensive report on the gastropods of those deposits. PQllnices (Eusplra) vlctorlanua Clark & Arnold, 1923 Plate 1, figures 23, 24 Folinices (Eusplra) vlctorlana Clark & Arnold, 1923t170, pi. 33, figs. la,b, 5a,bi Durham, 1944i160. Polin^g| (^o^rjices) ^|ctgrlana^Clark ^Arnold, Weaver, Pollnices^vi^torlan^ Clark^fii Arnold, Addlcott, 1970*67, Pollnlces canalIs Moore, 1963128, pi. 2, figs. 18, 221 Addlcott, 1970 *67 fas Junior synonym of £. vlctorl anua 7 i Moore, 197li23, pi. 7, figs. 1, 2. Natica (Natica) saxea Conrad, Etherington, 1931ipl. 12, figs. 2. 3. 7. 14 Tnot N. saxea Conrad, 18491. Natica (Tectomatlca) ««** Conrad, Weaver, 1942ipl. 68, fig. 4 Cnot saxea Conrad, 1849J. X21 Descriptioni Size.--Averagei height 33 nun, diameter 28 ram) larg est sped men i height 38.1 mm, diameter 36.3 ram £UCB 2713, Temblor Formation, Callente quadrangle, California}. Shell Form.--Shell globose to somewhat elongate, spire moderately to strongly elevatedi body whorl not greatly inflated) shoulder flattened near suture, may be slightly concavei shell thickness averagei whorls suture very slightly Impressed. Shell smooth except for fine incremental growth lines. Parietal callus thick, heavily filling posterior apertural angle) anterior lobe weak, with a very shallowly dimpled surface. Umbilicus narrowly open, may be slit-like, never completely closed) shallow, poorly defined sulcus may be present. Umbilical callus relatively narrow, gently tapers anteriorly) may have central swelling set off above by sulcus and below by indentation on umbilical side of inner lip, or swelling may be absent, with umbilical margin of callus straight) funicle low, broad, if present. Anterior inner lip and basal lip greatly thickened. Operculum.--Unknown. presumably chitinous. Specimens Examined! 450. Stratigraphlc Occurrence! Ranges from Lower to Middle Miocene. Lower Miocene (?)i Sooke Formation, Vancouver Island, British Columbia (Clark & Arnold, 1923). Lower 122 Miocenei basal Jewett Sand* Kern River areat California (Addlcotti 1970i USGS). Middle Miocenei middle and upper Olcese Sand and Round Mountain Silt* Kern River area, Cali fornia (Addlcott, 1970i USGSj UCB)* Temblor Formation, Reef Ridge, CoalInga district, California (SU)i Astoria Formation, Skamokawa quadrangle, southwestern Washington (Addlcott, 1970), Montesano quadrangle, Washington (USGS), and coastal Oregon (Moore, 1963i CASj UCB). lyre LrealUlgq« Pollnlces (Eusolra) vlctorlana - CAS 231, in sea cliffs east of the mouth of Kirby Creek, 6 miles west of Sooke, Vancouver Island, British Columbia. Sooke Formation, lower Miocene (7) (Addlcott, 1970). Polinioes canalis - USGS 18806, In beach cliffs form ing first headland south of fill at Spencer Creek, Lincoln County, Oregon. Astoria Formation, middle Miocene (Moore, 1963* Addlcott, 1970). Type Materiali Pollnlces (Eusolra) vlctorlana - Holotype, CAS 582. Pollnlces canalis - Holotype, USNM 563134 [not 561534 or 56134, as given by earlier workers]. Discussioni As noted by Addlcott (1970), the specimens with a swollen umbilical callus are smaller than those with a simple callus. Although these forms intergrade to 123 some extenti they occur In approximately equal numbers and may represent sexual dimorphism. Umbilical closure is a function of callus development, so specimens with a central callus swelling have narrower umbilici. Pollnlces (Eusolra) vlctorlanus seems most closely related to £. (£.) pallldus (Broderip & Sowerby, 1829), which occurs from Miocene to Recent, although the two species are easily told apart. Fossil specimens of £. (£.) pallldus are smaller than those of £. (£.) vlctorlanus. al though Recent specimens are larger. Pollnlces (£.) pal- 11 dua has more strongly Impressed sutures, so that the whorls stand out individually inproflle, whereas the whorls of £. (£.) vlctorlanus form nearly a single slope from apex to periphery. Some individuals of £. (£•) pal lldus have a closed umbilicus, which never occurs In £. (£.) vlctorlanus. but the umbilical callus does not characteristically taper anteriorly. In £. (£.) pallidus the parietal callus distinctly overhangs the umbilicus, and the parietal callus is much thinner than in £. (£.) vlctorlanus. The Late Miocene species £. (£.) dlabloensls Clark, 1915, differs from £. (£,) vlctorlanus by being larger (up to 51 mm in height) and more elevated, having a more strongly Impressed suture, a thinner parietal callus, and a thinner inner lip and umbilical callus. Pollnlces (Eusplra) vlctorlanus Is abundant In the 124 middle Miocene deposits of the Kern River area and rela tively uncommon elsewhere. It Is present In Oregon and Washington, but unknown In the extensively collected California Miocene deposits south of the San Joaquin basin. The record of this species from Upper Ollgocene beds of Kings County, Washington (Durham, 1944), Is questionable. The hypotype In the UCB collection Is not recognizable except as a natlcld. Pollnlces (Eusolra) aauJanus Dali, 1908 Plate 1, figure 25 Pollnlces (Eusplra) aaulanus Dali, 1908i334, pi. 9, fig. Tt Dali, 1909123oi M. Smith, 1944112, fig. 127. Pollnlces (g^llnlces £.1. ) aaulanus Dali, Keen, 197li480, Descriptioni Color.--Shell white* Interior and callus white. Perlstracum thin, olive brown. Size.— Holotypet height 26.0 mm, diameter 24.0 mm. Shell Form.--Shell globose, spire low* body whorl moderately inflated\ whorls slightly compressed laterally, weakly and narrowly shoulderedi shell thlnj whorls about 5 Ctype wom]t suture slightly impressed, minutely chan neled. Spiral sculpture of minute, weakly developed, ob scure llneatlonst axial sculpture of incremental growth lines. Parietal callus thin, lightly filling posterior 125 apertural anglei anterior lobe weak. Umbilicus open* relatively small. Umbilical callus broken off of holo type, was apparently very slender. Basal lip thickened. Operculum.--Unknown. presumably chltlnous. Specimens Examinedi Holotype. Geographic Occurrence and Ecologyi Known with certainty only from off of Punta Aguja, northern Peru (type locality), also reported as "probably" £, (£.) aguianus from the Gulf of Panama (Dali, 1908). Type Localityi U.S.S. "Albatross" station 4653, 17 miles N61W from Punta Aguja, northern Peru (6°S), on a mud bottom at 981 meters depth, bottom temperature 5.2°C (Dali, 1908). Type Materlalt Holotype, USNM 110566. Discussioni In addition to the holotype, Dali (1908) referred to a second specimen Identified "probably" as this species, from the Gulf of Panama In 3058 meters depth, which was supposedly dead and worn. I did not locate this specimen In the USNM collections* 126 Pollnlces (Eusolra) crawfordlanus Dali, 1908 Plate I. figure 26 Pollnlces (Eusplra) crawfordlanus Dali, 1908*335-336, pi. 11, fig. 7i M. Smith, 1944*12, fig. 138. Pollnl^s (ggllnlces^g.l.) crawfordlanus Dali, Keen, Description * Color.--Shell whitish. Perlostracum thin, pale yellowish brown* Size.--Ranges from height 6.5 mm, diameter 5.8 mm USNM 123051 to height 15.0 mm, diameter 13.5 mm 0JSNM 110650J, based on three specimens. Shell Form. — Shell elongate, spire moderately elevated* whorls evenly rounded* shell thin* whorls 4 or more apex eroded * suture moderately Impressed. Whorls smooth, with only Incremental axial growth lines. Parietal callus thin, lightly filling posterior apertural angle* anterior lobe weak. Umbilicus open, slit-like, Inconspicuous. Umbilical callus not distinct from Inner lip, slightly conceals part of umbilicus. Inner lip and basal lip thickened* outer lip smooth. Operculum.--ChitInous. filling aperture. Specimens Examined* Holotype and two paratypes• Geographic Occurrence and Ecology* Mazatlan, Mexico 127 (23°N), Co Punta Aguja, Peru (5°40'S), and the Galapagos Islands. In depths of 996 to 1895 meters, from substrates of mud and "ooze," and with prevailing bottom-water temperatures of 37°F to 40.1°F (2.8°C to 5°C), Tvoe Localityi Gulf of Panama, 996 meters depth, U.S.S. "Albatross" station 3356 (Dali, 1908). Type Materiali Holotype, USNM 123044. Pi acubaIoni Dali mentions five specimens in his descrip tion of £. (£.) crawfordlanus. of which four are consider ed here to be correctly assigned. The fifth specimen is from off southern Chile at about 50°S latitude and is 32 mm in height and 28 mm in diameter. It is characterized by a greatly thickened inner lip, slightly swollen umbilical callus, and Imperforate umbilicus. I did not locate this specimen in the USNM collection, but I did find a similar specimen, USNM 102586, labeled "Pollnlces (Eusplra) crawfordlanus Dali. Cotype. Patagonia. Dr. Crawford." This specimen is 30.8 mm in height and 28*6 mm in diameter and was apparently considered by Dali when he proposed the species, although he did not mention it. This specimen and the one noted by Dali (1908) from off southern Chile are considered to be distinct from £. (£.) crawfordlanus and are not assignable to any species within my study area. 128 Pollnlces (Eusplra) lltorlnua Dali, 1908 Plate 1, figure 27 Pollnlces (Eusolra) lltorlnua Dali, 1908i337. Pollnlces lltorlnua Dali, Keen, 1971*480, Descriptioni Color*--Shell white. Perlostracum thin, pale yellow ish white. Size.--Height 9 on, diameter 8.5 mm (holotype). Shell Form.--Shell globose, spire lowi shoulder flattened, whorls evenly roundedi shell thickness averagei whorls about 3%, apex eroded) suture very slightly Im pressed. Spiral sculpture of Indistinct, minute strla- tionsi axial sculpture of minute Incremental growth linest shell smooth. Parietal callus of average thickness, moderately filling posterior apertural anglet anterior lobe weak. Umbilicus narrowly open, slit-like. Umbilical callus an indistinct swelling. Inner lip and basal lip slightly thickened. £Q£ESUlmB*~~Chltinous, filling aperture. Geographic Occurrence and Ecologyi Known only from type locality, In 1485 meters depth on a bottom of "ooze,” with temperature of 38,4°F (3.5°C). 129 / Type Localityi Near Galapagos Islands, U.S.S. "Albatross” station 2807, 1845 meters (Dali, 1908). Type Materiali Holotype, USNM 96481. Discussioni Because it is known from a single specimen, this species is difficult to evaluate as a member of a family whose species are often very similar and show wide lntraapeclfic variation. If it occurred in shallow water it might be mistaken for a Juvenile Pollnlces (Pollnlces) intemeratus (Philippi), fgUalygg (fiugplra) pardoanus Dali, 1908 Plate 1, figure 28 Pollnlces (Eusolra) pardoanus Dali, 1908i336. Pollnlces CPollnlces $.1.) pardoanus Dali, Keen, 1971i480, fig. 887. Descriptioni Color.--Shell white. Periostracum thin, pale yellow ish brown. Size.--Ranges from height 5.9 mm, diameter 7.0 mm ClJSNM 123050} to height 13 mm (broken), diameter 14 mm (broken) [USNM 123046}, based on three specimens. Shell Form.— Shell globose, spire low to moderately elevatedi whorls evenly roundedi shell thickness average% whorls about 4| suture moderately impressed. Spiral 130 sculpture of occasional, indistinct, minute striatlonsi axial sculpture of incremental growth lines. Parietal callus thin, moderately filling posterior apertural anglei anterior lobe weak, not overhanging umbilicus. Umbilicus open, narrow. Umbilical callus a simple thickening of inner lip, partly reflected over umbilicus. Inner and basal lip thickened. Operculum.--Chitinous. filling aperture. Specimens Examinedi Holotype and two paratypes. Geokraphic Occurrence and EcqIorvi Gulf of Panama and near the Galapagos Islands. In depths of 1915 meters and 2690 meters in the Gulf of Panama and 1620 meters near the Galapagos, on NoozeN substrates, with water temperatures between 36.2°F and 37°F (2.3°C to 2.9°C). Type Localityi Gulf of Panama, 2690 meters depth, U.S.S. HAlbatrossN station 3361 (Dalit 1908). Type Materiali Holotype, USNM 123046. Discussion i This species is known only from the three specimens described by Dali. Its shape is like that of some Juvenile £. (£.) pallldus (Broderip & Sowerby), but with a lower spire and more open umbilicus. 131 Pollnlces sookenais Clark & Arnold, 1923 Plate 1« figure 29 Pollnlces (Ammiii^na?^ sookensla Clark & Arnold, 1923i170, pi, 33, figs. 4a,b. Amnul\Ina sookensla (Clark & Arnold), Weaver, 1942i347, pi. 7, figs. 3, 6. Description (from Clark. & Arnold, 1923) i Shell medium in size, suborbicular* whorls five in number* spire about one*third height of shell. Suture slightly appressed. Whorls gently convex, greatest con vexity near base of whorl. Aperture broadly subovate. Inner lip reflexed anteriorly and nearly covering the closed to subperforate umbilicus* posteriorly there is a thin wash of callus on body whorl. Dimensions of typei Height, 15 mm* greatest width of body whorl, 12 mm* height of body whorl, 10.5 mm. Specimens Examined* "Holotype" only. Stratlaraphic Occurrencei Known only from type locality. Type Localityi SU NP129, sea cliffs between mouths of Muir and Kirby creeks, west of Otter Point, Sooke, Van couver Island, British Columbia, Canada. Sooke Formation, Lower Miocene (7) (Clark & Arnold, 1923). 132 Type Materiali Unknown* apparently lost* See discussion below. Discussioni The labeled holotype of £. sookensla in the Stanford University collection is not the specimen figured as the type by Clark & Arnold (1923) or by Weaver (1942). The Stanford specimen has the same lot number (30205) and locality number (NP 129) inked on It as were given for the holotypei the number 245 has been more recently inked on the Stanford specimen. The present Stanford specimen Is nearly the same size as the figured holotype, with a height of 14.4 mm, diameter of 12.5 mm, and body whorl height of 10.0 mm, but It differs from the figured holotype mainly by having an open umbilicus. The umbilicus of the figured type is des cribed as "closed to subperforate" (Clark & Arnold, 1923i 347) and is clearly shown as closed and represented by a shallow depression. This is unquestionably different from the narrow but clearly open umbilicus of the Stanford specimen. In addition, the figured holotype has more acutely tabulate whorls, the sutural area of its body whorl is not as badly decorticated, and it bears a dif ferent pattern of minor fractures and breaks than the Stanford specimen. Thus, the Stanford specimen is clearly not the holotype of Clark & Arnold (1923), which appears to be lost. 1 examined the Sooke Formation collection made 133 by Clark and Arnold at Stanford* but no specimens similar to their figured holotype was found. 1 have done no preparation work on the Stanford specimen. The Stanford specimen Is most similar to Natica (Natica) clarkl Etherington, 1931* a rare species from the Middle Miocene Astoria Formation of Oregon and Washington. Natica (Natica) clarkl Is known from only 4 specimens• so its range of morphologic variation is not known. The Stanford specimen has a higher spire than any of the (££.) clarkl specimens* but is otherwise identical. If this identification is correct* it is the first record of £ ! • (£.) clarkl in Lower Miocene (?) deposits. PgUnl99S 9lYTU?UH Reagan* 1909* nomen UybUtt Pollnlces (Lunatla?) olvmpldil Reagan* 1909i171, 194, pi. 3, fig. 291 Reagan, 19101648 Cas "£. (L.?) olympldea Arnold & Hannibal* 1913* sp. list facing p. 566, 584. Natica olvmpldil (Reagan), Weaver, 1916*172, 176. This species is considered to be a nomen dubium be cause it cannot be identified from its original descrip tion and figure and there are no unaccounted for natlcid species in its type area. The location of the type speci men is unknown. The type locality of £. olvmpldil is Gettysburg* Washington* in beds of the middle Miocene Clallam Forma tion* and the species has been reported in the Astoria 134 Formation of western Washington (Arnold & Hannibal, 1913* Weaver, 1916). There are several Middle Miocene natlclds In Washington that could be Identical with £. olvmpldil. but Reagan's (1909) crude drawing of the type specimen does not allow comparison with any of them. Subgenus Hvoterlta Woodrlng, 1957 Hvpterita Woodrlng, 1957*92. Tvne-at>eclea * Natica hellcoldes Gray, 1825, by original designation. Miocene to Recent* eastern Pacific. Figured herein. niftttnoaiq * Shell thin, greatly depressed. Umbilicus very broadly open, with gently sloping wall. Umbilical callus a thin lobe suspended anteriorly from the pillar-like funicle. Parietal callus very thin. Discussion* Hypterlta Is characterized by Its flattened shell and peculiar umbilical callus. The type-species Is the only known living representative of this subgenus * with a Caribbean Miocene species also placed here by Woodrlng (1957), 135 PoUnices (Hypterlta) hellcoldea (Gray, 1825) Plate 1, figures 30, 31 Natica hellcoldes Gray. 1825i5ll Tnew name for N. patula G, B. Sowerby, I, 1824, not J. Sowerby, 18123. Pollni^gs^gltcoldes (Gray), Hertlein & Strong, 1955* Pollnlces (Pollnlces) hellcoldes (Gray), Keen, 1958t322, fig. 268i Keen? 197174/o7 fig* 876. Never^a^CHvp^erlga) hellc^de^(Grav)■ Woodrlng, 19571 not Natica hellcoldea Johnston, 1835i69 C* Amauropsls is* landlea (Gmelln. 1788), Recent, northern oceans!• Natica patula G, B. Sowerby, I, 1824i60, pi. 5, fig. 6| Carpenter, 1864i522. not Natica patula J. Sowerby, 1822, vol. 4i99 C“ fossil, England?. not Natica patula Wood, 1828145 C* Nerita~ ) . not Natica patula Deshayes, 1832i16 C“ "Ampullarla." fossllj. Natica Rlauca Lesson, 1830*369, pi. 11, fig. 1. Natica alauca "Humboldt" Calc]I Orblgny, 1840t403i Menke, 1851i165* Philippi, 1852*42, pi. 6, fig. 3 (plate 1849)i Troschel, 1852i161i Reeve, 1855ipl. 2, figs. 5a,bi G. B. Sowerby, 11, 1883i76, pi* 1, fig* 5| Tryon, 1886i34, pi. 11, figs. 97-98. Pollnlces (Neverita) alauca ("Humboldt"), Dali, 1909t236. Polln^jgs a^guca ("Humboldt"), M. Smith, 1944i12, figs. Natica bonylandi Valenciennes, 1832*264-265, pi. 57, figs. 3a,b [new name for N. patula G. B. Sowerby, 1, 1824, not J. Sowerby, I82zji Carpenter, 1864i521. 136 Baasrtetlon* Color.--Nuclear whorls light browni early adult whorls grey-brown with prominent white band at suturei later adult whorls pinkish gray or brown, usually lighter near the suture, with a diffuse white band below peripheryj callus and Inner lip dark orange-browni Interior chestnut brown, with white band at suture and below periphery. Slze.--Averaaei height 27 mm, diameter 50 mmi larg est specimeni height 34.4 mm, diameter 62.2 mm CSU 1377, Acapulco, MexicoJ. Shell Form.--Shell low, spire depressed) shoulder tabulate, and tabulate surface may be slightly concavei base flattened) shell thin) nuclear whorls 1^, smooth, postnuclear whorls 3*fi suture moderately Impressed. Spiral sculpture of microscopic, closely spaced strlatlonsi axial sculpture of incremental growth lines that strongly disrupt the spiral striations and are heaviest near the suture. Parietal callus very thin, transparent, lightly filling posterior apertural anglet anterior lobe weak. Umbilicus very widely open, with gently sloping walls. Umbilical callus a thin lobe concealing posterior part of umbilicus, suspended anteriorly from the pillar-like funicle, tapering anteriorly) funicle strongly developed, nearly circular In section. Inner lip and basal lip not thickened) outer lip smooth. Operculum.--ChitInous. filling aperture. 137 Specimens Examined i 81. Geographic Occurrence and Ecology> Magdalena Bay, western Baja California, Mexico (24°30'N) to Faita, Peru (5°S). Most common throughout the Gulf of California, rare south of Acapulco, Mexico (17°N). Sparse habitat data indicate a depth range of 9 to 46 meters, and Keen (1971) suggests Intertidal occurrences. Earlier workers beginning with Tryon (1886) extend the southern range limit to Callao, Peru (12°S), but it Is unlikely that this tropical species ranges below Paita into the cool waters of the Peruvian zoological province. Srratigraphlc Occurrencei Ranges from Middle Miocene to Recent. Middle Miocenei lower and middle parts of Gatun Formation, Panama (Woodrlng, 1957)» 7Banana River area, eastern Costa Rica (Woodrlng, 1957). Pliocenei western Costa Rica (Woodrlng, 1957). Pleistocenei Ecuador (Hert- lein & Strong, 1955). lYfig Lw?HtUg« Natica helicoidea - Unknown (Sowerby, 1824, for £ 1 . MtUUO* Natica patula - Unknown (Sowerby, 1824). Natica alauca - Colan, near Faita, Peru (Lesson, 1830). 138 Natica bonolandi - Acapulco, Mexico (Valenciennes, 1832). Type Materiali Natica helicoldes - None designated (Gray, 1825). Natica patula - Unknown, probably in BM(NH) (Dance, 1966). Natica ftlauca - Unknown, probably in Museum d' Histolre Naturelie, Laboratoire de Malacologle, Paris (Dance, 1966). Natica bonolandi - As above. Nomenclatural Commentary t The earliest proposed name for this species, Natica patula G. B* Sowerby, I, 1824, was preoccupied by £. patula J. Sowerby, 1822. Gray proposed the replacement name £. helicoldes In 1825 and Valencien nes proposed N* bonolandi in 1832. Gray cited £ j . heli coldes as "Mr. Barnes's MS name" (Barnes, 1824), but It is still available as of Gray's (1825) mention of it. Discussloni Because of its shell form and umbilical features, this is the most distinct of all the eastern Pacific naticlds, although its rarity in older collections and Its wide geographic distribution caused it to be named more than once. It is related to £. (jj.) nereidla Maury, 1917(Miocene, Dominican Republic), which differs by being smaller (diameter 35 mm) and having a broader funlcle* 139 Natica eurvontphala Philippi, 1887 (Tertiary, Chile) was considered somewhat similar to £. (Jj.) helicoldes by Hert- leln & Strong (1955), but Philippifs original description is not diagnostic, making comparisons with other species difficult. The record of £. ( ]j, ) helicoldes from the Miocene of Costa Rica Is based on Woodrlng*s (1957) doubtful identification of a fragment earlier identified by Olsson (1942) as £. (iJ.) nereldls. Subgenus Mammilla Schumacher, 1817 Mammilla Schumacher, 1817i58. Type-speciesi Mammilla fasciata Schumacher, 1817 (- Albula mammata Roding, 1798), by monotypy. Recent, western Pacific. Diagnosisi Shell small to medium in size, elongate pyri- forra, thini suture slightly Impressed. Umbilical callus slender, elongate. Parietal callus thin. Discussioni This subgenus is characterized by Its elong ate pyrlform shape and thin shell. Mammilla is a distinct ly tropical group, confined largely to the Indo-Pacific region, with only a single species In the eastern Pacific. 140 Polinlces Q jafflfflU ls) caprae (Philippi, 1852) Plate 1, figure 32 Natica caprae Philippi, 1852i56, pi, 9, fig. 2 (plate 1850)7 Follnlces caprae (Philippi), Hertlein & Strong, 1955i139i Shasky & Campbell, 1964i115. PqU nices (Polinlces) caprae (Philippi), Keen, 1958i322, fig. 267j Keen, 19717378, fig. 874. Pollnlcefl nrirkmavt Palmer & Hertlein, 1936*77-78, pi. 19, ?igs. 15, 14. Pol inices (Magna lja) clarki M. Smith, 1950*60-61, pi. 4, figs. 7, 7a. Description* Color.--Shell pale greyish white to tan, with a wide band of mottled dark brown on the shoulder and a narrower band on the base* color bands may be poorly developed, reduced to series of Irregular blotches* entire adult shell may be covered by series of closely spaced spiral lines consisting of minute orange-brown dots* umbilicus and anterior half of callus dark brownt interior light to dark brown, reflecting exterior color pattern. Nuclear whorls buff, Perlostracum thin, pale yellowish white. Size.--Average* height 25 mm, diameter 22 mm* larg est specimen* height 32.1 mm, diameter 25.5 mm fMCZ, Pearl Islands, Panama]. Shell Form.--Shell elongate, spire moderately 141 elevated} body whorl greatly inflated anteriorly% shoulder steeply sloping) shell thinj nuclear whorls 2%, smooth) postnuclear whorls 3) suture very slightly im pressed. Spiral sculpture of minute* poorly defined, wavy costellae) axial sculpture of incremental growth lines. Parietal callus thin, moderately filling posterior apertural angle) anterior lobe weak. Umbilicus open, elongate, with broad channel. Umbilical callus narrow, occupying posterior half of inner lip) broadest near anterior end, where an abrupt swelling is dimpled by a short transverse groove) funlcle broad, with a more abrupt posterior face. Anterior inner lip thickened, basal lip not thickened. Operculum.--Not observed, presumably chitlnous. Specimens Examinedi 59. Geographic Occurrence and Ecologyi Cabo San Lucas, southernmost Baja California, Mexico (22°45*N), through out the southern end of the Gulf of California, and south to Paita, Peru (5°S), and the Galapagos Islands, Ecuador. Based on very limited collecting data, it is found in depths of 45 to 65 meters, and Keen (1971) reports it intertldally. Stratlaraohic Occurrencei Reported in Pleistocene de posits along the "coast of Oaxaca," southern Mexico (Palmer & Hertlein, 1936). 142 Type Localities t Natica caprae - Mazatlan, Mexico (Philippi, 1852). Polinlces crlckmavl - CAS locality 1299, coast of Oaxaca, southern Mexico, Pleistocene (Palmer & Hertlein, 1936). Polinicea clarki - Panama (Smith, 1950). Type Materiali Natica caprae - Unknown, presumably in BM(NH) or Museo Nacional de Historla Natural, Santiago, Chile (Dance, 1966). Polinlces crlckmavl - Holotype, CAS 5615. Polinlces clarki - Holotype, ANSF 189043. Discussioni The anteriorly Inflated whorls of £, ($.) caorae distinguish it from other eastern Pacific natlclds, to which it is not closely related. However, £. (£1.) caorae is close to several Indo-Paclfic species of similar form, including Polinlces (Mammilla) maurus (Lamarck, 1816), £. Of.) melanoatomus (Gmelin, 1791), and £. (if.) melanostomoides (Quoy & Gaimard, 1833), among others (figured in Cemohorsky, 1971). This general shell shape seems to be confined to Indo-Pacific natlclds, so that £. (If.) caorae Is apparently a Pleistocene migrant from the western Pacific. No similar species Is known as a fossil 143 in the eastern Pacific. Genus Neverita Rlsso, 1826 NeverIta Rlsso, 1826*149. Type-speciesi Neverita losephina Rlsso, 1826, by mono- typy. Eocene to Recent, Europe. Figured in Wenz (1941i 1031, fig. 2952). Diagnosisi Shell small to large, globose to ovate» whorls not greatly inflated, smooth except for incremental growth lines and microscopic spiral costellaei suture slightly to moderately Impressedi shell thickness average or greater. Umbilicus closed, except for a few species or variations within a given species. Umbilical callus broad and mas sive, either entire or divided into two lobes by a trans verse groove. Parietal callus moderately thick to heavy. Operculum chitlnous, entirely filling aperture. Discussioni This genus is characterized by its closed or nearly closed umbilicus and generally massive umbilical callus. Neverita is considered to be of full generic rank because the degree of closure of the umbilicus, which is of major importance in a subfamily composed of otherwise generally similar and nearly featureless species. Umbilical closure is thus deemed more significant in the Pollnlclnae than in the Natlcinae, where there are more morphologic features to aid in evaluating generic taxa. This genus Is worldwide In distribution) and as de fined here It ranges from the Arctic to the tropics In the eastern Pacific. Subgenus Neverita Rlsso, 1826* sensu stricto Diagnosisi Umbilical callus entirely covering umbilicus, not divided into two lobes by transverse groove. Discussioni This subgenus Is characterized by Its massive smooth umbilical callus. It occurs living in temperate and Arctic waters of the eastern Pacific and elsewhere* and is found in Upper Miocene deposits of western America. Neverita (Neverita) nana (MSller, 1842) Plate 1, figure 33 Natica nana Moller, 1842i80 [p. 7 of separate]! Philippi, 1851*108, pi. 15, fig. 13. Lunatia nana (Moller), Sars, 1878*159, pi. 21, figs. 16a, b7 plTV, fig. 14i Verrill, 1882*516, pi. 42, fig. 9» Odhner, 1913*8, 30, pi. 4, figs. 22-25) Okutani, 1964 394, pi. 1, fig. 20, pi, 5, fig. 6* Okutani, 1966* 16, pi. 2, fig. 7 Os L* nana "Muller"]. Lunatia (Mamma) nana (Moller), Dali, 1885*182. Mamma nana (Moller), Dali, 1878*12. Natica (Limatla) nana (Moller), Frlele & Grieg, 1901*69. faltnlffM (JEttUnlfiSa) (toller), Dali, 1921*165) Oldroyd, 19Z7*732) Keen, 1937*44) Burch, 1944*32. 145 not Natica nana Tentson-Woods, 1875i149 [Recent, Tasmania], Descriptioni Color.--Shell off-white, with a brighter white sub- sutural band. Periostracura very thin, inconspicuous, pale yellowish white or yellowish brown. Size.--Averagei height 9 ram, diameter 7 mmj largest speciment height 17.3 mm, diameter 14.8 mm [USNM 267170, Santa Barbara, California]. Shell Form.--Shell elongate, spire low to moderately elevatedj shoulder flattened, very slightly concavei average thicknessi nuclear whorls 1^, smooth except for minute radial grooves extending short way from the suturet postnuclear whorls 3i suture very slightly im pressed. Spiral sculpture of obscure microscopic stria- tions. Axial sculpture of minute incremental growth lines. Parietal callus thin at its center but thickens toward posterior apertural angle and umbilicusi anterior lobe well-developed, merges smoothly with umbilical callus. Umbilicus closed. Umbilical callus broad, semi circular, flat, depressed anteriorly, with its margin smoothly blending Into underlying whorl* Inner lip and basal lip thickened. Operculum.— Chltinous, filling aperture. Specimens Examinedi 477. 146 Geographic Occurrence and Ecologyi From Point Belcher, Arctic coast of Alaska (70o4d'N, 159°39*W) and Aika Island in the Aleutians, south to Cabo San Quintin, western Baja California, Mexico (30°20'N), and from Labrador south to Newport, Rhode Island (41°25'N), and Georges Bank (41° 23 *N, 68°46*W). Also known on Greenland, Spitzbergen (77°N), and in the Minch area of Scotland (58°N), Vadso, Norway (70°08*N) and in the White Sea, Russia. In Japan, at Sagaml Bay and nearby Islands (Okutani, 1964, 1966). Odhner (1913) reported specimens from Iceland and south in the western Atlantic to Cape Hatteras, North Carolina (35°N). The distribution is discontinuous, so that there are no known specimens from Arctic Canada (Macpherson, 1971), none from between the southeastern Alaska Peninsula and Monterey, California (36°39'N), and very few from the Aleutian.Islands. Most specimens I have seen are from Alaska, The total depth range for this species is 11 to 1710 meters, but Its presence at a given locality is a function of water temperature, so it lives at progressive ly greater depths southward. Depth occurrences in the eastern Pacific are as follows■ Point Belcher, Arctic Alaska, 16 metersi Bering Sea and Bristol Bay, Alaska, 43 to 91 metersi western Aleutian Islands, 27 to 30 metersi Monterey, California, 1065 metersi southern California islands, 976 to 1281 metersi San Diego, California, 658 to 1506 metersi San Quintin, Baja California, Mexico, 658 147 meters. Okutani (1964, 1966) gives depths of 620 to 1350 meters for Sagami Bay, Japan. Eastern Pacific specimens from several localities between Alaska and San Diego were collected at water temperatures of 39°F to 41.8°F (3.9°C to 5.4°C), in sand and mud substrates. Type Localityi Greenland (Moller, 1842). Tvoe Materiali Probably in the Zoological Museum, Copen hagen, Denmark, because Odhner (1913) examined the type> may be in BM(NH) (Dance, 1966). Nomenclatural Commentaryi It is remarkable for such a widely distributed species, especially a natlcid, to be free of synonymous names, but this seems to be the case. Tryon (1886) considered ft. (ft.) nana a Junior synonym of Natica limnaculata Totten, 1835, but the latter species has an open umbilicus. Morch (in Rink, 1857) and Sars (1878) thought ft . (ft.) nana synonymous with f t . borealis Gray, 1839, but that species Is itself a Junior synonym of Polinlces (Eusplra) palIldus (Broderlp & Sowerby, 1829). Discussioni All of the eastern Pacific adult specimens 1 have examined have moderately elevated spires, but Odhner (1913) illustrates some Atlantic specimens that have low spires. This species Is commonly confused with Juveniles of Natica (Crvptonatica) clausa Broderlp & Sowerby in col 148 lections, understandably so considering their similar basic shapes. Living specimens are easily distinguished because £ 1 . (£.) clausa has a calcareous operculum, and shells of N. (N.) nana are translucent, whereas those of £ J , (£.) clausa are not. Shell form must be used to distinguish dead specimens, howeveri shells of N. (N.) nana are more elongate, with higher spires and more steeply sloping shoulders. In addition, the umbilical and parietal call! of £$. (£(. ) nana merge together more smoothly than do those of £1. (£.) clausa, the filling of the posterior apertural angle of £4 . (££.) nana is thicker and sticks out farther ahead of the aperture than in £ 4 . clausa. The cool-water habitat of £ 1, (££.) nana prevents its being found commonly in areas that have warm-temperate surface waters. However, it would have lived in shallower depths during the cool-water intervals of the Pleistocene, and its occurrence in the abundant marine Pleistocene deposits of southern California is not unlikely, although it has not yet been found there. flgverlta klrkensls (Clark, 1915) Plate 1, figures 34, 35 Natica (Eusplra) klrkensls Clark, 1915bi487, pi. 69, figs. 4, 9i Keen & Bentson, 1944i177. Natica klrkensls Clark, Trask, 1922i143. 149 Polinlces (EuanIra) klrkensls (Clark), Clark, 1929ipl. 34, fig, 2ui Keen & Bentson, 1944i186. Natica (Neverita) amoldl Clark, 1915b*488, pi. 68, figs. 13, 15i Keen & Bentaon, 1944*176. Natica amoldl Clark, Trask, 1922*143. Neverita amoldl (Clark), Adegoke, 1969*168-169. Description* Size.--Average* height 25 mm, diameter 20 mmi larg est specimen* height 43.5 mm, diameter 33.7 CUCB 1227, San Pablo Group, Concord quadrangle, California, Miocene]. Shell Form.--Shell usually elongate, rarely globose, spire moderately to greatly elevated* body whorl not greatly Inflated* whorls evenly rounded except for flatten ing below suture* shell thickness average* whorls 5^* suture slightly impressed, rarely moderately impressed. Shell surface smooth except for fine incremental growth lines. Parietal callus thick, heavily filling posterior apertural angle* anterior lobe inconspicuous. Umbilicus closed. Umbilical callus relatively narrow, evenly tapers anteriorly, smoothly merges with parietal callus* oc casionally depressed at anterior end. Anterior inner lip greatly thickened, elevated* basal lip slightly thickened. Operculum. - -Unknown. SraUgna fixftfU n ed i 152. 150 Stratlaraohlc Occurrence i Known only from Upper Miocene strata. Upper Miocenei Cierbo Formation, Mount Diablo quadrangle, California UCfi)i San Pablo Formation, Contra Costa County (Clark, 1915b), Concord quadrangle (UCB), Californiaf Neroly Formation, Mount Diablo quadrangle (UCB), Tassajara quadrangle (UCB), Californiai Briones Formation, Concord quadrangle, California (UCB)i Santa Margarita Formation, Coalinga area, California (Adegoke, 1969), Miocenei Pittsburg Bluff, Oregon (CAS)i Clata- kanie, Oregon (CAS). Type Localities« Natica (Euspira) kirkensis - UCB 467, southeast of Oyster Point, NE^, NE%;, section 23, T1S, R1E, Contra Costa County, California, Lower San Pablo Forma tion, upper Miocene (Clark, 1915bt Keen & Bentson, 1944). Natica (Neverita) amoldl - UCB 505, a little west of center of SE^, NEfc, section 9, T1S, R2W, Concord quadrangle, Contra Costa County, California. Upper San Pablo Formation, Upper Miocene (Clark, 1915bi Keen & Bentson, 1944), Iym Material* Natica (Euspira) klrkensls - Holotype, UCB 11591. Natica (Neverita) amoldl - Holotype, UCB 11594. 151 Nomenclature! Commentaryi Merita (Neverita) k^rkenaifl and N* (£}•) amoldl are synonyralzed here for the first time* The holotype of £J . (£.) klrkensls is more globose than the more typically elongate holotype of £ 4 . (£J.) amoldl. but the specimens are otherwise identical. Both species were described by Clark (1915b) In the same pub lication, and N. (£}.) klrkensls is chosen arbitrarily as the senior synonym because it is described one page prior to N. (£J.) amoldl. Discussioni The most noticeable variation in this species is in shell proportions. The great majority of specimens are distinctly elongate, but individuals occasionally occur that are fairly globose. Juveniles are slightly less slender than adults. The width and taper of the umbilical callus also varies noticeably and some specimens, es pecially juveniles, have a nearly imperceptible anterior taper to the callus. When this occurs, specimens may re semble Polinlces (Eusolra) palIldus (Broderlp & Sowerby, 1829), although most Recent specimens and all fossils of £. (£.) t>alIldus have a narrowly open umbilicus. Recent individuals of £. (£.) pallidus with closed umbilici are distinguished from £f . (£.) klrkensls by their centrally swollen umbilical call! and more strongly impressed sutures. Polinlces (Eusolra) pallidus is known from the Middle Pliocene to Recent of western North America, and 152 may be related to J*. (£}.) klrkensls. although similar forms are not known from Lower Pliocene rocks. Neverita (Neverita) polltiana (Dali, 1919) Plate 1, figure 36 EugcUfl PPUUftna Dali, 1919i353. Polinlces (Eusolra) polltianus (Dali), Dgll, 1921i164 fas (E.) polltlana"1i Oldrovd. 1927i727i Burch, 1946i Polln|^es polltianus (Dali), Keen, 1937*44* Burch, 1944* Description* £212£•- •Shell white. Periostracum thin, pale yellow ish brown. Size.--Heiaht 16 mm, diameter 13 mm (holotype only). Shell Form.--Shell globose, spire low* whorls evenly roundedj shell thin* whorls about 4* suture slightly im pressed and narrowly channeled. Spiral sculpture lack ing* axial sculpture of Incremental growth lines and sharply Incised grooves radiating from the suture part way to the periphery. Parietal callus thin, but thickens to moderately fill posterior apertural angle* anterior lobe Indistinct* Umbilicus closed. Umbilical callus small, low, with a small dimple at its growing margin. Inner lip slightly thickened, basal lip thin. Operculum.--Chitinous, filling aperture. 153 Geographic Occurranee and Ecologyi Known only from the type locality in the Bering Sea, at a depth of 1098 meters, on sand and pebbles. Tvoe Localityi Petrel Bank, Bering Sea Qabout 52°08'N, 179°48* e3i in 1098 meters, U.S.S. "Albatross'* station 4779 (Dali, 1919). Tvoe Materiali Holotype, USNM 205653 (Dali, 1919). Discussioni This species is known only from the type specimen. Neverita (Neverita) new species Plate 1, figure 37 Description! Color. — Shell exterior white, with narrow dark brown or white band just below suturet interior, parietal callus and umbilical callus white. Perlostracum thin, light gray ish brown. Shell Form.--Shell elongate, spire elevated, body whorl noticeably inflated, with a distinct, narrow groove shallowly incised a short way below suturei shell thlm whorls 4^, nuclear whorls not clearly differentiated, earliest nuclear whorl sunk into following whorli suture moderately impressed. Spiral sculpture of minute, wavy, closely spaced coatellae, absent from base and earliest 1% 154 whorlsi axial sculpture of incremental growth lines. Parietal callus thick, moderately to heavily filling posterior apertural angle % anterior lobe weak but general- ly distinct* Umbilicus usually closed, may be extremely narrowly open along entire margin. Umbilical callus elongate, with greatest width near or at posterior end, smoothly merging with parietal callust tapers anteriorly nearly to a pointt callus evenly thins toward growing margin or more abruptly ends at a marginal depression that may be deepest posteriorly. Umbilical callus bounded anteriorly by low, sometimes indistinct rib on body whorl. Anterior inner lip thickened. Dimensions of holotypei height 18,5 mm, diameter 16.1 mm. Operculum■--Chitinous. filling aperture. Type Localityi Off central Oregon at 44°34.8*N lat*, 125°33.6*W long., 2816 meters depth. Collected Department of Oceanography, Oregon State University, R/V Yaquina, sample BMT 186, 16 March 1970. Type Material! Holotype, LACMt 1 paratype, LACMj 1 paratype, USNMi 1 paratype, CASi 1 paratype, AWH. Height and diameter dimensions for paratypes are* 18.6 mm X 17.1 mm, 19.0 mm X 16.6 mm, 19.9 mm X 17.6 mm, 18.7 mm X 17.3 mm. Discussioni Characterized by its elongate shape, thin 155 shell* elongate umbilical callus* and shallow groove Just below the suture, £. n. sp. is not easily confused with any other eastern Pacific natlcld. It most closely re sembles (£!•) nana (Moller, 1842), a clrcumboreal species that ranges southward in progressively deeper to Cabo San Quintin* western Baja California* in depths of 11 to 1281 meters. Neverita (Neverita) nana is smaller (maximum sizet height 17.3 mm, diameter 14.8 mm, but usually much * less) than £. n. sp., with a thinner parietal callus, more weakly impressed suture, less inflated body whorl, thicker shell, and lacks a narrow groove below the suture and a depressed first protoconch whorl. The differences between these two species are clear, and n. sp. seems to be un related to any other eastern Pacific natlcid. The holotype and all but one paratype were alive when collected. The paratypes are from depths of 2760 to 2860 meters in the vicinity of the type locality. All living specimens but one have a dark brown band Just below the suture. The lone exception has a white band Instead of a brown one, which is still distinct for being more brightly white than the rest of the shell. One paratype has a very narrowly open umbilicus, along the entire growing margin of the umbilical callus. This feature is atypical, be cause all other specimens have firmly closed umbilici. The radula was removed from one paratype. It is not illustrated here because, as for nearly all other eastern 156 Pacific natlclds, its detention is not taxonomlcally use ful. Subgenus Glossaulax Filsbry, 1929 Glossaulax Pllsbry, 1929i113, Type-speciesi Neverita reclusiana (Deshayes, 1839), by original designation. Miocene to Recent, eastern Pacific. Figured herein. Diagnosisi Umbilical callus entirely covering umbilicus, divided into anterior and posterior lobes by narrow trans verse groove. Discussioni Glossaulax is characterized by its massive grooved callus. Some species, including the type-species, show variations with the umbilicus open, although the callus groove always remains distinct. In the eastern Pacific, this subgenus occurs mainly in temperate waters, but ranges into the tropical Panamic province. In western North America it is found in deposits as old as Late Oligocene, Neverita (Glossaulax) andersonl (Clark, 1918) Plate 2, figures 1, 2 Natica reclusiana andersonl Nomland, I9l7b|301 fnomen nudum» as “N. recluziana andersonl Clark (MS)"]. 157 a&a Natica (Neverita) reclusiana andersonl Clark. 1918i168- 169, pi. JO, figs. 3, 10-lJ pis " f i. (£.) recluzlan andersonl. n, var."3i Keen & Bentson, 1944i176 "N. (N.) recluziana andersonl"!. Natica andersonl (Clark), Hanna, 1924*173. Polinlces (Neverita) recluslanus andersonl (Clark), Grant & Gale. 19311802 fas "P. (N.) recluslanus variety andersonl"1i Loel & Corey, 1932i270, pi. 65, figs. 4a,b [as "£. (fj.) recluzlanus andersonl"! . not figs. 5a,b f- £. recluslanus (Deshayes. 1539)1i Keen U Bent son, 1944*185 fas "£. Qj.) rec luzlana var. andersonl" and "£. (fi.) recluzlanus andersonl"7. Polinlces recluslanus andersonl (Clark), Lutz, 1951*380, 39l, pi. 18, fig. 4 fas "£. recluslanus var. andersonl"1. Never^a andersonl (Clark), Keen, 1943*37j Stanton, 1966* Neverita reclusiana andersonl (Clark), Adegoke, 1969*168. andersonl (Clark), Addlcott, 19 d| Addlcott, 1970*67-69, pi. 5, figs. 22-24. Neverita reclusiana (Deshayes), Woodrlng gp ftl*, 1940*86 6 n part], pi. 15, figs. 19, 20, pi, 20., figs. 2, 4 piot N* reclusiana (Deshayes, 1839)3, not pi. 20, figs. 1, 3, 5 C* £ i . reclusiana (Deshayes, 1839)3. not Natica anderssonl Strebel, 1906*142-143, pi. 11, figs, 67, o/a,b QRecent, Magellanic region!* ?Natica ocovana Conrad, 1855*18i Conrad, 1857*pi. 7, fig. 57i Addlcott, 1970*68 [as noaen dublunf. Natica (Neverita) pabloensls Clark, 1915b*488-489, pi. 68, figs. 12, 14) Keen & Bentson, 1944*177. Natica pabloensls (Clark), Trask, 1922*143-145. Polinlces recluslanus pabloensls (Clark), Clark, I929tpl. 34Tfig. l21 Grant & Gale, 1931*802-803 [as "£. (fi.) r^gju^|gnus variety pabloensls"3 * Keen & Bentson, Neverita (Glossaulax) andersonl (Clark), Addlcott, 1965* 105, pi. A, fig. 158 Descriptioni Color.--Addlcott (1970) described cream-colored sub- sutural and basal color bands for Middle Miocene specimens of Neverita aj£a O N. reclusiana J from the Round Mountain Silt of the Kern River area, California, and noted similar banding on specimens of £• andersonl. 1 have seen one specimen from the Middle Miocene Topanga Formation of southern California with a dark brown parietal and umbili cal callus. Sl2e.— Averagei height 32 mm, diameter 29 mmi larg est speclment height 62.0 mm, diameter 58.3 mm [[CAS 69, Pyramid Hill, Kern County, California^. Shell Form.--Shell relatively low to elongate, spire usually low, may be elevated* body whorl not greatly in flated' sides of body whorl flattened, narrowly tabulate' shell thickness average' whorls about 5%i suture very slightly impressed. Spiral sculpture of minute, weakly developed, closely spaced, minutely wavy, costellae, usually not preserved' axial sculpture of incremental growth lines that are heaviest on base. Parietal callus heavy, thickly filling posterior apertural anglei anterior lobe weak to indistinct, merges with umbilical callus. Umbilicus usually closed, may be partly open, especially in Juveniles* Umbilical callus massive, divided into two lobes by weak transverse groove' posterior callus lobe is always larger one, is elongate 159 anteriorly to plug or nearly plug umbilicust callus thick, generally with swollen appearance* Anterior Inner lip thickened. Operculum *--Unknown. presumably chitinous* Specimens Examinedi 1511. Stratlaraphlc Occurrencei Ranges from Lower Miocene to Upper Pliocene. Lower Miocene (?)i San Ramon Formation, Contra Costa County, California (Clark, 1918). Lower Miocenei Vaqueros Formation, Santa Cruz Mountains, northern Gabilan Range, Santa Lucia Range, La Panza Range, San Emigdlo Mountains, western Santa Ynez Range, Ventura basin, Channel Islands, western Santa Monica Mountains, San Joaquin Hills, Santa Ana Mountains (Loel & Corey, 1932), Mount Pinos quadrangle, Junlpero Serra quadrangle, CoalInga quadrangle, (UCB), Hew Idria quad rangle (USGS), and Callente Range (Eaton si £i*» 1941), Californiai Fainted Rock Sandstone, La Panza quadrangle (USGS) and Pozo quadrangle (USGS)j Jewett Sand, Kern River area, California (Addlcott, 19701 USGS). Middle Miocenei Astoria Formation 7, Claskanle, Oregon (CAS)i Sobrante Formation, Pacheco-Walnut Creek area, California (Lutz, 1951)t Monterey Formation, Saltos Shale Member, Callente Range, Chlmineas quadrangle, California (USGS)i Gould Shale, Temblor Range, California (USGS)i Oursan Sandstone, Pleasanton area, California (Hall, 1958)i 160 Oursan (?) Sandstone* Tesla quadrangle, California (Clark in Huey, 1948)i McClure Shale, Reef Ridge quadrangle, California (USGS)t Temblor Formation, Coalinga area, La Panza Range (Loel & Corey, 1932), Callente Range (Eaton et al.. 1941), Calabasas quadrangle, San Joaquin Rocks quadrangle, Priest Valley quadrangle, Dominguez Ranch quadrangle (UCB), Panoche Valley (USGS), California) Topanga Formation, Santa Monica Mountains (LACMf USCi USGS) Woodrlng in Hoots, 1931, "cf.") Loel & Corey, 1932), Santa Ana Mountains (Loel & Corey, 1932), California, Round Mountain Silt, and upper and lower Olcese Sand, Kern River area, California (USGS) UCB) Addlcott, 1970). Upper Miocenei Neroly Formation, Tassajara quadrangle, California (UCB)f upper and lower San Pablo Group, central California (Clark, 1915b)f Santa Margarita Formation, Coalinga area (Nomland, 1917bi Clark, 1918) UCB), Stock- dale Mountain quadrangle, Chalome Hills (USGS), Priest Valley quadrangle (USGS), Pozo quadrangle, San Luis Obispo quadrangle (USGS), California. Upper Pliocenei San Joaquin Formation, Kettleman Hills, California (Wood- ring fit al,, 1940). Pliocenei "Lower Fernando" (Clark, 1918). Tvoe Localities) Natica (Neverita) recluziana - UCB 1131, in creekbed half a mile southwest of walnut Creek 161 about 100 yards east of Oakland and Antioch bridge altitude 150 feet. Contra Costa County, California* San Ramon Formation, upper Oligocene or lower Miocene (Clark, 1918i Addicott, 1970). Nat lea ocovana - Ocoya Creek fFoso Creek]}, Kern County, California. Miocene (Conrad, 1855). Nattea (Neverita) pabloensls - U C B 231, 38, 1227. Upper and lower San Pablo Group, Central California, upper Miocene (Clark, 1915b). Type Material< Natlea (Neverlta) recluzlana anderaonl - Holotype, UCB 11212. Natlea ocovana - Unknown, presumably lost (Keen & Bentson, 1944). Natlea (Neverlta) pabloensls - Hoiotype, UCB 11592. Discussioni Neverlta (Glossaulax) andersonl is closely related to ££ . (G.) recluslana (Deshayes, 1839), and specimens are sometimes difficult to assign to one species or the other. When typically developed, as are the great majority of specimens, (£.) andersonl is characterized by a body whorl with flattened sides and tabulate shoulder, massive callus thickly filling the umbilicus, and low spire. The most common variation is for a deep pit to form at the anterior margin of the umbilical callus, at the site of the closed umbilicus, or, less commonly, for 162 the umbilicus to be slightly open. An open umbilicus is not common, but Is most often seen on immature specimens, although a closed umbilicus still predominates among these younger specimens. Adult or Juvenile specimens with an open umbilicus are always high*spired and look somewhat like the so-called alta form of (G.) recluaiana■ al though the posterior callus lobe is never as elongate as in that form. The alta form also never has the flat-sided whorls of N. (G.) andersonl. Some specimens have more elongate shells than usual, although they retain their low spires. Specimens occur rarely with greatly elevated spires, but retain other specific features such as flattened, slightly tabulate shoulders and massive callus closing the umbilicus. Neverlta (Glosaaulax) andersonl is most similar to the so-called imperforata form of £. (G.) recluaiana. which Is known from Pleistocene and Recent assemblages. The main differences between the two are that the imper- forata form has a less elongate shell, has its outer apertural lip attached lower on the preceding whorl, and has more gently sloping margins to Its umbilical callus. The Imperforata form often has an evenly rounded shoulder profile, but some living and fossil specimens have whorls nearly as flattened as typical £, (£.) andersonl. The margins of the umbilical callus of f j. (£.) andersonl are Invariably steep, especially anteriorly, whether the callus 163 is complete or noti unlike most (G.) recluaiana. The transverse groove of |J. (G.) andersonl Is less strongly impressed than on any variety of £1. (G.) recluaiana ex cept the so-called imperforata form. (G.) andersonl is the most common natlcid in Miocene deposits of California. It is sometimes found in huge accumulations of specimens that show consistent morphology not Intergrading with that of other species. Specimens of the related species (G.) recluaiana and ££. (G.) lamesae (Moore, 1963) are relatively uncommon whether or not they are found along with (G.) andersonl. (G.) andersonl is known from a single occurrence north of California, In probable beds of the Astoria Formation at Clatskanle, Oregon. It is replaced as the common Mio cene natlcid in the Pacific Northwest by £. (£.) gallanol Dali, 1909, which was apparently adapted to cooler-water conditions. I have seen specimens essentially identical with those of N. (G.) andersonl in collections of Miocene fossils from Chile and the Caribbean region. (G.) andersonl and related species and forms in the eastern Pacific are thus only a part of a larger occurrence of similar or Identical species throughout the Miocene tropics. It is not yet possible to describe the origin of this diverse and widespread group. There are also similar species in Ollgocene deposits of the eastern 164 Pacific, and future study of these may uncover the begin nings of the f $ . (G.) andersonl-N. (G.) recluaiana stock. Neverita (Glossaulax) recluaiana (Deshayes, 1839) Plate 2, figures 3, 4, 5, 6, 7 Natlea recluaiana Deshayes, 1839i361, Deshayes, 1841ipl. 371 Menke, 1851*165 fas recluzlana"1 * Philippi, 1852*38-39, pi. 5, fig. 5 (plate lo49)* Hupe, 1854* 219-220 fas " l i- " * Reeve, 1855 ipl. 1, fig. 3 fas " J S« recluziana"!* Sowerby, 1883*76, pi. 1, fig. 6 fas " | * . recluzlana"! * Nomland, 1917a*221 fas recluzlana Petit"J* Nomland, I9l7b*301 fas "I*, recluzlana Petit"}i Wagner & Schilling, 1923*243- 246, 248* Bremner, 1933*16 [as cf.}. Neverlta recluaiana (Deshayes), H. & A. Adams, 1853*vol. 1, p. 208i Carpenter, 1864*661 [as "H. recluzlana Petit"}* Carpenter, 1878*12 fas recluzlana Petit"}i Dali, 1878*12 fas " J J. recluzlana Petit"}* Keep, 1888*46, fig. 26 fas recluzlana Petit"3* Dali, 1892*369 fas "JJ. recluzlana"!* Arnold. 1907a* 544, pi. 48, fig. 6 fas "$f. recluzlana Petit"}* Arnold, 1907b*pi. 38, fig, 6 ras "N. recluzlana Petit"}* Arnold, 1907c*2o, pi. 54, figs. l4a,b.15 fas "U. recluzlana Petit"}* Arnold & Anderson, 1907* 144, pi. 21, figs. 14a,b, 15 fas " | * . recluzlana Petit"}* Arnold & Anderson, 1910*pi, 22, fig. 2 fas "N. re- cluziana Petit"}* Arnold & Anderson, 1910*110, 130, 133, pi. 42, fig. 2 fas "JJ. recluzlana Petit"}* Arnold, 1910*31, 146, pi. 20, rigT 2 [as " £ J. re cluzlana Petit"}* Nomland, I9l7a»213, 221 fas "N. recluzlana Petit"}* Pllabry, 1929*109, pi. 6, fig. 1| Clark, 1931* sp. list* Woodring fit a£* * 1940*86 fin part}, pi. 20, figs. 1, 3, 5, not pTT 15, figs. 19, 20, pi. 20, figs. 2, 4 C- £. andersonl (Clark, 1918)}* Keen & Bentson, 1944*180 fas "N. recluzlana"}* Stewart, 1946itable 2* Durham, 1950*1277 pi. 54, fig. 8* Woodring & Bramlette, 1950*73, pi. 20, fig. 4* Valentine, 1956*200* Rodda, 1957*2483* Valentine, 1960*20 fas "N. recluslana. £.!•"}* Valentine, 1961* 332, 333, 336, 338, 5457 361, *70, 372, 387, 389* Winterer & Durham, 1962*296* Addlcott & Vedder, 165 1963i67i Durham & Addlcott, 1965t12t Stanton, 1966i 23, 33i Adegoke, 1969i167-168* Kern, 1971t813. Natlea (Neverlta) recluslana (Deshayes), Tryon, 1886*34, pi. 2, fig, 1 Cas "N. QJ.) recluzlana") i Williamson, 1892*211 (as "N. (N.) recluzlana PetTt"Ji Moody, 1916*44* Martin, 1916*555" 239, 245, 257. Pollnlces (Neverlta) recluslanus (Deshayes), Steams, 1894*196 Cas "Polynlces (N.) recluaiana"3* Axnol 1903*314-315, pi _ _ recluslanus (Deshayes), Steams, "Polynlces (N.) recluaiana"3* Arnold, . lfl, fig, 12 fas ^Pojvnices (£.) "]* Reagan, 1909*171 (as "P. (JJ.) recluzlana Petit"]* Reagan, 1909*171 Cas "P. recluslana"?. 193 [as "P. (N.) recluzlana" 1. pi. 3, fig. 28* Dali, 1909a*236* Reagen, 19l0*648* Dali, fOlUllCgg (Neverlta) recluslanus recluslanus Grant & Gale, 1931*801-802 (As (N.) recluslanus variety recluslanus. s.s."]. PolInIce9 recluslanus (Deshayes), Jordan, 1924*150-152 fas "P. recluzianus"?* Jordan, 1926*246* Jordan, 1936*114* Keen, 1937*44 (as "P. recluzianus"1* Smith & Gordon, 1948*199* MacGinitle & MacGlnitle, 1959* 330, 355, figs. 170, 202 Cas recluzlana"1 * Hertlein & Emerson, 1956*166* Kanakofi & Emerson, 1959*30* Addlcott A Emerson, 1959*16 (as "cf."]* McLean, 1969*37, fig. 19.3. Neverlta callosa Gabb, 1866*10-11, pi. 2, figs. 17, l7a,b (plate 1869)* Arnold, 1907a*542, pi. 44, figs. 4,4a* Arnold, 1907b*234, pi. 31, figs. 4,4a* Eldrldge & Arnold, 1907*147* Arnold, 1908*350* Arnold & Ander son, 1910*85, 86* Anderson & Martin, 1914*43* Keen, 1943*12* Keen & Bentsor., 1944*179* Adegoke, 1969* 167* Addlcott, 1970*68, 69. Pollnlces calloaus (Gabb), Arnold U Hannibal, 1913*576 &s "£. gaHaa»"]. Pollnlces (Neverlta) recluslanus calloaus (Gabb), Grant & Gale, 1931*803, text-fig. 14 fas "£. (£1.) re cluslanus variety calloaus"?. 166 Neverlta recluaiana alta Dali, 1878i12 Cnomen nudumi as var. alta"1i Arnold, 1910*32, 146, pi. var, alta Dall"]| pi. 45, fig8, 5, recmsLana aica uau, lo/onz "N. recluzlana var. alta"!* Amoli 20, figs. 5,5a Cas "N. recluzlana Arnold & Anderson, 1910 *i30,133, 5a Cas "N. recluzlana var. alta Dali"]* Valentine, 1956i200T Valentine, 1957i30(Jj Valentine, 1961*357, 360, 361, 364, 376, 377, 386, 388, 408i Valentine L Meade, 1961*8, 12, 17, 19, 20, 23, 27f Adegoke, 1969*168. _ _ recu________ Dall"]( Oldroyd, 1927*732 Cas "£. (N.)recluzianus alta Dall"]i Grant S t Gale, 1931*801-802 Cas probable male form of "£. (£J.) recluslanus reclusianus"!. Natlea recluslana alta (Arnold), Nomland, 19l7a*221 Cf t s "N. recluzlana alta Dali"]. Neverlta alta (Arnold), Pllsbry, 1929*110-111, pi. 6, figs75^9 Cas alta 'Dali' Arnold"] i DeLong, 1941* sp. list. PollnlcfB altus Arnold, Keen, 1937*44i Kanakoff S t Emerson, 19?9«301 McLean, 1969*37-38, fig. 19.4 Cas "£. altus Pllsbry"]. Pollnlces (Neverlta) altus Arnold, Willett, 1937*400 Cas "£. (N . V altus DallH]i Burch, 1946*31. Pollnlces recluslanus altus Arnold, Smith S t Gordon, 1948* 199 Cas "£. recluslanus altus Dali”]. Neverlta (Glossaulax) alta (Arnold), Addlcott, 1970*69, pi. 5, figsT 19, 25. Pollnlces (Neverlta) recluslanus imperforatus Dali, 1909b* 88 Las "P. (N.) recluzlanavar. Imperforata Stearns"]t Dali, 1921 * 155 Cas £7 (£1.) recluzlana Imperforata"!i Oldroyd, 1927*731 Cas "£. (N.) recluslanus imper forata Steams"]) Grant & Gale, 1931*8021 Burch, 1946730-31 Cas "£. (£}.) recluzianus Imperforatus"!. Neverlta recluslana imperforata (Dali), Pllsbry, 1929* Ul-1157 pl. 6, figs. 2-ei DeLong, 1941isp. list Cas "N. recluzlana imperforata"!i Woodring sL a£*» 1946*72| Valentine, 1901*3017^376. 167 Pollnlces Imperforatus Dali, Keen, 1937i44. not Natlca Imperforata Gray, 1839*135, pi. 37, fig, 1 [Recent, Cape of Good Hopei. Description* Color.-*Shell buff to pale gray, with a darker band high on the shoulder and a narrower white band usually at the suturei base white or at least paler than rest of whorl* umbilical callus white to medium brown, may be white centrally with a brown margini parietal callus white, may be brown on Its anterior half) Interior clouded with white and light to dark brown, white on basej nuclear whorls dark brown. Perlostracum thin, whitish on base, pale grayish brown to chocolate and rust brown above base, often with irregular rusty staining. Dorsal part of shell, not covered by mantle in life, may be darker than remainder of shell. Size.--Averagei height 40 mm, diameter 38 rami larg est specimen* height 90.5 mm, diameter 82.0 mm CCAS 1937, San Diego, California?. Shell Form.--Shell low to elongate, spire moderately to greatly elevated) body whorl not greatly inflated) shoulder slightly to distinctly flattened, commonly slightly concave, especially in high-spired forms) shell thick to average) nuclear whorls 2k, smooth) postnuclear whorls 4) suture slightly impressed. Spiral sculpture of minute,%weakly developed, closely spaced, minutely wavy, 168 costellaei axial sculpture of incremental growth lines chat are heaviest on base« Parietal callus heavy, thickly filling in posterior apertural anglei anterior lobe weak to indistinct, merges with umbilical callus. Umbilicus broadly open to closed, with all intermediate gradationst broadly open umbilicus shows weakly developed spiral cords in channel* Umbilical callus clearly divided into two lobes by strong transverse groovef groove may be simple or terminate as a deep pit at inner lip margini posterior callus lobe is always larger one, may be confined to posterior end of umbilicus or may be elongate anteriorly to plug or nearly plug umbilical opening. Anterior inner lip and basal lip thickened. Operculum,— Chltinous, filling aperture. Specimens Examinedt 3200. Geographic Occurrence and Ecologyi Crescent City, Cali fornia (41°46'N), south to the Gulf of California, through out which it occurs, and to Just south of Mazatlan (23°14'N), and at Islas las Tres Marlas (2l°30'N, 106o40'W), Mexico. Uncommon north of Mugu Lagoon, California (34°05'N) and south of the Gulf of California. Living on sandy and muddy bottoms interltdally and in depths to 50 meters, common in embayments. Stratigraohic Occurrence« Ranges from Lower Miocene to 169 Recent. Lower Miocene (?)i Pliest Formation, Echino- phorla apta zone, King County, Washington (Durham, 1944)i San Lorenzo series, central California (Clark, 1918)t "Vaqueros Formation," San Miguel Island, California (Bremner, 1933). Lower Miocene (?)i Vaqueros Formation, Trabuco Canyon, Los Angeles County, California (LACM). Middle Miocenei Temblor Formation and McClure Shale, Reef Ridge, Coalings area California (Stewart, 1946t USGS)i Temblor Formation, Coalings area (Adegoke, 1969j UCB), Wilson Corner quadrangle (USGS), and Panoche Valley quadrangle (USGS), California» Clallam Formation, north western Washington (Reagan, 1909, 1910)t Astoria Forma tion, Montesano quadrangle, Washington (USGS)i Topanga Formation, Topanga Canyon, Los Angeles County, California (LACMi USC)t upper and lower Olcese Sand (Addlcott, 1970i USGS) and Jewett Sand (Addlcott, 1970i USGS), Kern River area, California. Upper Miocenei Santa Margarita and San Pablo formations, central California (Nomland, I9l7b)i Santa Margarita Formation, north Coalings area (Nomland, 1917b), La Panza quadrangle (USGS), and Coallnga area (UCB), California. Miocene to Pliocenei Imperial Forma tion, Carrlzo Creek, California (UCB)i Towsley Formation, southeastern Ventura basin, California (Winterer & Durham, 1962). Lower Pliocenei Jacalltos Formation, Coallnga area (Arnold & Anderson, 1910), and Priest Valley quad rangle (UCB), Californiai Empire Formation, Coos Bay, 170 Oregon (UCB)t Pancho Rico Formation, Hanes Valley quad rangle (USGS) and Salinas Valley (Durham & Addlcott, 1965), Californiai Repetto Formation, San Fernando quadrangle, California (CASt UCB)i Slsquoc Formation, Lompoc quad rangle, California (UCB), Middle Pliocenei Etchegoln Formation, Coallnga area (Martin, 1916j Nomland, I9l7at UCBj Arnold, 1910), and Palvadero Gap quadrangle (UCB), California, Carmen Island, Gulf of California, Mexico (IXirham, 1950), Middle Pliocene to Pleistocenei Merced Formation, HeaIdsburg quadrangle (UCB) and Sargent oil field, Santa Clara County (Martin, 1916), California. Upper Pliocenei Pico Formation, San Fernando quadrangle (UCB), southeastern Ventura basin (Winterer A Durham, 1962), Plru quadrangle (UCB), Triunfo quadrangle (UCB), Val Verde quadrangle (UCB), and Elsmere Canyon (Arnold, 1907a), Californiai San Joaquin Formation, Pecten and Trachvcardlum zones, Neverlta zone, Acila zone Ccf.3, Cascajo Conglomerate Member fas M?"J, Kettleman Hills, California (Woodring ££ a^,, 1940)j San Joaquin Formation, Los Vlejoa quadrangle (UCB) and Reef Ridge quadrangle (UCB), California. Fernando Formation, central Los Angeles Basin, California (LACM)i San Diego Formation, San Diego County (LACM), San Ysidro quadrangle (UCB), Pacific Beach (CAS), Balboa Park (CAS), and U.S.-Mexican border (CAS), California. Pliocenei Purlsima Formation, Ano Nuevo quadrangle, Sgn Mateo County, California (SU)i 171 Falor Formation, Trinidad Head, Humboldt County, Cali fornia (UCB)i Newhall Canyon, Los Angeles County, Cali fornia (CAS). Upper Pliocene to Pleistocene* Saugus Formation, Ventura quadrangle (UCB) and Piru quadrangle (UCB), California. Lower Pleistocenei Santa Barbara Formation, Santa Barbara, California (LACM) UCB)j "San Pedro Formation," Ventura County, California (CAS)j San Pedro Sand Member of San Pedro Formation, San Pedro area, California (UCBj CASj LACM) Oldroyd, 1924i Arnold, 1903t Woodring si &1., 1946)) Loraita Marl Member (Wood ring ££ al., 1946i LACM) and Timms Point Silt Member (Woodring ££ al«» 1946* Clark, 1931j LACM) of San Pedro Formation, San Pedro, California. Upper Pleistocene! Palos Verdes Sand, Palos Verdes Hills and San Pedro area (LACMj Woodring ££ al., 1946i Arnold, 1903i Valentine, 1961), Playa del Rey (LACM* CASi Willett, 1937), Newport Bay (LACMiCASi Kanakoff & Emerson, 1959), and Potrero Canyon, Pacific Palisades (Valentine, 1956), California) southwestern San Diego County, California (Emerson & Addlcott, 1953i Valentine, 1961)* Pacific Beach, San Diego County, California (Valentine, 1961)» San Clemente, Orange County, California (Valentine, 1961)i terrace deposits, Suiamerland and El Cap!tan Beach, Santa Barbara County, California (Valentine, 1961). 172 Type Localitiesi Natlea recluslana - "Mers de California” (Deshayes, 1839). Neverlta callosa - Walnut Creek, California, Miocene (Gabb, 1866). Polynlces (Neverlta) recluzianus var. alta - Pacific and Oliver Streets, San Pedro, California fpalos Verdes Sand, upper Pleistocene3 (Pllsbry, 1929). Pollnlces (Neverlta) recluzlana var. Imperforata - "living In the vicinity of San Diego and In the Pleistocene of the upper San Pedro fPalos Verdes Sand, upper Pleistocene3 at Deadman Island fsan Pedro, Califomlaj” (Dali* 1909), Type Material» / Natlea recluslana - Unknown, presumably in Ecole des Mines, Paris, or BM(NH) (Dance, 1966). Neverlta callosa - Unknown. Polynlces (Neverlta) recluzianus var. alta - Neotype, ANSP 147585 (designated by Pllsbry, 1929). EflllalCgg (flE Y O rU ft) recluzlana var. Imperforata - Neotype, ANSP 147436 (designated by Pllsbry, 1929). Nomenclature! Cof«^ntarvi The authorship of the synony- 4 mous species J J . (G.) alta has been variously attributed to Arnold (1903), Dali (1909), and Pllsbry (1929). The earliest mention of the name by Dali (1878) is a noman 173 nudum. Arnold (1903i315) noted that "Dr. Dell has des cribed a variety Q>f £) . (G.) recluslana! with an elevated spire which he calls var. alta. * * This is sufficient des cription to differentiate the taxon, so that authorship is correctly attributed to Arnold. In a discussion of (G.) recluslana. Dali (1909*88) later stated that "There is a variety alta Dali* with small narrow shell and ex ceptionally elevated spire*” which would have been enough to validate the taxon if Arnold had not already done so. Discounting both of these brief descriptions. Pllsbry (1929i110-111, pi, 6, figs. 5-9) redescribed the taxon more thoroughly as Neverlta alta and designated and figured a neotype. Most subsequent workers have used Pllsbry's concept of the taxon, although its authorship still be longs to Arnold (1903). Further consideration of (G.) recluslana and the so-called alta form of it is given be low. There was also early misunderstanding about authorship of the so-called imperforata form of (G.) recluslana. which Dali (1909) first cited as £. (Neverlta) recluzlana var. imperforata Steams. However, the name never appear ed in print prior to Dali's citation, but was probably a manuscript name used by Steams. Discussioni Although (G.) recluslana does not have a large number of synonyms, the allocation of given trivial 174 names to the subspecific, specific or varietal level by various workers indicates the instability of its morpho logic features. Shell proportions, umbilical morphology and color vary so widely that it has been difficult for past workers to assign all of the observed features to a single taxon, as is done here. The trivial names have thus been arranged in the past to account for portions of the whole range of morphology and color exhibited by the species. 1 believe that the synonymous names given here do not represent discrete morphologic or color entities, or consistent ecotypic forms. Further detailed collecting and observation may show that the variations are related to differences in local environments or microhabitats. The principal criterion for recognition of the so- called alta form has been its elongate proportions, even though Woodring fit a^, (1946»7l) noted that both £J . (G.) recluslana and its alta form have spires of variable height. Figure 5 shows a comparison of "typical” £ 4 . (G.) recluslana and so-called "typical" alta. based on ratios of shell height to diameter, taken from a variety of Recent LACM localities. This graph shows that "typical** alta specimens tend to have a slightly more elongate shape than "typical" £. (G.) recluaiana. but that the two forms thoroughly Intergrade. The measured specimens of alta also have a smaller average size than those of . (G.) re el us iarya . which confirms my observation that all of the FIGURE 5 Comparison of height to diameter ratios of Atlantic and Pacific specimens of Amaurop- als islandlea (Gmelln). Based on specimens from several institutions. 175 HEIGHT I N M M 176 100- N«v«rita rvclusiana N«v*rita alia 80- 60- 20- ♦ ♦ 't 40- • ; • . A • • "it:* + + * * . * * * • * T i---------- 1 ---------- r 20 40 60 80 DIAMETER IN MM 177 large specimens 1 have seen have "typical* J *. (G.) re cluslana features. It is difficult to describe all of the umbilical callus variations of (G.) recluslana without going into excessive detail. There are more recognizable variants than have been named, which is highly unusual for natlcids. As noted earlier, the umbilical callus is divided into an anterior and posterior lobe by a trans verse groove, and the umbilical variations largely depend on the degree to which this callus conceals the umbilicus, plus the relative sizes of the callus lobes. Completely Imperforate specimens have been called imperforata. moderately umbllicate specimens (£.) recluslana. and widely umbllicate specimens alta. Neverlta imperforata is the taxon most easily dispensed with* Its closed umbilicus is known only among relatively low-spired shells of the otherwise "typical" (G.) recluslana type. In a single population of imperforata are shells with every gradation of umbilical closure from Imperforate to narrow ly perforate to the moderately perforate shells of "typical" U. (G.) recluslana. Specimens of £. imperforata are usually present In Recent and fossil populations of IJ. (G.) recluslana in various proportions, and there seems to be no meaningful way to segregate them as a distinct species-level taxon. Neverlta alta is supposed to have a brown callus that 178 Is smaller than the white callus of (G.) recluslana. but callus size and color show a broad and nearly con tinuous range of variation between these two extremes. Entirely brown call! are common among both the smaIl eal lused and large-callused forms, although more common among the former. 1 have not seen a brown callus on an imperforate specimen, although some nearly imperforate specimens, including some of the largest specimens known, have partly pale brown calll. Many specimens of both callus types have mostly white calli with brown margins. Individuals showing all manner of callus colors may be found together, although one color type will usually pre dominate in a local population. Except in fortuitous cases, the colors of fossil specimens are not preserved. Diet is known to affect the coloration of other gastropod species, including naticids. Turner (1958) has shown that Individuals of duplicate (Say, 1822), the western Atlantic analogue of (G.) recluslana. have a brown or white umbilical callus, depending on diet* Those snails that feed upon the clam Mva arenarla Linnaeus, 1758, develop a white callus with a faintly pink margin, whereas those that feed on other claras develop a dark brown callus. Turner (1958) found that specimens with a dark callus developed white ones when fed a steady diet of J g . granarUi Despite supposed differences in umbilical coverage by 179 the callus between (G.) recluslana and alta. there is little difference between most specimens of the two types. In "typical" (G.) recluslana the two callus lobes are elongate to about the same extent, no matter what the absolute size or coverage of the callus. In "typical" I* , alta the posterior lobe Is distinctly more elongate and covers most of the umbilicus. As with shell height and callus color, however, there Is a continuous gradation In relative sizes of the umbilical callus lobes, with no clear distinctions between the two forms. More elevated shells tend to have elongate posterior callus lobes, but a large percentage do not, and many low-spired shells have elongate posterior lobes. Local Recent and fossil accumulations are predominantly of one type, but more extensive collecting shows a patchwork of such populations with intergrading umbilical features. Oddly enough, one of the more distinctive callus morphologies In the (G.) recluslana complex has not been given a subspecific name * In this form, the callus is relatively small, with the two lobes nearly equal in size, and the transverse groove Is deep and terminates In a strong pit at the Inner lip. The umbilicus Is broadly open and reveals some of the earlier whorls, and the shell Is relatively elevated* This form does Intergrade with "typical" (G.) recluslana. although not to the extent seen In J J . alta and £. Imperforata. Past workers 180 have presumably Included this form In £ j . alta. Except for the strong callus groove, this form resembles the Peruvian and Ecuadorian Recent species Pollnlces (Pollnlces) ravldus (Souleyet, 1852) In shell and callus form. As noted above, specimens of £J . (G.) recluslana and the forms Imperforata and alta are found together In both Recent and fossil populations. This Is especially well shown In prolific fossil assemblages such as those of the Upper Pleistocene deposits at San Pedro, California, where hundreds of specimens may be collected in a short time. All of the morphologic varieties of fJ. (G.) recluslana are found In such deposits. The present study has not confirm ed the observations of McLean (1969) that Individuals of N. (G.) recluslana are more likely to live In shallow bays and lagoons, whereas those of N* £l£a are more character istic of offshore sandy bottoms. Although the N. alta form may predominate in local populations, such popula tions are as commonly found In erobayments as in offshore habitats. For example, abundant specimens of alta have been collected in Newport Bay, Anaheim Bay, San Pedro Bay, Magdalena Bay, Bahia Adair, Morro Bay and Mugu Lagoon, whereas populations of (G.) recluslana occur at numerous open-shore localities throughout the range of the species. There appears to be no consistent distinction in habitats between the two forms. 181 Individuals of the alta form with a mostly brown callus range from Point Conception, California (Berry collection 18895) to Islas las Tres Marlas* Mexico (USNM 46535), whereas specimens of (G.) recluslana range from Crescent City, California (USNM 104130) to Islas las Tres Marias (USNM 46535). Their known northern range limits are thus different, which might relate to prey species available at different latitudes, because diet can affect callus color in naticids, as noted above. In rare Instances, individuals of J J. (2*) recluslana are essentially indistinguishable from those of Pollnices (Eusplra) lewis11 (Gould, 1847). Such individuals are globose, with relatively small call! and sometimes even have the slight shoulder flexure that otherwise character izes £. (£.) lewlsll. It is possible that such specimens represent hybrids of (£•) recluslana and £. (£.) lewlsll. although such a hypothesis would require experi mental proof. The few such specimens of (G.) re cluslana I have seen differ from £, (£.) lewlsll by having an elongate posterior callus lobe, a slightly thicker parietal callus, and a more abrupt anterior termination of the umbilical callus. Additional variations within the f t . (G.) recluaiana comples are the Neogene species (£*) andersonl (Clark, 1918) and (G.) lansgae (Moore, 1963), which are dis cussed separately. 182 Neverlta (Gloasauiax) jaroesae Moore, 1963 Plate 2, figures 8, 9 Neverlta (Glossaulax) lamesae Moore, 1963*28-29, pi, 2* figs. 5, 15, 19f Addlcott, 1970*70, pi. 6, figs. 8, 10* Moore, 1971*23, pi. 7, figs. 3-5. Description* Size. - -Average * height 21 ram, diameter 24 mini larg est specimen* height 22.6 ram, diameter 28.6 mm £CAS 2064, Kern River area, California]. Shell Form.--Shell relatively elongate, spire moderately elevatedi body whorl Inflatedi shoulder usually evenly rounded, may be flattened near suturei base distinctly flattenedi shell thini whorls 5j suture very slightly impressed. Shell smooth except for fine incremental growth lines that are heaviest on base. Parietal callus thin, lightly filling posterior apertural angle* anterior lobe weak. Umbilicus broadly open but not revealing earlier whorls, with its posterior portion oc cupied by funicle. Umbilical callus small, confined to posterior half of inner lip, clearly divided into two lobes by sharply incised transverse groovei both lobes rounded, with swollen appearancei lobes equally elongate or posterior lobe slightly longer* posterior margin of callus rarely detached from underlying whorli funicle occupies posterior half of umbilicus. Anterior inner lip 183 and basal lip not thickened. Operculum.--Unknown, presumably chitinous. Specimens Examinedi 25. Stratlgraohlc Occurrencei Ranges from Lower to Middle Miocene. Lower Miocene (?)i upper part of Twin River Formation, northwestern Washington (Durham, 1944, as Pollnlces vaneouverens1s Clark & Arnold, 1923). Middle Miocenei Astoria Formation, coastal Oregon (Moore, 1963) Washington ? (Moore, 1963)i Temblor Formation, Caliente quadrangle, California (UCB)t upper part of Olcese Sand, lower part of Round Mountain Silt (as cf.), Olcese Sand or Round Mountain Silt, Kern River area, California (Addlcott, 1970)f Round Mountain Silt, Caliente quad rangle, California (UCB). Middle Miocene (?)i near mouth of Fall Creek, where it enters Scappoose Creek, Oregon (USGS). Type Localityi About 17 feet above base of section ex posed in big headland in beach cliff about 600 yards north of Spencer Creek, Lincoln County, Oregon. Astoria Forma tion, middle Miocene (Moore, 1963). Type Materiali Holotype, USNM 563129, Discussiont £. (G.) lamesae belongs to the complex of species and forms associated with g. (G.) recluslana 184 (Deshayes, 1839), among which it is one of the more dis tinct taxa. It is characterized by a flattened base, by a relatively small, swollen umbilical callus that is bi sected by a sharply incised groove, and by a thin parietal callus* The spire always remains moderately elevated, but the body whorls of some specimens are more elongate than average. Juvenile specimens may be diffi cult to differentiate from (G.) recluslana. There are no living forms of (G.) recluslana similar to £1. (G.) lamesae. The compact, bilobed umbilical callus of £4. (G.) lamesae is similar to that of the Miocene and Pliocene species Pollnlces (Euspira) gaHanoi Dali, 1909, which occurs in the same region. The latter species is easily distinguished by its larger size, thicker parietal callus and higher spire. ]4. (G.) andersonl (Clark, 1918) occurs with (G.) lamesae. but is also much larger, with dis tinctly flat-sided whorls, lower spire, larger size, and much more massive callus entirely filling the umbilicus. Genus Callnatlclna Burch & Campbell, 1963 Callnatlclna Burch & Campbell, 1963*22.1. Type-speciesi Slgaretus qldfgyd-U Ball* 1397, by Original designation. Recent, eastern Pacific. Figured herein. Diagnosisi shells medium to large in size, globose* thin, 185 base distinctly flattened! whorls inflated) shoulder flattened) slightly concave below suture! sculpture of minute* closely spaced spiral costellaei suture slightly impressed. Umbilicus broadly open* simple. Inner lip thickened* lacking distinct umbilical callus* reflected to partly conceal umbilicus. Parietal callus very thin. Operculum chitinous, not entirely filling aperture. Radula typically natlcid* with tricuspate rachidian* one raulticuspate lateral, one blcuspate inner marginal, and one raonocuspate outer marginal tooth per half-row. Discussioni Callnatlclna is characterized by its thin shell, flattened base with widely open umbilicus, in flated whorls, and lack of an umbilical callus. As noted by Dali (1899*85), the type-species seems intermediate in shell form between Slnum. Eusolra. and Eunatlcina. The genus is monotyplc. Callnatlclna oldrovdli (Dali, 1897) Plate 2, figure 10 SlRaretus oldrovdli Dali, I897i85| Dali, 1899i85. Eunatlcina oldrovdli (Dali), Packard, 1918i325-326, pi. 35, figs. lOa,b| Dali, 1921*165, pi. 14, figs. 1, 3* Oldroyd, 1927*734, pi. 92, figs. 11, 11a! Keen, 1937i36i Burch, 1946i32» Smith & Gordon, 1948*199. Callnatlclna oldrovdli (Dali), Burch & Campbell, 1963* 221-553, pi. 347 fie®• 4-7, text-fig. 2. 186 Descriptioni Color.--Shell pale grayish brown, often slightly pinkish, with a narrow band of yellowish orange at the suturei base white. Interior white and medium brown, axially streaked, white at basei umbilical callus usually blotched with brown and white. Periostracura thin, pale yellowish white. Size.--Averagei height 42 mm, diameter 40 mmi larg est speciment height 81.4 mm, diameter 71.5 mm fSU 27053, Monterey Bay, California}. Shell Form.--Shell globose, spire moderately elevated; body whorl greatly inflated; shoulder flattened, slightly concave Just below suture; shell thin; whorls about 7 (apex always eroded); suture slightly Impressed. Spiral sculpture of minute, wavy, closely and irregularly spaced costellae that are best developed above the periphery; axial sculpture of incremental growth lines. Parietal callus very thin, lightly filling posterior apertural angle, anterior lobe weak but distinct. Umbilicus broadly open, partly concealed by reflected inner lip. Inner lip thickened, lacking a distinct umbilical callus, with a shallow but distinct notch at its posterior end, where it meets the parietal callus. Basal lip not thickened. Operculum.--Chitlnous, very thin, not filling aperture. Specimens 73. 187 Geographic Occurrence and Ecologyi Eureka (40°45'N) to San Diego (32°43fN) and Catalina Island, California. Found in sandy mud substrates in depths of 73 to 365 meters* most common in depths of 73 to 110 meters* Type Localityi Catalina Island, California, in deep water (Dali, 1897). Type Materiali Holotype, SU 6447. Discussioni The adult animal is too large to retract com pletely Into Its shell, as seen in numerous preserved specimens. However, one dried Juvenile frjSNM 111307i height 12.8 mm, diameter 14.5 nan] has its operculum tight ly fitting the aperture, sealing the desiccated animal within the shell. From this one example, it may be that Juveniles can fully retract their soft parts into the shell, whereas adults, being less susceptible to predators, are unable to do so. The shallow notch at the posterior end of the inner lip seems analogous to the callus groove of the subgenus Glosaaulax among the Follnlclnae, perhaps indicating a phyletlc link with that group. Genus Bulbua Brown, 1839 Buibus Brown in J. Smith, 1839*104, not Humphrey, 1797*31 (non-binomial]. 188 Type-aoeclea i Bulbua amlthll Brown In J. Smith, 1839 ( • " Natlea fraallls Leach, 1819), by monotypy, Late Cenozolc to Recent, Clrcumboreal. Figured herein. Diagnosisi Shells snail to medium In size, globose to slightly elongate, thlnt whorla evenly rounded to slightly flattened at suture, sculptured with nlnute, weak spiral costellae. Umbilicus closed to very narrowly open and silt-like. Umbilical callus quite thin, slender, Parietal callus very thin. Operculum chltlnoua, thin, entirely filling aperture. Radula with monocuspate rachidlan, one monocuspate lateral tooth and outer marginal tooth, and one blcuspate Inner marginal tooth per half-row. Discussioni Buibus Is characterized by Its extremely thin shell, rounded base and slit-like umbilicus. It lacks the deeply channeled suture and more elongate proportions of AMUEfiBfll&e And the more widely open umbilicus of Chorlstes. Bui bus is known living only In Polar or sub polar regions. Among eastern Pacific naticids, only £• fraallla (Leach) and the two Chorlstes species have monocuspate rachldlans• 189 Bulbua fragllla (Leach, 1819) Plate 2, figure 11 Natlea fraallla Leach, 1819tApp. 2, p. 62t Philippi, 1853i151, 7 Natlea fraallla Conrad, 1830i222, pi, 9, fig. 3. 7 Natlea fraallla J. Smith, 1841*l56. Natlea flava Gould, 1839i196i Gould, 1841i239-240, fig, 132i Middendorff, 1849*422-424i Philippi, 1851*114- 115, pi. 16, fig, 5j Sowerby, 1883*79, pi, 8, fig. 175. Bulbua flavua (Gould), Gould, 1870*347-348, fig. 616i Habe & Ito, 1965*31, pi. 8, fig. 9. Lunatla (Bulbua) flavua (Gould), Dali, 1885*182. Amaura (Aervbia) flava (Gould), Filabry, 1895*72. Acrvbla flava (Gould), Odhner, 1913*9, 46-47, pi. 4, figs. 26-28. Bulb^ fl|vua elonaatua Habe & Ito, 1965*31, pi. 8, Bulbua anlthli Broun In J. Smith, 1839*104, pi. 1, fig. 18. Natlea amlthll (Brown), Jeffreys, 1867ivol. 4, p. 230. Acrv^a amlthll (Brown), Harmer, 1921*699-700, pi. 55, Natlea aoerta Loven, 1847*149) Middendorff, 1849*419* Middendorff, 1851*206, pi. 11, figs. 1-3) Philippi, 1853*146) Danielssen, 1861*31. Bulbua aoertua (Loven), Keen, 1937*31) Burch, 1946*33. X (Brown), Sara, 1878* pi. 21, fig. 18, pi. V 1 L 5 8 « p U f Alf i l^a lU| p i a v | fig. 9) Frlele & Grieg, 1901*111, 68 ), Sara, 1878*155-156, pi. 12, fig. 18, pi. V, fig. 9, pi. XVIII nudumJi Lea, 1846* 190 7 Naj^ca (Po Unices) tenuicula Sowerby, 1915i166, pi. 10, ? Na^^ca alacl^^Janielssen. 1861131 £not £. Rlacialls Description* Color.--Shell exterior and Interior white. Perlo st racum thin, resistant, colors range from light yellow ish brown and gray to dark olive brownj colors often axially streaked* rust brown staining common on upper parts of whorlsi base paler. Size.--Averagei height 23 mm, diameter 18 ramj larg est specimen* height 51.0 mm, diameter 39.0 mm fCAS 39460, Bristol Bay, Alaska^. Shell Form.--Shell globose to somewhat elongate, spire low to moderately elevated* body whorl inflated* shoulder flattened* shell thin* whorls about 5 (apex always eroded)* suture slightly impressed. Spiral sculpture of minute, closely and irregularly spaced costellae that are strong est above periphery* axial sculpture of Incremental growth lines. Parietal callus very thin, often transparent and nearly Impreceptlble* anterior lobe indistinct* Umbilicus open as a very narrow slit at base of umbilical callus, rarely closed* more widely open in younger shells, but never conspicuously open. Umbilical callus thin, narrow, applied as a narrow semicircle upon underlying whorl, scarcely broader than Inner lip In young Individuals* 191 Basal lip thin. Operculum.--ChitInoua. very thin, filling aperture. Specimens Examinedi 60. Geographic Occurrence and Ecologyi Circumboreal. In the eastern Pacific, south to Atlca Island and Unalaska In the Aleutians and the Shumagin Islands (55°10'N, 160°W)| In the western Pacific, south to Xwate Prefecture, Japani in the western Atlantic, south to Massachusetts Bay» in the eastern Atlantic, south to the Lofoten Islands, Nor way. Known on sand and mud bottoms in depths of 37 to 317 meters in the eastern Pacific, and in depths to 600 meters In the northern Atlantic (Odhner, 1913). Stratlgraohic Occurrencei Reported as a "fossil in Eng land and northern Russia" by Frlele & Grieg (1901), but without further details, and from the Pleistocene of Ardlncaple, Scotland, and Iceland by Harmer (1921). Natica fragllls Conrad, £. fragills Smith, and f i . aperta Lea, described from Neogene deposits of eastern America and frigland, would provide further fossil records If they could be verified as Junior synonyms of £• fragllls. Type Localitiesi Natica fraallls - Baffin Bay, between Greenland and Canada (Leach, 1819). 192 Natica flava - "stomachs of fishes" (Gould, 1839). Bulbua flavua elonaatus - Nemuro, Hokkaido, Japan (Habe & Ito, 1965). Bulbua snlthll - Ardlncaple, near Helensburgh, south western Scotland (Brown, 1839). Natica aperta - FInnark, Norway (Loven, 1847). IYB9 Natica fraallla - Unknown, presumably In BM(NH) (Dance, 1966). Natica flava - Unknown, presumably lost (Johnson, 1964). Bulbua flavua elomnatus - Holotype, National Science Museum, Tokyo (Habe & Ito, 1965). Bulbua amlthll - Unknown, presumably In Manchester Museum, Manchester, England (Dance, 1966). Nomenclature! Commentary i There Is a confusing array of northern Atlantic species that may or may not be identical with £. amlthll. These species generally have poor litera ture citations, are rare or absent In American collections, and have type specimens unavailable to me. Included as possible synonyms are Natica tenulcula Sowerby, f g . Klaclalla Daniellsen, and perhaps £. tenulatrlata Dautzen- berg & Fischer, 1911. Settling of their taxonomic rela tionships must await study of type specimens in European collections• 193 DiscussIoni Variations In the relative height and thick ness of shells have resulted In the naming of species that are distinct only when considered alone. The range In morphology Is from an elongate* higher-spired form with a thin shell* to a more globose, lower-spired form with a thicker shell. The lower spired form also has a slight flattening of the shoulder and a stronger deflection of the Inner lip by the penultimate whorl. However, these two extremes Intergrade* with no apparent geographic separation, and collecting data are not precise enough to demonstrate ecological distinctions. I have not made a radular preparation for lack of suitable material* but Odhner (1913), referring to Acrvbia flava (Gould), stated that It has a monocuspate rachidlan. If true, this would distinguish B. fragllls. and presumably all Bulbua species, from most other naticlds. In the eastern Pacific, only the two Chorlstes species have monocuspate rachidians. Thorson (1951) figures a monocuspate rachidlan and lateral teeth In the original description of £. alaclalis. so the monocuspate rachidlan may be a general feature of Bulbus. It la unclear from Brown's (1839) original descrip tion whether the type of £• amlthll la a Pleistocene or Recent shell. Harmer (1921) seems to Indicate that It Is a fossil. 194 Genus Chorlstes Carpenter, 1872 Chorlstes Carpenter In Dawson, 1872i392. Type-speciesi Chorlstes elesans Carpenter In Dawson, 1872, by monotypy. Late Cenozolc, Montreal, Canada, Figured by Carpenter In Dawson (1872, pi. 7, fig. 13, 13a). Diagnosisi Shells small to medium In size, globose, thlnj whorls moderately Inflated, sculptured with weak spiral costellaet suture may be narrowly channeled. Umbilicus narrow to relatively broad, simple. Inner lip slightly thickened, simple, lacking umbilical callus. Parietal callus thin to average. Operculum chltlnous, entirely filling aperture. Radula with monocuspate rachidlan, one monocuspate lateral tooth, and two monocuspate marginal teeth per half-row. Discussioni Chorlstes Is characterized by Its thin shell, rounded base, and distinctly open (not silt-like) umbilicus, and by the lack of a distinct umbilical callus. It Is found living only in bathyal and abyssal depths, and Is known from one Atlantic and two Pacific species* The radulae of the two Chorlstes species I have examined both have monocuspate rachldlans, one monocuspate lateral tooth, and two monocuspate marginal teeth per half-row. This differs clearly from the usual natlcld pattern of a trieuspate rachidlan, one blcuspate lateral 195 tooth« and two monocuspate laterals per half-row. Choxiatea carpenter! Dali. 1896 Plate 2, figure 12 Chorlstes carpenter! Dali* 1896t10-11, Dali, 1908i328-329, pi. 3, fig. 4) Keen, 1971t388, fig. 424. PWflPtVWM Color.--Shell whitish. Ferlostracum thin, pale green ish brown. Size.--Average (based on 4 specimens)i height 20 mm, diameter 20 runj largest specimen (holotype)i height 20.5 mm, diameter 20.4 mm. Shell Form.--Shell somewhat elongate, spire moderately elevated) body whorl not greatly inflated! suture channel ed) shell thini whorls more than apices eroded on all specimens . Spiral sculpture of minute, weakly developed, obscure costellaei axial sculpture of incremental growth lines. Parietal callus thin, not thickening Into posterior apertural angle) anterior lobe lacking. Umbilicus open, narrow to relatively broad. Inner lip slightly thickened, simple, lacking umbilical callus. Basal lip not thickened. Operculum.--Chitlnoua. filling aperture) margin of penultimate whorl extends beyond Inner edge of operculum. 196 Specimens Examined* 4. Geographic Occurrence and EcoIqrvi Known from the Gulf of Panama (about 13°N), from off southernmost Mexico (14° 46'N), and from off central Oregon (44°41, N* 125°37, W), The Oregon record (1ACM) la baaed on two recently col lected specimens* one alive. Reported In depths of 2693 meters (live) and 3291 meters (dead) In the Gulf of Panama* 3436 meters (dead) off of southern Mexico* and 2800 meters off of Oregon (live)* on substrates of mud and "ooze." Tvne Localityi USFC station 3382* Gulf of Panama* 2693 meters depth* mud (Dali* 1896). Tvoe Materiali Holotype* USNM 123039. DiscussIoni The combination of clearly open umbilicus* thin shell* and channeled suture characterizes this species. A detailed comparison with Chorlstes n. sp. Is given later. The radula was removed from an Oregon specimen* and the rachidlan was found to be monocuspate rather than trlcuspate as In most natlclds. There Is a large central cusp* as In most Follnlclnae and Natlclanae* but there are no smaller cusps flanking It on either side. In ad dition* the Inner marginal tooth Is also monocuspate* whereas that of nearly every other eastern Pacific species 197 Is blcuspate. These features also occur In Chorlstes n. sp., which reinforces the generic allocation of these two species and further distinguishes them from other naticids. Chorlstes new species Plate 2, figure 13 Description of Holotypei Color.-*Shell exterior and interior whitish. Perio- stracum thin, pale yellow to brownish white. Shell Form.--Shell globose, spire moderately elevatedi body whorl not greatly Inflated, narrowly flattened to slightly concave Just below suture shell thickness averagei whorls 4%, nuclear whorls not clearly dif ferentiated! earliest nuclear whorl sunk into succeeding whorli microscopic radial wrinkles occur discontinuously below suture of nuclear whorlsi suture slightly im pressed. Spiral sculpture of very faint, minute, irregu larly spaced costellae,slightly stronger near suturei axial sculpture of Incremental growth lines. Parietal callus of average thickness, moderately filling posterior apertural anglei anterior lobe weak. Umblllcua open, small. Inner lip thickened,lacking distinct umbilical side, and expanded posteriorly to match width of perietal callus. Anterior inner lip and basal lip slightly 198 thickened. Dimensions! height 13.9 mm, diameter 13.0 mm. Operculum.--Chltlnous. filling aperture. Radula.--Each row of teeth consists of a rachidlan flanked on each side by one lateral and two marginals. Rachidlan with a single anterior cusp and prominent basal processes. Lateral tooth elongate* with one strong anterior cusp and smaller basal process on Inner end. Inner and outer marginal teeth simple* evenly curving, lacking denticles. Tvoe Localityi Off central Oregon at 2853 meters depth. Collected by Department of Oceanography, Oregon State University* R/V Yaoulna. sample OBT 132. Type Materiali Holotype, LACM. Discussioni Compared to £. carpenter!. £. n. sp. has a lower spire, a more narrowly and shallowly channeled suture, much more distinct spiral sculpture (especially near suture), a thicker shell, and a much heavier filling of parietal callus in the posterior apertural angle. Specimens of £• carpenterl were found off of Oregon along with £• n. sp., and were easily distinguished. The radula was removed from a paratype of £. n. sp. The radulae of £. n. sp. and £. caroenterl differ markedly, although they share features that distinguish them from 199 all ocher eastern Pacific natlclds. The rachidlan of each species Is monocuspate, but the rachidlan of £. n. sp. has a more quadrate shape. The basal processes of £. n. sp. project abruptly from each comer, whereas those of £, carpenterl are more massive and have less sinuous lateral margins. The lateral teeth of these two species are more similar than the rachldlans, but the laterals of £. n. sp. have more broadly rounded and prominent inner basal processes. The marginal teeth are simple and very similar between the species. As noted In the discussion of £. carpenter!. the monocuspate rachldlans and lateral teeth of these two species distinguish them from all other eastern Pacific natlclds. Subfamily Slninae Wenz, 1941 Slnlnae Wenz, 1941t1037. Diagnosisi Usually thin-shelled, auriform to nearly globose, rarely elongate) final whorl greatly enlarged) ornamented with spiral costellae) animal too large to retract Into shell) operculum cliltinous, rarely partly calcareous• Discussioni This subfamily is almost exclusively a tropi cal group, with only a few species ranging into temperate waters. A key to the two genera used here follows) 200 la. Shells auriform* extremely depressed to moderately elevated* umbilicus slit-like to closed* umbilical callus lacking .................. sinum lb. Shells ovate, not autiform, umbilicus distinctly open, umbilical callus slender .................. Eunatlclna j* Genus £< num Rodlng, 1798 Slnum Rodlng* 1798t14, Tvoe-speciesi Helix haliotoldea Linnaeus, 1758, by sub sequent designation (Dall( 1915i109), Recent* West Africa. Slaaretus Lamarck, 1799i77. Type-speciesi Helix haliotoldea Linnaeus, 1758, by mono- typy. Diagnosisi Shells small to largei whorls typically very depressed, may be globose, rarely elongatet sculptured with distinct, closely spaced spiral costellae. Umbilicus extremely narrow and slit-like or closed. Outer lip may be weakly crenulated. Operculum chltlnous, much smaller than aperture. Radula with trlcuspate rachidlan in which central tooth is often very small, one nultlcuspate lateral tooth, one bleuspate inner marginal tooth, and one 201 monocuspate outer marginal tooth per half-row. Discussloni Slnun is characterized in most cases by an extremely depressed, auriform (ear-shaped) shell with a slit-like or closed umbilicus. A few species have rela tively elevated, nearly globose shells. Only the extinct species treated here, Sinum perrini (Arnold), has an ex tremely elongate shell. Sinum cvmba (Menke, 1828) Plate 2, figures 14, 15, 16 Slaaretus cvmba Menke, 1828188, Menke, 1830*87» Orblgny, 1840i403, pi. 57, figs. 3-6 (plate 1837)| Troschel, 1852i161* Hupe, 1854t225-226i Philippi, 1860t186t Weinkauff, 1883i10-12, pi. A, figs. 1, 2, 3, pi. 1, figs. 10-12. Slaaret^a cvmba var. alba Weinkauff, 1883ipl. 3, figs. sinum cvmba (Menke), Keen, 1971i482, fig, 889. Slaaretus Philippi, 18441vol. 1, p. 143, Slaaretus. pi. 1, fig. 1* Philippi, I86O1I86 Cas synonym of 5 . cvmba MenkeJ* "Slaaretus concavus Lamarck" fnot 5 . concavus Lamarck, 18221 Recent, West Africa*1 Reeve, 18641pi. 1, figs. 3a,bi Sowerby, 1882*39-40, pi. 1, figs. 8-11, pi. 2, figs. 18, 19i Tryon, 1886(55, pi. 23, figs. 36, 37 Cnot pi. 22, fig. 34, haliotoldea Lin naeus , 1758, ?-5. concavus Lamarck, 15221 Recent, West Afrlcaji Dali, l$09ai2361 Dali, 1909b11751 Burch & Burch, 1964(109, pi. 5, figs. 2, 4. 202 "Sinum cgncavuip (Lamarck)" ^not S. concavus Lamarck, I822]i I. S. Oldroyd, I9l7*l3. Descriptioni Color.--Protoconch pale yellowish brownj early adult whorls dark brownish grayj last one to two whorls pale yellowish brown to purplish brownt base white. Narrow white band occurs below suture of postnuclear whorls. Ferlostracum thin, pale yellow, thickening toward growing margin of large adults. Interior chestnut browni internal peripheral ridge chocolate browni callus and margin of aperture white. Form.--Average sizet height 35 mm, diameter 40 mmj largest specimeni height 50.8 mm, diameter 59.4 mm [AM4H 155878, Caldera, Chile]. Spire moderately elevated, shell high for the genust shoulder slightly flattened) shell thick for the genusi nuclear whorls 2%, with indis tinct spiral costellaei postnuclear whorls 3i suture slightly impressed and narrowly but distinctly channeled. Interior with low ridge along periphery. Spiral sculp ture of flat-topped costae separated by much narrower interspaces, both of which may bear minute costellaei axial sculpture of Incremental growth lines that disrupt the spiral costae and give them a wavy appearance. Parietal callus thin, transparent In young specimens, lightly filling posterior apertural anglei anterior lobe Indistinct. Umbilicus open, inconspicuous, slit-like. 203 Umbilical callus narrow, flattened, conceals umbilicus. Inner lip evenly roundedi basal lip not thickenedi outer lip weakly crenulate. £B£££iLLyffl.--Chitlnous, much smaller than aperture. Specimens Examinedi 351. Geographic Occurrence and Ecology! Galapagos Islands and Manta, Ecuador (1°S) to Caldera, Chile (27°s), and pos sibly farther south. Single adult specimens are present in collections from Nicaragua, and from Acapulco, Sinaloa, and Baja California, Mexico, but these localities are probably in error. The southern range-limit is even less certain. Two MCZ specimens are reportedly from the Magellanic region at "Port Gallant" and "Possession Bay," but these have not been located. Hupe's (1854) record from Valparaiso (33°S) has not been confirmed by modem collecting. Based on sparse collecting data, this species occurs alive in 24 to 100 meters depth and has been found abundantly in 91-100 meters off of northern Peru (LACM collection). It is not reported in shallow water from southern Chile by Dali (1971) or from northern Chile by Marincovich (1973). Because this species is fairly well represented in collections, its habitat probably ranges into shallow water, but ap parently not into the intertidal. 204 Stratlaraphlc Occurrencei Reported from "Cenozolc" beds on Seymour Island, Galapagos, by Dali & Ochsner (1928), and represented by CAS collections by one specimen from the "Pliocene" of Indefatigable Island, Galapagos. It Is not reported from the Pliocene or Pleistocene of northern or central Chile by Herm (1969). Type Localitiesi Slaaretus cvmba - "In oceano peruviana" (Menke, 1828). Slaaretus cvmba var. alba - Unknown (Weinkauff, 1883). Slaaretus maxlaus - Peru (Philippi, 1844). lYPfi Material, Slaaretus cvmba - Unknown, presumably sold to a dealer and lost (Zilch, 1958i53). Slaaretus cvmba var. alba - Unknown. Slaaretus maxlmus - Unknown, presumably In Museo Naclonal de Historla Natural, Santiago, Chile (Dance, 1966). Nomenclatural Commentaryi The West African species £. pnnpaviin is similar to £• cvmba and has been widely con fused with it until recently. There are at least four named species In West Africa that are similar to £• con- and that have been considered synonymous In various combinations by earlier workers. 205 Discussioni This species has the largest and most solid shell of the eastern Pacific Sinums and is also the most colorful. It is most similar to £. sconulosum (Conrad) but is distinguished by Its darker color, thicker shell, narrower spiral interspaces, higher spire with more steeply-sloping whorl profile, and more broadly rounded Inner lip. The spiral costae of £. cvmba also maintain their integrity better than do those of £. scopulosum. tending less to become subdivided into costellae. Some specimens of £* cvmba may also appear close to £. aravl (Deshayes), but the latter species differs by having lighter-colored early whorls, a proportionately higher and thicker shell, angulate inner lip, smaller size, wider spiral interspaces, and usually a brown margin within the aperture. Sinum cvmba and £. concavum of West Africa are closely related. Individuals of each species are similar in size and whorl profile. The principal differences are that £• concavum has lighter-colored early whorls and completely lacks a white band below the suture, has a paler brown Interior, narrowed and wavier spiral costae, and has Interspaces equal to or greater in width to the costae. It would be Interesting to compare fossils of each species to see If the differences are less than those between Re cent shells. Slnun Rravl (Deahayes, 1843) 206 Plate 2t figures 18, 19, 20, 21 Slaaretus Rravl Deshayes In Lamarck, 1843ivol. 9, p. 12t Troschel, 18521161 [[as synonym of £• cvmba Menke, 1828], ‘ tVVHI aravl (Deshayes), Shasky, 1961123, pi. 4, figs. 13, 14t Keen, 1971>482, fig. 891. Sinum cortez1 Burch & Burch, 1964i109-110, pi. 5, figs. i ,37 Descriptioni Color.--Shell tan to medium brown, with a narrow white band Just below the suturei vague axial banding of lighter and darker colors may occur i base whitei Interior and callus white, usually with light brown margin to outer lip. Periostracum thin, pale yellowish brown. Form.-‘Average sizei height 25 mm, diameter 27 mmi largest specimen! height 36.6 mm, diameter 34,8 mm [LACM, uncataloged, off northern Peru in 160 mj. Shell globose, spire low to moderately elevated, shell high for the genust whorls evenly roundedi shell moderately thicki nuclear whorls 2fc, with microscopic spiral costellaei postnuclear whorls 3t suture slightly Impressed. Spiral sculpture of flat-topped costae separated by interspaces of about equal or greater wldthi Interspaces usually with one or more well-developed costellaei axial sculpture of incremental growth lines that may disrupt the spiral 207 costae and give them a slightly wavy appearance. Parietal callus thin, transparent In young specimens, moderately filling posterior apertural anglet anterior lobe Indis tinct. Umbilicus open, Inconspicuous, slit-like. Umbilical callus narrow, flattened, conceals umbilicus. Inner lip slightly angulate, not evenly roundedi basal lip thickenedi outer lip weakly crenulate. Operculum.--Chitlnous. much smaller than aperture. Geographic Occurrence and Ecology Guaymas, Mexico (28°N) to near Lambayeque, Peru (6°27*S). Except at the range end-points, reported only from Panama Bay (Keen, 1971) and the Golfo de Fonseca, El Salvador (Shasky, 1961). Found in depths of 25 to 160 meters. Rare throughout its range except off northern Peru, where about 300 LACM specimens were collected in 160 meters depth. Type Localitiesi Sl&aretua firflyj - Unknown. cortez1 - MShrimp trawlers working between Mazatlan and Altata [Sinaloa, west Mexico^ in 15 fathoms'1 (Burch & Burch, 1964). Type Material! Slaaretus aravl - Unknown, presumably in BM(NH) or Ecole de Mines, Paris (Dance, 1966). Sinum cortez1 - Holotype* CAS 12601. 208 Discussion* Sinum grayl has the most globose and pro portionately high shell of the living eastern Pacific Sinums. Its shape, thickened basal lip and light interior coloration distinguish it in most cases from £. cvmba. and other differences are noted in the discussion of the latter species, sinum cvmba may range into shallower depths than £. Rravl■ although the habitats of these species are imprecisely known. Sinum «pnnii1n«nni (Conrad* 1849) Plate 2* figures 22* 23* 24, 25 Slaaretus scoduIqsus Conrad, 1849(appendix p. 727* pi. 19, figs. 6, 6a [not figs. 6b,c, ■Natica oregcnensls (Con rad, 1849)i not fig. 6d, ■ not identifiableJt Meek, 1864i31i Dali, 1892i380t Dali in Diller, 1896*474) Anderson, 1905(188, 203, pi. 16, figs. 72, 73) Arnold, 1906(17, 19) Arnold, 1908*350) Dali, 1909b(125) Reagen, 1909*194-195, pi. 3, fig. 30) Reagen, 1910* 648i Arnold, 1910■154, pi. 24, fig. 1) Arnold 6 * Anderson, 1910(127, pi, 46, fig. 1) Anderson, 1911* 100) J. P. Smith, 1912*167, 176) Anderson & Martin, 1914*43) Keen & Bents cm, 1944*192. Catin^s acopuloaua (Conrad), Meek, 1864*31) Conrad, 1865* scopulosa (Conrad), Meek, 1864131. siflmi gypnyilqauw (Conrad), Meek, 1864i31) Gabb, 1869*114) Merriam, 1896*103, 106) Dali, 1909b*12, 18, 91-92, pi. 4, fig. 10, pi. 5, fig. 8) Arnold A Hannibal, 1913(584, 588) fingllsh, 1914(211) Clark, 1915(15) Weaver, 1916*172, 176, 216, 218) Nomland, 1917ai213, 221) Nomland, 19l7b*30l) Clark, 1918(169-170, pi. 22, fig. 8) Trask, 1922*153-154) Stewart, 1927*327-328, pi. 32, fig. 4) Grant A Gale, 1931(806) Etherington, 209 * fig. 13* Loel & Corey* 1932>270* ,3a,b* Tegland, 1933*140, pi, 14* 1931*95-96, pi 349-350, 1945i57i Smith Burch •v r * * 7 * flgs. 6 * S i n u m n | a n l o n g t ! u iii G a b b * 1 8 6 9 i4 9 - 5 0 * 7 8 * p i . 1 4 * f i f t . 6 | D a l i , l v M b *91-92 f a s J u n i o r s y n o n y m o f bcopu- l o a u m l . S l a a r e t u s p ! a n i e n s t u n ( G a b b ) , C o o p e r i n W a t t s * 1 8 9 6 t 8 1 1 A n d e r s o n , 1 9 0 51204 . filn V lT 7 * S . O l d r o y d , 1 9 1 7 *1 3 * D a l i * 1 9 2 1 i 1 6 51 T . S . O l d r o y d , 1 9 2 411 9 * I . S . O l d r o y d , 1 9 2 7 *7 3 2 , p i . 9 2 * f i g s . 13 - 14 * S t e w a r t * 1927*327 f a s J u n i o r s y n o n y m o f £ . s c o p u l o s u a l . S i n u m k e r a t l u m D a l i * 1 9 1 9 *3 5 4 * D a l i , 1921 *1 6 5 * 1 . S . O l d r o y d * 1 9 2 7 *733 * K e e n , 1 9 3 7 *4 5 * B u r c h * 1946 *3 2 . ( S l f i M S t l l l ) t r i a e n a r l u m T r a s k , 1 922 *153 - 1 5 4 , p i . 7 , f i g s . 2a , b * S t e w a r t , 1927*327 C a s J u n i o r s y n o n y m o f £ • ffffftP V llftfftifflT K e e n & B e n t5 o n , 1944 *1 9 3 . " S l a a r e t u s d e b l l i s G o u l d " C n o t S . d g b l l g G o u l d , 18531 * K e e p , 1 888 *4 7 * A r n o l d , 1 9 0 3 *3 1 6 * A r n o l d , 1 906 *2 8 , 3 6 * A r n o l d , 1908 b *4 2 3 . ■ s i m p d e b l i e ( G o u l d ) " C n o t 3 . d e b l i e G o u l d , 1853 J * V a l e n t i n e , 1 9 5 6 *2 0 0 . P«srlBfcUa« Color.--Shell exterior and Interior white. Perlo st racum thin, pale yellowish white, may have dark rust 210 brown staining above basei base white. Site. - - Average i height 21 nun, diameter 26 ntmi largest Recent speciment height 31.2 mm, diameter 38.2 mm CAS, Newport Bay, California j largest fossil specimeni height 33.0 mm, diameter 35.0 mm £CAS 1, Empire Formation, Coos Bay, Oregon, lower Pliocene^. Shell Form.--Shell high, spire moderately elevatedi body whorl Inflated and laterally expandedi shoulder broadly convex, base flattenedi shell thini nuclear whorls 2 smooth, postnuclear whorls 2* suture slightly impressed. Spiral sculpture of low, flat-topped costae separated by Interspaces of equal or lesser widthi costae usu \y bear up to 5 weak to well-developed costellaei Interspaces usually lack costellae, may bear 1 weak onei axial sculpture of incremental growth lines. Parietal callus thin, transparent, lightly filling posterior apertural anglei anterior lobe indistinct. Umbilicus open, inconspicuous, slit-like. Umbilical callus narrow, flattened, conceals umbilicus. Inner lip angulate, thickenedi basal lip not thickenedt outer lip smooth to weakly crenulate. Operculum.--Chitlnous, much smaller than aperture. Specimens 427. 211 Geographic Occurrence and Ecologyi Monterey, California (36°30’N) to Bahia Iortolo, western Baja California, Mexico (27°39*N), including the southern California is lands. Principally living in undisturbed sandy and muddy substrates nearshore, in depths of 15 to 171 meters, and In shallow embaymentsi reportedly found at low water oc casionally in bays (McLean, 1969). Its shallowest known occurrence at the southern limit of its range (which is also the southern limit of the Californian molluscan province) is 48 to 60 meters, indicating its aversion to warm southern surface water. Stratlaraphic Occurrencei Ranges from Upper Oligocene to Recent. Upper Oligocenei Twin River Formation CEchlno- phorla rex zone3, northern Olympic Peninsula, Washington (Durham, 1944)j type Blakeley Formation of Weaver (1912), Puget Sound, Washington (Tegland, 1933). Lower Miocene (7)i Twin River Formation, northern Olympic Peninsula, Washington (Arnold A Hannibal, 1913t Durham, 1944 CEchlno- phorla rex zone])t San Ramon Sandstone, Contra Costa County, California (Clark, 1918t Weaver, 1953). Lower Miocenei Vaqueros Formation, northern Santa Lucia Range, La Panza Range, western Santa Ynez Range, Ventura basin (Loel & Corey, 1932)i Callente Range (Eaton & others, I941)i Jewett Sand, Kern River area, California (Addicott, 1970). Middle Miocenei Clallam Formation, northern Olympic 212 Peninsula, Washington (Reagan, 1909, 1910i Arnold A Han nibal, 1913| Durham, 1944)i Astoria Formation, south western Washington (Etherington, 1931), coastal Oregon (Conrad, 18491 Moore, 1963)i Monterey Shale, Cameros Creek, Napa County, California (Weaver, 1949)i Monterey Group, western Contra Costa County (Weaver, 1949)i Temblor Formation, Vallecitos area (Schenck & Keen, 1940), Reef Ridge (Stewart, 1946, cf.), La Panza Range (Loel & Corey, 1932)i Saltos Shale Member of the Monterey Shale of Hill, Carlson and Dlbblee (1958), Caliente Range (J. G. Vedder, fide Addicott, 1970)i Topanga Formation, Santa Monica Mountains (Susuki, 1951), Santa Ana Mountains (Vedder & Woodrlng, unpub., fide Addicott, 1970)i Upper Olcese Sand and lower Round Mountain Silt, Kern River area, California (Addicott, 1970). Upper Miocenei Monte- sano Formation of Weaver (1912), southwestern Washington (Weaver, 1916| Ehterlngton, 1931)i Santa Margarita Forma tion, Coalings area (Nomland, 1917a), Comanche Point (Addicott, 1970)i Briones Formation, Solano County, Cali fornia (Trask, 1922)i Castaic Formation of Crowell (1955), eastern Ventura basin (Stanton, 1966, 1967). Pliocenei Empire Formation, southwestern Oregon (Dali, 1909i Weaver, 1942, 1945i CAS) j Pancho Rico Formatlorn, Salinas Valley (Durham & Addicott, 1965t USGS)i Etchegoin Forma tion, Coallnga area (Nomland, 19l7bi Adegoke, 1969), Careaga Sandstone, Santa Marla basin (Woodrlng & Bramlette, 213 1950)i Pico Formation, eastern Ventura basin (Gabb, 1869j Kew, 19241 Grant & Gale, 19311 Woodrlng and others in Winterer and Durham, 1962), central Los Angeles County (LACM)i Fernando Formation, Puente Hills (Vedder In Dur ham & Yerkes, 1964, cf.). Lower Pleistocenei San Pedro Formation, San Pedro, California (Oldroyd, 1925). Upper Pleistocenei Palos Verdes Sand, Palos Verdes Hills, ter race 12 (Marlncovlch, 1970), Playa del Rey (Willett, 1937), Potrero Canyon (Valentine, 1956), Newport Bay area (Kana- koff & t Emerson, 1959. Type Localities» Slaaretus scooulosus - Astoria, Oregon (Conrad, 1849) ^presumably from the Astoria Formation, middle Miocene]. Sinum planicostum - Pliocene of San Fernando (Gabb, 1869)i Pico Formation, Ventura basin, California, Pliocene (Addicott, 1970). . < 54mm TflllfVFJilEM” - San Pedro, California (Oldroyd, 1917). * 54mils " Catalina Island, California (Dali, 1919). SlJam (si&aretus) trlaenarlua - UCB locality 3576, Mare Island quadrangle, near San Pablo Bay, Solano County, California, upper Miocene (Trask, I922i Keen & Bentson, 1944). 214 Tvoe Materiali SUaretus scopulosus - USNM 3553, lectotype (desig nated by Moore, 1963). Slnum planlcostum - ANSP 4326, lectotype (designated by Stewart, 1927). Slnun callfomicum - SU 6446, paleontology type col lection, holotype. Slnun keratlun - USNM 206152, holotype, Slnun (SUaretua) trl&enarlun - UCB 12386, holotype. Nonenclatural Connentaryi The lectotype of £. planlcostun is badly worn, with only vestiges of shell material show ing spiral sculpture identical to that of £. nmim. Slnun callfomlcun was compared in its type description to 5. deblle (Gould) and "£. concavum" - cvnba (Menke), not concavum (Lamarck) , but not to £. scopuloaum. which even then was well known at the type locality of call fomlcun. The holotype of 5. keratlun is a Juvenile scopuloaum. with the sculpture and Juvenile proportions of this species. The holotype of £• triaenarlun is a cast without shell material and has been abraded and deformed during deposition. Deformation has altered the proportions of the shell, to make the base appear more flattened than usual for the genus. Spacing of the spiral costae Is the same as on j, ffgfrgUlgfl'JB- but costellae are not preserved. Trask (1922) stated that £. firlfigflfTTimi bas 30 spiral 215 costae on Its body whorl, compared to 45 for £• scopuloaum. The poorly preserved spiral costae of Trask's holotype are difficult to accurately count* but at least 30 are pre sent* with others apparently missing, whereas Recent and Neogene individuals of £. scopuloaum have 35 to 50 costae on their body whorls* Trask's species is thus not separable from £. scopuloaum. Discussioni Individuals of S. scopulosum vary noticeably in inflation of the body whorl, proportionate height of the shell, and the degree to which the body whorl is angled downward to give the shell an "oblique" look. These variations have been partly responsible for the nam ing of synonymous species. Relatively flattened shells from the southern California Pleistocene have sometimes been referred erroneously to £. debile (Gould), but I have seen no specimens of that species in the several extensive Pleistocene collections that are in West Coast institu tions. Neogene specimens show the same range of variation in shell characters as to Recent ones. Some Miocene specimens have thickened shells but are otherwise typical. The largest known fossils are about the same size as the largest Recent specimens, but with slightly higher shells (see "Size" heading above). &QUB debile (Gould, 1853) 216 Plate 2, figures 27, 28, 29 Slaaretus debllls Gould, 1853(379-380, pi. 14, fig. 17, Carpenter, 1857ai207i Tryon, 1886i57, pi. 24, fig. 65i Keep, 1888>471 Steams, 189411961 Arnold, 1903(316 scopuloaum (Conrad, 1849)3i Johnson, 1964(66. Sinum debile (Gould), Dali, 1921(65* Oldroyd, 1927(733, pi. 95, figs. 3, 7i Grant & Gale, 1931(806-807| M. Smith, 1944(13, fig. 136* Keen, 1958(324, fig. 275i Parker, 1964(153, pi. 4, fig. 6* Keen, 1971(482, fig. 890. Sinum oazianum Dali, 1919(354i Dali, 1921(165i Oldroyd, 1957(734, pi. 92, fig. 10* Keen, 1937(45* M. Smith, 1944(13i Burch, 1946(32* Keen, 1958(325, fig. 277. Descriptioni Color.--Shell white, covered with thin yellowish white periostracumi callus and interior white. Form.--Average sizei height 9 mm, diameter 20 mmj largest specimen( height 12.0 mm, diameter 29.0 mm ftJSNM 46554, Gulf of California]. Apex strongly depressed, shell outline very lowi shell thin* nuclear whorls 2, with microscopic spiral costellaei postnuclear whorls 2^j suture very slightly impressed. Spiral sculpture of flat- topped costae separated by sharply incised interspaces of equal or greater widthi inconspicuous costallae may be present in interspaces! axial sculpture of incremental growth lines that often disrupt the spiral costae and give them a wavy appearance. Parietal callus extremely thin, transparent, only slightly filling posterior apertural 217 anglei anterior lobe broad. Umbilicus open, incon spicuous, slit-like. Umbilical callus narrow, flattened, conceans umbilicus. Interior of outer lip weakly crenu- lated. Operculum.--Chitinous, much smaller than aperture. Geographic Occurrence and Ecology« La Paz, Baja,Cali fornia, Mexico (24°N) to Panama Bay, Panama (8°N)i most common throughout the Gulf of California, rare elsewhere. Dali (1919) and some later workers listed £. debile from Catalina Island, California (33°N), but specimens so named from that locality are actually £• ffWPVlflftVW (Con rad). One specimen from Salinas, Ecuador [CAS, no number j, is close to £. debile but too worn to identify with cer tainty. Living intertidally on sand and mud, dead shells found to 73 meters. Stratlaraphlc Occurrencei Reported from upper Pleistocene deposits at San Pedro and Long Beach, California (Arnold, 1903), and from the Pliocene in a well at San Diego, Cali fornia (Cooper, fide Arnold, 1903). These records are based on specimens of £. «r>ontiin*nw (Conrad), the only recognized sinua species in Neogene deposits of these areas. 218 Slaaretus debllla - La Paz, Baja California, Mexico (Gould, 1853). Slnun na7l*n’ H - Off La Paz, Baja California, Mexico, USFC station 2823, 26*f fathoms (Dali, 1919). Type Materiali Sl&aretua debllla - Holotype, MCZ 169117. <amu pazlanun - Holotype, USNM 211406. Dlscusslom This species has the most flattened outline among the eastern Pacific Sinums. Occasional specimens have shells that are much thicker than average for the genus. Although faint costellae may occur in the spiral interspaces, they are absent on most specimens. Sinum sanctilohannla (Pilsbry & Lowe) is similar to this species, but has a higher shell, occasional rust-colored stains on the perlostracum, and more frequent and stronger costellae In its spiral interspaces. The morphology of the Carib bean species £• (Say, 1831) is identical to that of £. debile, although the former species attains a greater maximum size (to a diameter of at least 36 mm). 219 Slnun novesll Dali, 1903 Plate 2, figures 30, 31, 32 simiM noveall Dali, 1903t37, M. Smith, 1944i13i Keen, 1958i 324, ftg. 276j Keen, 197li482, fig. 892. Jte2££U&l2Q' Color.--Earlieat part of protoconch dark brown, other wise whitei early poatnuclear whorls white with two spiral rows of pale brown spots above periphery, which become darker and spread to color entire upper half of final adult whorl mahogany browni interior repeats exterior color pattern through semi-transparent shelli callus white. Perlostracum thin, pale yellowish brown. Form.--Average size* height 13 mm, diameter 32 mmi largest specimen* height 42.1 mm, diameter 21.7 mm [USNM 622799, Isla Tortola, Panama Bayl. Shell flattened, spire low* shoulder flat or slightly convexi shell thlni nuclear whorls 2, with microscopic spiral costellaei postnuclear whorls 2^i suture concealed by perlostracum, lies within narrow channel of uneven depth formed by overlapping of earlier whorl by later whorl. Spiral sculpture of low, narrow, very weak costae separated by Interspaces of lesser to much greater (to about 6 times) wldthi interspaces with several indistinct costellaei axial sculpture of Incremental growth lines that disrupt 220 the spiral sculpture and give it a minutely wavy appear ance. Parietal callus thin, transparent, lightly filling posterior apertural anglei anterior lobe indistinct. Umbilicus closed. Umbilical callus not distinct, reduced to an anterior extension of parietal callus. Inner lip and basal lip slightly thickened, evenly rounded) outer lip may be very weakly crenulate in Juveniles, not in adults. Operculum.--Chitinous. much smaller than aperture. Geographic Occurrence and Ecologyi Isla San Benito, outer coast of Baja California, Mexico (28°20'N, 115°30'W) to Isla Gorgona, Colombia (3°N, 78°19'W), Between these range end-points, specimens have been collected only at Isla Esp£ritu Santo, Gulf of California, Mexico (24°30'N, ll0o20'W), at San Juan del Sur, Nicaragua (11°15'N), and in the Gulf of Panama. Reported living intertldally at San Juan del Sur (Lowe, 1932) in a substrate of small stones and sandy mud, and found elsewhere living in depths to 89 meters. Type Locality) Isla Gorgona, Colombia C3°N 1st., 78°19'W long] (Dali, 1903). Tvoe Material! Holotype* USNM 170298. 221 Discussion* Slnun noveali Is easily distinguished fron other eastern Pacific Slnuns by Its closed umbilicus. The other two extremely flattened species* £• debLie (Gould) and £. sanctljphannia (Pllsbry & Lowe), also lack the dark brown color of £. noveali. Siam 02XSAU 4fibUS have been found living together In the Bay of Panama. simia sanctllot^aifnig (Pllsbry & Lowe* 1932) Plate 2, figures 33, 34, 35 Slaaretus sanctllohannls Pllsbry & Lowe, 1932i84, pi. 9, fig- 7. Slnun lohannls (Pllsbry & Lowe), M. Smith* 1944t13, fIg. 135i KeenT 1958i325, fig, 278i Keen, 1071*482, fig* 893. Description* Color.--Shell exterior and Interior white. Perio- stracum thin, pale yellowish white, rust brown staining at periphery and above. Size.--Averaxe * height 11 on, diameter 21 nmi larg est specimen* height 15.1 mm, diameter 25.9 mm [LACM, no number, 160 meters depth off northern PeruJ• Shell Form.— Shell flattened, spire low* shoulder slightly concave Just below suture, otherwise broadly con vex i shell thlni nuclear whorls 3, with cloaely spaced 222 microscopic spiral lines* postnuclear whorls 2i suture slightly Impressed. Spiral sculpture of low, flat-topped costae separated by Interspaces of equal or greater widthi interspaces usually with one costella, may have four or moret axial sculpture of incremental growth lines that disrupt the spiral sculpture and gives it minutely wavy appearance. Parietal callus thin, transparent, lightly filling posterior apertural anglej anterior lobe indis tinct. Umbilicus open, inconspicuous, slit-like. Umbilical callus narrow, flattened, conceals umbilicus. Inner lip evenly rounded, basal lip very slightly thicken ed t outer lip smooth to weakly crenulate* Operculum.--Chitinous. much smaller than aperture. Geographic Occurrence and Ecology» Isla San Benito [AHF 1250-41], outer coast of Baja California, Mexico <28°20*N, ll5o30*W), to off northern Peru (6027*S, 80°56'W), Found most commonly throughout the Gulf of California and off northern Peru, rarely between these two places* In depths of 18 to 165 meters In the Gulf of California, 27 to 73 meters off of southern Mexico, and 118 to 160 meters off of northern Peru (where it Is found with £. cvmba). Iy m faoctllty San Juan del Sur, Nicaragua (Pllsbry & Lowe, 1932). 223 Type Materiali ANSP 155436, holotype* height 16.5 mm, diameter 27.4 ora« Discussion. This species is most similar to S. debile (Gould), which differs in having a more flattened shell, broader and more persistent spiral costae, inconspicuous costellae in its Interspaces, and a perlostracum lacking rust-brown stains. The spiral costae of £. sanct1johennla may be so narrow that they are not distinct from the costellae, which never happens in £. deblie. Sinum perrinl (Arnold, 1907) Plate 2, figure 26 Siaaretua perrinl Arnold, I907a.532, pi. 51, fig. 5| Arnold, 196/bi228, pi. 28, fig. 5| Eldridge & Arnold, 1907i147i Grant A Gale, 1931i807f Keen & Bentson, 1944.192, Descriptioni Size.--Height 19,9 mm, diameter 10.1 mm (holotype). Shell Form.— Shell extremely high, spire greatly elevatedi body whorl axially elongate, anteriorly inflat ed) shoulder broadly convex, sloping steeply from suturei shell thini 2^ whorls preserved, apex misslngi suture strongly impressed for the genus, overhung by preceding whorl. Spiral sculpture of low, narrow and rounded costae separated by interspaces that are twice as wldei 224 Interspaces each bear one well-developed costellai axial sculpture of incremental growth lines. Parietal callus thint poorly exposed on holotype. Umbilicus concealed by matrix. Specimens Examinedi Holotype. StratlRraphlc Occurrencet Middle Miocenei Topanga Forma tion! Topanga Canyon, Los Angeles, California (Arnold, 1907a). Tvoe Localityi Head of Topanga Canyon, 3 miles south of Calabasas, western Los Angeles County, California (Arnold, 1907a). Camulos quadrangle, south % section 33T1N, R17W (Keen & Bentson, 1944). Topanga Formation, Middle Miocene. Tvoe Materiali USNM 164979, holotype. Discussioni This species has a more elevated shell than any other stmim species known to me. It is known only from the holotype, even though its type locality is heavily collected by professional and amateur paleontologists. Genus EtflfltlEUli Fischer, 1885 Eunatlclna Fischer, 1885i768. Type-speciesi Natlcina papilla (Gmelln, 1791), by original designation. Recent, western Pacific. Figured by Cernohorsky (1971 *201-202, fig. 69). 225 Diagnosisi Shells small to medium In size, globose to elongate* thin to average thicknessi whorls moderately In flated, sculptured with spiral costae and sharply Incised groovesi shoulder slightly flattened) suture narrowly and weakly channeled. Umbilicus broadly open. Umbilical callus slender, may be Indistinct from Inner lipi funicle low to indistinct. Farietal callus thin. Operculum partly calcified in some species, may possibly be entirely chitlnous or entirely calcareous in other species. Radula with multicuspate rachldian, one multlcuspate lateral tooth, one mono- or bicuspate inner marginal tooth, and one nonocuspate outer marginal tooth per half-row. DiscussIoni Eunatlclna Is characterized by Its ovate shape, broadly open umbilicus with slender callus, and sharply incised spiral grooves over the shell surface. It never has the depressed auriform shape of most Sinums. This genus is entirely tropical in distribution and has only a few species, perhaps as few as five or six. The multlcuspate rachldian of £. insculpta (Carpenter) is unique among eastern Pacific natlcids. 226 Eunatlclna Inaculota (Carpenter* 1865) Plate 2« figures 36* 37* 38 Narlca insculpta Carpenter* 1865*280) Carpenter* 1872* 273) Palmer, 1963*296. Eunatlclna he1ml Jordan In Hertleln* 1934*68* pi. 21* fig. 4) Jordan* 1936*161* pi. 19, fig. 6) Keen, 1958*322, fig. 265j Keen, 1971*477, fig. 872. Description* Color.--Shell exterior and Interior whltei nuclear whorls medium brown. Perlostracum very thin, pale yellow ish white. Size.--Average* height 13 mm* diameter 11.5 rami larg est specimen* height 14.3 mm, diameter 12.9 mm CAHF 1725-49, Cabeza Ballena* Baja California* MexlcoJ. Shell Form.--Shell globose, spire moderately elevated) body whorl Inflated) shoulder slightly flattened) shell thickness average, shell with solid appearance) nuclear whorls 2%, smooth) postnuclear whorls 3) suture narrowly channeled) Inner lip may be weakly crenulated. Spiral sculpture of flat-topped costae separated by much narrower* sharply Incised grooves) spiral costae are narrower near suture and on base, are sometimes subdivided Into poorly defined costellae) bottoms of the grooves are Irregular, sometimes with closely spaced shallow pits. Axial sculp ture of Incremental growth lines. Parietal callus thin, transparent, lightly filling posterior apertural angle) i n anterior lobe weak. Umbilicus widely open, exposing earlier whorls. Inner lip thickened, with a low forward projection at its midpoint, not forming a distinct umbilical callusi funicle low, very narrow, sometimes in distinct. Basal lip not thickened. Operculum. — Inner surface chltinousi outer surface Incompletely calcified, white, formed of radial rows of minute, coalesced beads, with occasional open Inter stices! outer calcareous layer thickest at inner part of final whorl, thinning and disappearing toward the upward- curved outer margin. Specimens Examined) 11. Geographic Occurrence and Ecologyi Cabeza Ballena, southernmost Baja California, Mexico (22054*N), in the southern Gulf of California, and south to Corlnto, Nicaragua (12°30*N), and the Galapagos Islands, Ecuador. Only one live-collected specimen with depth data is known, from Bahia de la Concepcion, eastern Baja California, In 22 meters. Stratlgraphic Occurrencei Pleistocenei Magdalena Bay, western Baja California, and Maria Madre Island, west Mexico (Jordan in Hertlein, 1934). 228 Tvoe Localitiesi Narlca lnsculota - Acapulco, Mexico (Carpenter, 1865), Eunatlclna helm! - Magdalena Bay, western Baja Call- fomtaj Pleistocene (Jordan In Hertleln, 1934). Tvoe Materlali Narlca lnsculota - Holotype, USNM 11841. Eunatlclna helnl - Holotype, CAS 5557. Nonenclatural Commentaryi 1 fortuitously found the holo type of Narlca lnsculota In the USNM general collection, where It had lain undetected since 1865. This specimen Is a young adult that Is 6.9 mm In height and 6.3 mm In diameter, and Is Identical In form with the type of £. DiscussIoni The radular dentition of £. lnsculota differs from that of every other eastern Pacific natlcld in having a multlcuspate rather than a tricuspate or monocuspate rachldian* In this respect, the dentition of £. lnsculota resembles that of the western Pacific species £• papilla Gmelln, 1791, the type species of Eunatlclna. The radular dentition of £. papilla (see Azuma, 1961, pi. 13, fig. 5t Oyama, 1969, text-flg. 7) has a prominent central cusp and downtumed basal processes of the rachldian to distinguish It from that of £. lnsculota. Although clearly different, the dentitions of these two species are 229 similar to each other and distinct from most other naticid dentitions in having multlcuspate rachldlans. The radular drawings by Azuma (1961) and Oyama (1969) of £. papilla differ in details of the lateral and marginal teeth, making these hard to compare with £* lnsculota■ However, both workers show the inner marginal teeth with a single apical termination, whereas the corresponding teeth of £. lnsculota have double terminations. The operculum differs from other calcareous naticid opercula known to me in being only partially calcified. I have seen only two opercula of £. lnsculota. but both lack calcification along the outer margin, with the thickest calcareous deposits along the inner margin and inner part of the final opercular whorl. The operculum of the geno type, £. papilla, is apparently entirely chltinous (Souverble & Montrouzier, 1874, pi. 7, fig. 8i Oyama, 1969, text-fig. 6), without the outer calcareous layer of £. lnsculota. It Is not possible to say whether the primitive opercular condition of Eunatlclna is chltinous or calcareous) no opercula are known from the Pleistocene specimens of £. Insculpta. Subfamily Natlcinae Gray, 1834 Nomen translatem W«nz, 1941, ex Naticldae Gray, 1834, * Verenaticinae Cossmarm, 1925. 230 Diagnosist Usually globose, mostly smooth) spire low to moderately elevatedi umbillcate, with funicle separated from parietal callus by a sulcust if imperforate, umbilical callus is a semicircular plugt external cal careous layer on operculum is usually thick and ornament ed. Discussioni This subfamily is characterized by its en tirely calcareous operculum and its funicle within the umbilicus of most species. It is principally a tropical group and the species are often brightly colored and patterned, and it is the subfamily most highly developed In opercular sculpture and umbilical morphology. Genus Natlea Scopoll, 1777 Natica Scopoli, 1777i392. Type-species» Nerlta vltellus Linnaeus, 1758, by sub sequent designation (Harris, 1897). Recent, Indo- Paciflc. Figured in Wenz (1941t1039, fig. 2974). Natica Lamarck, 1799i77. Tvoe-speciesi Nerlta vltellus Linnaeus, 1758, by sub sequent designation (Anton, 1839). Recent, Indo-Pacific. Diagnosisi Shells small to medium In size, globose, with low to moderately elevated spiresi outer surface often brightly colored and patterned, smooth or with incised 231 axial and/or spiral lines* Umbilicus usually narrowly to broadly open, sometimes closedi umbilical callus and funicle robust to weak, callus located Just below middle of inner lip. Parietal callus thin to moderately thick* Operculum calcareous, usually with one to many spiral ribs on outer face, sometimes smooth. Radula typically naticid, with a tricuspate rachldian, one multlcuspate lateral, one bicuspate inner marginal, and one mono- cuspate outer marginal tooth per half-row* Discussioni This genus is characterized by its cal careous operculum and, in most cases, its open umbilicus with distinct funicle. In his type designation, Scopoli (1777) refers to "Natica. Adanson," meaning Adanson (1757*172), a pre- Linnaean description of the genus* Anton (1839*31) ap parently designated Natica vltellus as type-species of Natlea Lamarck, 1799, not Scopoli* Anton (1839ivi) clear ly stated that specific names printed in capitals in his work were type-species of their genera. He listed 6 species (a through f) under Natica. and none of them is capitalized, although fflfnlIlfrift U-ated first and printed in ordinary letters and the others In Italics. However, under vltellus he listed several synonymous species, Including "VITELLUS Lam* ■ Nerlta vltellus L*," which presumably designates £• vltellus as type-species of 232 Natica Lamarck. This designation was first noted by Woodrlng (1957i84) and further brought to my attention by Cemohorsky (1971* personal communication). Because Anton's (1839) designation Is for Natica Lamarck* not Scopolii and Is therefore of doubtful validity* and be cause it is done In a somewhat ambiguous manner* It seems best to disregard It. Harris' (1897) designation of Nerlta vltellus as type-species of Natica Scopoli is con sidered the first valid designation. Fortunately* this is the same species used for Lamarck's taxa* and the type concept has remained stable. A key to the subgenera of Natica. s.l., used here follows I la. Shells smooth* lacking sculpture ..... 2 lb. Shells sculptured* usually with axial grooves 3 2a. Shells umbllicate, umbilical callus slender Natica. s.s 2b. Shells umbllicate* umbilical callus semicircular* not 2c. Shells Imperforate . . • 3a. Axial grooves extend from suture to base ....... slender IfiCtWMtUa Crragnat left Stlgmaulax 233 3b• Axial grooves occur only above periphery .................... 4 4a. Posterior margin of umbilical callus isolated as low, sharp riblet.............. GlvphepiEl^gifrft 4b. Umbilical callus slender, funicle reduced to narrow twisted vertical ridge along inner lipi operculum with one marginal riblet .............. Lunala 4c. Umbilical callus slender, with a posterior sulcusi funicle robust to lowi operculum usually with several spiral rlblets over outer portion ............. Naticarlus Subgenus Natica Scopoli, 1777, sensu stricto Diagnosisi Shells globose, smooth, lacking sculpture. Umbilicus relatively narrow, but not slit-likei umbilical callus slender, funicle low. Operculum with one too many spiral ribs on outer surface. Discussioni Natica. a *A* , is characterized by a smooth shell, open umbilicus, and slender umbilical callus. It is essentially a worldwide group in the tropics, with few species known In cooler climates. 234 Natica (Natica) claifel Etherlngton, 1931 Plate 2, figure 39 Natica (Natica) clarki Etherlngton, 1931*93, pi. 12, fig, 12. Nati^a^(Tectog^tl^a) c^rkl Etherlngton, Weaver, 1942* "Cryptonatlea oreRonensls (Conrad)-, Moore, 1963.pl. 2, fig. 2 (not £. oreRonensls (Conrad, 1857)J, Description. Size.--Baaed on 4 known specimens, size ranges from height 16.9 mm, diameter 15.8 mm (holotype) to height 9.6 mm (incomplete), diameter 9.2 mm (Incomplete). Shell Form.-"Shell globose, spire low to moderately elevated) body whorl not greatly inflated* whorls evenly rounded except for distinct flattening below suturet shell thin* whorls 5| suture strongly impressed. Shell smooth except for minute Incremental growth lines. Parietal callus thin, lightly filling posterior apertural anglei anterior lobe weak, projecting above umbilicus. Umbilicus narrowly openi sulcus distinct, with more abruptly excavated posterior margin and more gently slop ing anterior margini channel narrow, may be reduced to groove, tapering anteriorly. Umbilical callus narrow to somewhat swollen, with more abrupt posterior tapert funicle low, broad. Anterior Inner lip and basal lip thickened* 235 Operculum.--Unknown. Specimens Examined! 4. stratlaraohlc Occurrencei Ranges from Lower Miocene (?) to Middle Miocene. Lower Miocene (?)t Sooke Formation* Vancouver Island* British Columbia (Clark & Arnold* 1923). Middle Miocenei Astoria Formation* southwestern Washington (Etherlngton* 1931)i Montesano quadrangle (USGS)* and Grays Harbor County (CAS)* Washington* and Astoria* Oregon (Moore* 1963). Tvne Localityi UCB A-373* seam of fossils in bed of Rock Creek* about one-fourth mile downstream from the falls and Just below the old dam site* Section 7 T16N* R5W* Grays Harbor County* Washington (Etherlngton* 1931). Tvoe Materiali Holotype* UCB 31996, Discuss ion i Among the few known specimens of . (£.) clarki. the greatest morphologic variation Is in the form of the umbilical callus. The holotype has a slender callus that does not greatly restrict the umbilicus* but other specimens have a more swollen callus whose Increased width reduces the umbilical channel to a groove. This species is related to Natica (Tectomatlca) lanthosmm Deshayes* 1841, a Lower Pliocene to Recent species. Fossils of (J.) ianthostoma are much larger 236 than specimens of f i . (fl.) clarkl. however* and Recent specimens are larger still. The umbilical callus of J j . (J.) lanthostoma Is much broader and more semi-circular in outline than that of £ $ . (2 *) clarkl. and the umbilical sulcus is more narrowly excavated. The posterior apertural angle of (X.) ls al*° more thick ly filled than on £. (££.) clarkl and the shell wall is somewhat thicker. Their general similarity* however* suggests a possible ancestral relationship. Natlea (Natlea) clarkl was not found by Moore (1963) in her extensive report on the Astoria Formation in Oregon* so the species was apparently restricted to more northern cool-water habitats. Etherlngton (1931) com mented that £. CH.) clarkl "occurs abundantly at Kern River, California" and "at the type Astoria," but this species is unknown there. He was apparently referring to Polinices (Eusplra) vlctorlanua Clark A Arnold* 1923, an early to Middle Miocene species that occurs In the Astoria Formation and abundantly in the Kern River Miocene deposits. The figure in Weaver (1942) is mistakenly referred to as the holotype of Jg. (fg.) clarkl. but obviously differs In shell form and umbilical morphology. His figure may be of an atypical specimen of £. (£.) vlctorlanua. but is otherwise unknown to me. As explained more fully in the discussion of £• 237 gookansia Clark & Arnold) 1923, the holotype of that species has been lost* and the specimen in Its place in the Stanford University collection Is apparently an (£.) clarkl. This specimen tentatively establishes clarkl in the Lower Miocene, Natlea (Natlea) lnexpectans 01sson, 1971 Plate 2, figures 40, 41 Naticajjiexg^ tgyg 01ssan^l97l.69-7l, figs. 31a,b, 32i Description i Color.--Shell with axial stripes of white and dark brown extending from suture to base* sometimes with zig zag pattern on shoulderi base whitej callus whitej base white. Nuclear whorls white, translucent* Perio- stracum thin, pale yellowish brown. Size.--Averagei height 17.7 mm, diameter 18.8 mmi largest specimeni height 23.1 mm, diameter 22.6 mm (holo type). Shell Form.--Shell globose, spire lowj whorls evenly roundedt shell thlcki whorls 5, nuclear whorls not clearly set-offi suture slightly Impressed. Whorls smooth except for Incremental growth lines. Parietal callus thin, but thickens to moderately fill posterior apertural anglet anterior lobe distinct. Umbilicus open, rather narrowi 238 channel narrow, tapers anteriorly. Umbilical callus narrow, tapers gently anteriorly, expands posteriorly to meet parietal callusi funicle broad, low. Inner lip and basal lip slightly thickened. Operculum.--Calcareous. white, mostly smooth, with 4 smooth, flat-topped ribs along the outer margin and separated by narrow Interspaces! nucleus slightly swollen. Specimens Examinedi Holotype and 3 paratypes. Geographic Occurrence and Ecologyi Known only from the Gulf of Panama, northeast of Cabo Mala (about 7°39'N, 79°40*W), in 117 to 119 meters depth. Type Localityi Gulf of Panama, east northeast of Cabo Mala, 7°39.5*N, 79°40.7*W, 117 meters depth (Olsson, 1971). Type Materiali Holotype, USNM 701161. Discussioni The color pattern is unique among eastern Pacific species* Watle. (Natlca) .lalllata McLMn, 1970 Plate 3, figures 1, 2 11*0^ al&lllata McLean, 1970i313-314, pi. 46, figs. 11, Natlgg^CNatlea) stalllata McLean, Keen, 1971*475, fig. 239 SMStiUBtlflD* Color.--Shell chestnut-brown with closely spaced tri angular white markingst callus and umbilical area whitei Interior white with brown outer lip. Nuclear whorls yellowish brown. Perlostracum thin, yellowish white. Size.--Average» height 10 mm, diameter 10.5 mmi larg est specimen! height 19.5 ism, diameter 19.2 mm CSU 49428, Carmen Island, Gulf of California, Mexico!. Shell Form.--Shell globose, spire moderately elevated to lowi body whorl Inflatedi whorls evenly roundedi shell thickness averagei nuclear whorls 2, smoothi postnuclear whorls 3^i suture moderately Impressed. Spiral sculpture of minute, poorly developed, obscure costellaet axial sculpture of incremental growth lines that are best- developed near the suture. Parietal callus very thin, lightly fills posterior apertural anglet anterior lobe weak. Umbilicus open, narrowi sulcus very weakf channel tapers gently, not to a sharp point. Umbilical callus narrow, little more than thickened inner lip, then broadens more abruptly to bridge posterior end of umbili cus \ funlcle essentially lacking. Basal lip thickened. Operculum.--Calcareous. white, darker centrally from Incorporated sediment grains. Outer surface smooth ex cept for 4 flat-topped ribs near outer margin, the inner 2 broader than the others. 240 [ i Holotype and 3 paratypes. Geographic Occurrence and Ecologyi Near Carmen Island, Gulf of California, Mexico (26°N), and the Galapagos Is lands* Ecuador (0°N), unknown in intervening area. Found at 37 to 82 meters depth near Carmen Island and at 91 * meters depth in the Galapagos. Type Localityi Tagus Cove, Isabella Island* Galapagos Is lands, Ecuador, 0°16'5, 91°22’W, 91 meters (McLean* 1970). Tvoe Materiali Holotype, LACM 1284. Discussioni This is the only eastern Pacific species with triangular markings on its shell. Natica (Natlca) vokesi Addicott, 1966 Plate 3, figures 3* 4 Natica (Natica) vokesi Addicott, 1966ai638-639, pi. 77* figs. 2-5. PflggrtFUflBf Size.--Height 20.1 mm* diameter 19.4 mm (holotype). Shell Form.--Shell globose, spire moderately elevatedi body whorl moderately Inflated\ whorls distinctly flatten ed below suturet shell thlni suture moderately to strongly impressed. Shell smooth except for Incremental growth lines. Parietal callus thin* moderately filling posterior 241 apertural angle\ anterior lobe weak. Umbilicus open, sulcus indistinct. Umbilical callus slender* gently tapering anteriorly and posteriorlyi funicle low. poorly differentiated. Anterior inner lip thickened. Operculum.--Unknown. Specimens Examinedi Holotype. StratUraphlc Occurrence i Middle Miocene i Astoria Forma tion. Lincoln County. Oregon (Addicott. 1966a). Type Localityi USGS Cenozolc locality M1803. Intertidal zone at western edge of point due east of Gull Rock. 550 feet south. 4150 feet west of northeast corner section 32, T9S, R11W, Cape Foulweather quadrangle, Oregon. Tvoe Materiali Holotype, USNM 649130. Discussioni This species differs from Natica (Natlcarlus) ooauncula Hanna & Hertleln and (£.) t eft land 1 Hanna & Hertlein by having a much more slender umbilical callus, a more indistinct funicle, and a less distinct umbilical sulcus. It also lacks the axial wrinkles below the suture on Juveniles of the other two species that place them in the subgenus Natlcarlus. 242 Subgenus Natlcarlus IXunerll, 1806 Natlcarlus Dumeril, 1806i164. Type-speciesi Nerlta canrena Linnaeus* 1758, by monotypy. Recent, West Indies and southeastern U.S.A. Figured in Abbott (1954, pi. 5, fig. 1). Diagnosisi Shell snail to medium in size, globose to somewhat elongate, body whorl distinctly inflated. Axial sculpture of sharply incised grooves that extend from suture part way to periphery. Umbilicus usually narrow, rarely broad, with moderately to deeply excavated sulcus. Umbilical callus slenderi funicle ranges from robust to low and indistinct. Parietal callus thin to slightly thickened. Operculum with one or two ribs at outer margin or several ribs over outer portion. Discussion. This is a worldwide group, confined largely to the tropics, and is among the larger subgenera of naticids in number of species. Natlcarlus is character ized by its distinct radial grooves above the periphery, slender umbilical callus, and an operculum usually with several spiral ribs. Natica (Natlcarlus) chemnltzli Pfeiffer, 1840, nomen ln^uir«»r|fliim 243 Plate 3, figure 5 Natica chemnitz 11 Pfeiffer, 18401villi Steams, 18941195. not Natica chemnltzli "Recluz", Reeve, 1855(pi. 2, figs, sP.]. "Natica marochlensls Gmelln" [not J J . marochlensls Gmelln, 1791i Recent, Caribbean and Mediterranean^, Carpenter, I857t26l, 274t Sowerby, 1883*82, pi, 5, fig. 62, pi. 8, figs, 108, 109, pi. 9, fig. 151i M. Smith, 1944i 12, fig, 131. Natica naroohletfU var. chemnltzli Pfeiffer, Tryon, 1886i24, pi, 5, figs• 94-90. "Natica fyiryoranfl Chemnitz" C“ Nerlta MrorganaB Chemnitz, 1781, non-binomialKoch, lo44 »l52, 1551 Philippi, 1851(78-80, pi. 12, figs. 1-5 (plate 1850)i Carpenter, 1857(448-450) Carpenter, 1864(624) Steams, 1894(446) Menke, 1847i179 [as of Koch]i Menke, 1851(165 [as of Natica? Mroecana var. callfomlca Carpenter, 1857b(2271 Palmer, 1958(46. Natica nrltchardl Forbes, 1852(272, pi. 11, figs. 2a-ci Stearns, 1894(195• "Natica lurlda Philippi" [not N. Philippi, 1852, " " U* aaulterlana Recluz, 1844) Recent, Indo-Paciflcj) Carpenter, 1857(261. Natica undata Philippi, 1852(74, pi. 11, fig, 12 (plate 1850)7 Troschel, 1852*161) Dali, 1909(235. ? Natica undata Sassi, 1827(478 [not seen]). not Natica undata Philippi, 1844, vol. 2tl4l [■ fossil, Sicily], ? Natica undata Meusehen, 1767 [not seen}. Pfeiffer, Keen, 1958(320, fig 75, fig. 861. Koch]. 244 Description! Color.--Shell with axial zig-zag pattern of maroon brown and grayish yellow or grayish white, with bolder pattern on shoulderi pattern best-developed on Juveniles, often cloudy on adultsi adults often with whit1st blotches on shoulder that may coalesce into a solid bandi narrow whitish band is Just below suturei base and callus area white. Nuclear whorls dark maroon brown. Interior medium to dark brown, often axially streaked, with white band at shoulder and base. Perlostracum thin, light to medium yellowish brown. Size.--Averagei height 28 mm, diameter 25 mmj larg est specimen! height 39.1 mm, diameter 37.4 mm CiACM H-527, near San Felipe, Baja.California, Mexico^. Shell Form.--Shell elongate, spire elevatedi body whorl not greatly inflated! whorls evenly roundedi shell moderately thick, with solid appearancei nuclear whorls 2%, smooth! postnuclear whorls 3^i suture moderately to deeply impressed. Spiral sculpture of minute, poorly developed costellae! axial sculpture of closely spaced retractive grooves extending from suture half-way to periphery on early adult whorls, becoming indistinguish able from growth lines on final adult whorl. Parietal callus thin, moderately filling posterior apertural anglei anterior lobe very weak. Umbilicus open, narrow, sulcus well-developed, channel slightly tapered anteriorly. 245 Umbilical callus narrow, with evenly rounded margin raised into a ridge where it meets funlclei funlcle low broad. Basal lip thickened. Operculum.--Calcareous, white, darker on central area from incorporated sediment grainst thickest centrally, tapers toward outer marglm smooth except for one low, rounded rib at outer margin. Specimens Examined! 867. Geographic Occurrence and Ecologyi Bahia Magdalena, western Baja California, Mexico (23°30'N), throughout the Gulf of California, and south to Bahia de Sechura, Peru (5°40*S). Found mostly on sand and mud flats intertidal- ly, but known to 18 meters depth. * Stratl&raphlc Occurrencet Miocene-Pliocene ( ? ) i Bahia Magdalena, western Baja California, Mexico (Hanna, 1926). Pleistocene» Imperial County, California (Hanna, 1926). lyre U>caUtUg» Natica chemnltzli - Mexico (Pfeiffer, 1840). Natica? maroccana var. ggjUgffllSa - California (Carpenter, 1857). Natica orltchardi - Mazatlan, Mexico (Forbes, 1852). Natica undata - Panama (Philippi, 1852). 246 I m Natica chemnltzli - Unknown, either destroyed with Ffelffer collection in Stettin Museum during Second World War, or in Cuming collection in BM(NH) (Dance, 1966). Natica? naroccana var. callfomlca - Unknown, pre sumably in TJSNM, BM(NH), or Redpath Museum, McGill Univ., Montreal, Canada (Dance, 1966). Natica orltchardl - BM(NH) (Dance, 1966). Natica undata - Unknown, presumably In BM(NH) (Dance, 1966). Nomenclatural Commentaryi When Pfeiffer (1840) described £ j . ({£.) chemnltzli. he cited as examples figures 1905-6 of Chemnitz's (1781) six figures (1905-10) of the non-binomial Nerltae maroccanae. Chemnitz had recorded Neritae maroc- canae from Africa, especially Morocco, and the Antilles. Gmelln (l791) described Nerita marochlensls. basing his species on Chemnitz's six figures and locality data, and Dlllwyn (1817) renamed the species Nerita maroccana. cit ing all six of Chemnitz's figures and recording the species from the West Indies, Africa and the Bay of Naples. Quoy & Galmard (1833) were the first to report f l . marochlensls from the central Pacific, and were followed by many later workers. In a review of the taxonomic history of f i . mrnffhUniia. Cemohorsky (1971) concluded that the name 247 may only be applied correctly to Mediterranean and Carib bean specimens. The status of J g . (£.) chemnltzli Is uncertain. If the name N. (fg.) marochlensls applies only to Mediterranean and Caribbean specimens, as Is Indicated In the original description, then £. (Jg.) chemnltzli Is a Junior synonym of £. (££.) marochlensls. because g. (£.) chemnltzli is based upon two of the six original figures of £. (£j.) naroohUmta. If examination of the types of f g . (£g.) Mroohlens1s (whose location is unknown) shows the speci mens given as Chemnitz's figures 1905-6 to be from the eastern Pacific, then Pfeiffer's £ j . (Jg.) chemnltzli Is availablei this seems unlikely. Deciding on the avail ability of J g . Qg.) chemnltzli would still leave the question of whether eastern Pacific specimens are separ able from those of the Caribbean and Mediterranean. Specimens from the two regions are not clearly different, but 1 have not seen enough of the Caribbean and Mediter ranean examples to be sure they show the same range of variation in color and shell form as the Pacific specimens. Because it is possible that J g . (Jg.) is a Junior Mrnchlanili. and that specimens from the Pacific and At lantic represent one species, J g . Qg.) tzll is cited here as a nomen Inquirendum (name under inquiry). Discussioni Among eastern Pacific species, £* (Jg.) 248 Chemnitzil is nost similar to Q$.) unifasciata Lamarck. The distinctions between these species are given in the discussion of (£•) unifasciata. Natica (Natlcarlus) unifasciata Lamarck, 1822 Plate 3, figure 6 Natica unifasciata Lamarck, 1822, vol. 6, part 2t201i Troschel, 18*2i160i Reeve, 1855ipi. 12, fig. 49bt Dali, 1909i235i Jordan, 1924•156j Hanna, 1926*451i M. Smith, 1944112. Natica marochlensls var. unifasciata Lamarck, Tryon, 1886t 23-24, pi. 5, fig. 93. Itelica (Mllss) rnUMSlfttS Lmtck. (Srmt & Gale, 19311 *97, text-flg7 10i Keen, 1971*475, 477, fig. 868. Natic^(Na^ ca) chemnltzli Pfeiffer £in part'] , Keen, Descriptioni Color.--Shell light grayish brown to dark purple and nearly black, with a narrow band of dull yellow on the shoulderi colors often axially streaked, may become abruptly lighter below periphery! base white\ nuclear whorls dark purplish brown. Interior streaked with gray and brown. Perlostracum thin, light to medium yellowish brown. Size.--Averaaei height 35 mm, diameter 31 mmi larg est specimen* height 44.0 mm, diameter 37.8 mm. Shell Form.--Shell elongate, spire elevated* body whorl inflated! whorls evenly roundedi shell moderately 249 thick, with solid appearance* nuclear whorls 2%, snoothr postnuclear whorls 3%* suture moderately to deeply Im pressed. Spiral sculpture of minute, poorly developed, obscure costellaej axial sculpture of closely spaced re tractive grooves extending from suture half-way to peri phery on early adult whorls, becoming Indistinguishable from growth lines on final adult whorl. Parietal callus thin, moderately filling posterior apertural angle* anterior lobe very weak. Umbilicus open, narrow, sulcus well-developed, channel only slightly tapered anteriorly. Umbilical callus relatively narrow, with evenly rounded margini funicle low, broad. Basal lip thickened. £o&££Ulmi>--Calcareous, white, darker on central area from Incorporated sediment grains* thickest centrally, tapers toward outer margin* smooth except for one low, rounded rib at outer margin. S p e c i m e n s E x a m i n e d i 3 8 6 . Geographic Occurrence and Ecology* La Union, El Salvador (13°20'N), south to Golfo de Guayaquil, Ecuador (3°S). Specimens with questionable locality data are from the Golfo de Tehuantepec, Mexico <16°N), "Peru," and the G4lapagos Islands. Collected alive lntertldally on mud flats. 250 Stratlgraphlc Occurrencei Hanna (1926) reported this apeclea from the Pliocene? of Coyote Mountain, Imperial County, California, but CAS and UCB specimens from that area, labeled as unifasciata. are too poorly preserved to identify. Type Localityi Unknown (Lamarck, 1822). Type Materiali Unknown, presumably in Museum d'Histoire Naturalle, Geneva, Switzerland, or Museum d'Histoire Naturelle, Laboratoire de Malacologie, Paris, France (Dance, 1966). Nomenclatural Commentaryi As noted by Keen (1971), unifasciata is not recognizable from Lamarck's original description, but a figure by Delessert (1841) of a speci men from Lamarck's collection shows this to be an eastern Pacific species. Discuss ion i NfttU* (Natlcarlus) unifasciata is most similar to £. (£).) chemnltzli Pfeiffer, 1840, which ranges from Baja California, Mexico, to Peru. The most obvious difference between the species is in color patterns, (hO chemnltzli having a series of yellowish blotches on its shoulder instead of a solid yellowish band, and a zig-zag pattern of yellow and purple or brown over the rest of the shell, instead of a solid dark shell as in { j . (£J.) uni fasciata. The zig-zag pattern of £. (£).) chemnltzli is 251 best developed on Juveniles, whereas older specimens may have a mottled dark brown or purple shell similar to paler specimens of { j . (£.) unifasciata. Older Individuals of {J. ( £ J . ) chemnltzli may also have their yellowish shoulder mottlings coalesced Into a clouded band that is wider than the similar band in £. (JJ.) unifasciata. How ever, the early whorls of f i . (A*) chemnltzli always show their characteristic zig-zag pattern, whereas juveniles of (£J.) unifasciata never do. Development of the umbilical callus is also different between the two species. The callus of (12.) unifasciata Is wider and more swol len, and fills In the umbilicus more completely. As a result, the umbilical channel of (£).) chemnltzli Is broadly open to its termination at the Inner lip, whereas the channel of £. (f£.) unifasciata Is reduced to a narrow groove for most of Its length. The Junction of the callus surface and funic le surface In J J . (£4.) chemnltzli forms a sharp ridge along the free margin of the callus, unlike the broadly rounded callus surface of (£.) unifasciata that conceals most of the funlcle. Although these dif ferences are remarkably consistent, occasional specimens seem to have characteristics of both species. These usual ly have the solid color pattern of £. (£,) unifasciata and the more open umbilicus of g. (£.) chemnltzli. Such speci mens are either individuals of (£.) chemnltzli that duplicate the color of (£.) unifasciata as a gercntlc 252 feature, or are unusually large specimens of J g . (ft.) uni fasciata. which develop a more open umbilicus as they attain great size. The geographic range difference between (£.) uni- fas data and (£.) chemnltzli reinforces the distinction made on the basis of umbilical morphology and color pat tern. £ $ . (£.) unifasciata occurs only In the inner tropics, from El Salvador to Ecuador, whereas £. (£.) chemnltzli ranges from southwestern Baja California, Mexico, to Bahia de Sechura, Peru, at the northern and southern limits of the eastern Pacific tropics. Natlc^(Ng^ca) othg^o Dali, Keen, 1958, 3211 Keen, Description* Color.--Shell medium to dark brown, often grayish, with a narrow white band at the suture and a white baset shoulder with a narrow band of alternating white and dark brown spotsi a narrow white band is just below periphery, with a row of dark brown spots along the upper margin of the band, sometimes with an additional row of smaller spots along the lower margin. Nuclear whorls orange- Natica (flatlcarWg) othello Dali, 1908 Plate 3, figures 7, 8 ) othello Dali, 1908*332-333| M. Smith, 253 brown* becoming whitish at shoulder. Periostracum thin* pale yellowish white. Size.--Averagei height 20 mm* diameter 18 mmi larg est specimeni height 24.8 mm* diameter 22.0 mm QJSNM 96992* Panama Bay, Panama3. Shell Form.--Shell elongate* spire elevated* body whorl inflated} whorls evenly roundedi shell thlni nuclear whorls 3* with spiral sculpture of microscopic* closely spaced costellae on early whorls* and of re tractive axial grooves below the suture on third whorl, the two types overlapping) postnuclear whorls 3t suture moderately impressed. Axial sculpture of retractive grooves that extend from suture one-third of the way to periphery, better developed on early adult whorlst spiral sculpture of minute, poorly developed, obscure costellae. Parietal callus thin, transparent* lightly to moderately filling posterior apertural angle, with weak to moderate anterior lobe overhanging umbilicus. Umbilicus open, narrow, typically with deep sulcus that indents posterior margin of umbilical callust channel narrow, tapering anteriorly, but not to a point. Umbilical callus rela tively narrow, gently tapering anteriorly, but abruptly truncated and notched posteriorly by umbilical sulcusi funicle robust, filling anterior half of umbilicus* Anterior inner lip thickened* basal lip thin* Operculum.--Calcareous, white* thickened centrally and 254 thin along outer margini sculptured with two outer- marginal ribs separated by a narrow groove* the outer rib rounded and minutely pustulate* the inner rib sharply raised and more weakly pustulate. Specimens Examinedi 467. GeoRraohlc Occurrence and Ecologyi Isla San Benito, western Baja California* Mexico (28°17'N), throughout the Gulf of California, and south to Bahia Santa Elena (2°10*S) and the Galapagos Islands* Ecuadort rare north of the Gulf of California. Collected alive in depths of 4 to 265 meters* most common in 18 to 45 meters depth on a variety of soft substrates. Type Localityi Bay of Panama* Panama (Dali* 1908). Type Materiali Holotype* USNM 46446, DiscussIoni The deep sulcal notch at the posterior margin of the umbilical callus is the easiest means of identify ing this species* but It is not always well developed. Some individuals entirely lack this notch and have a relatively gentle posterior taper to the callus. Two specimens among those examined differ from typical X J . (E.) othello specimens in color and form, and may be considered a new subspecies or species when more specimens are available for study. These specimens fAHF 1703-49* 255 Cedros Island, Baja California, Mexico, and USNM 210762, Gulf of Califomlaj, differ from ordinary £. (g.) othello by having a lower spire and more globose whorls. In ad dition, they have 4 distinct spiral bands of alternating white and brown, instead of the 3 in J j , (£.) othelloi prominent ones on the shoulder and base, and two much narrower ones at the periphery. Also, the two ribs along the outer margin of the operculum are more widely spaced, and the inner rib not as sharply crested, as in typical (£J.) othello. The color pattern is nearly identical to that of ( £ J . ) gravL Philippi, although that species has a fifth brown and white band near its suture. More speci mens than the present two would be needed to erect a sub species of £. (£.) othello or a new species for this variant. Natica (Natlcarlus) caneloensls Hertleln & Strong, 1955 Plate 3, figures 9, 10 Natica caneloensls Hertleln A Strong, 1955*287, pi. 2, figs. l3, 18. Keen, 1 r l«473, fig. W&fliSBWBaSS ££r3):- K" ?n’ 195 8 ‘ 860. Descriptioni Color.--Shell pale greyish brown, with three rows of 256 quadrate chestnut brown spotsi rows of large spots on the shoulder and periphery, and a row of smaller spots (which may be lacking) on the basei narrow band of tan or dull yellow occurs Just below suture. Nuclear whorls medium orange-brown, becoming white* Interior pale violet. Perlostracum thin, pale to medium yellowish brown. Size.--Averagei height 28 mm, diameter 26 mmj larg est specimeni height 29.8 mm, diameter 28.0 mm QLACM, no number, off northern Peru]. Shell Form. — Shell globose, spire moderately elevated, body whorl inflatedi whorls evenly roundedi shell thim nuclear whorls 3, smootht postnuclear whorls 3%) suture moderately Impressed. Axial sculpture of fine, retractive grooves that extend from the suture less than half-way to the periphery, and become indistinct on the final adult whorlj spiral sculpture of minute, poorly developed, ob scure costellae. Parietal callus extremely thin, trans parent, lightly filling posterior apertural anglei anterior lobe well-developed. Umbilicus open, relatively narrow) sulcus narrow and shallow) channel tapers to a point anteriorly. Umbilical callus narrow, gently taper ed anteriorly and posteriorly. Basal lip not thickened. Qoerculum.--Calcareous. white, central area dark from incorporated sediment grains) outer margin with two raised ribs undercut by adjacent deep grooves) inward from marginal ribs are 4 to 7 lower and flatter ribs separated 257 by very narrow and shallow groovest outermost margin with one minute* weakly pustulate costella. Specimens Examinedi 150. Geographic Occurrence and Ecologyi Cabo San Lucas* southernmost Baja California (22°45,N), and Mazatlau (23°14*Ji), Mexico* south to off Paita* Peru (5°S). Found alive in depths of 118 to 133 meters* with empty shells known in as shallow as 24 meters. Type Localityi Port Parker* Costa Rica (Hertleln & Strong* 1955). Type Materiali Holotype* CAS 9891, Discussion» Hertlein & Strong (1955) state that (£ [. ) caneloensls may have four distinct spiral bands of brown spots per whorl* although their holotype shows only two such bands. Most specimens of J J . (£J.) caneloensls have two such bands* at the shoulder and periphery* and some individuals have a third band of smaller spots on the base* but 1 have never seen more than three bands per whorl. Hertlein A Strong (1955) described 4 ribs and 5 grooves on the operculum of their holotype* without not ing variations among other specimens. In a larger sample* the number of ribs Is seen to vary as described above* 258 with a total of 4 to 7 flattened Inner ribs and 2 stronger outer ribs. Natica (Natlcarlus) caneloensla differs clearly from £J. (£}.) brunneolinea McLean and £J. (g.) scethra Dali in shell color and opercular sculpture, but is very close to these species in shell form. Natica (Natlcarlus)caneloen sls differs from £ * . (£.) brunneolinea by lacking a low ridge or angulation along the anterior margin of its umbilical channel, by having a more sharply attenuated umbilical channel, and by having a thinner parietal callus without a distinct growing margin. Natica (Natlcarlus) caneloensls differs from £ 1 . (£•) scethra by having its umbilical callus more attenuated posteriorly, having a deeper and narrower umbilical sulcus, having more in flated whorls, and by lacking a low ridge or angulation along the anterior margin of its umbilical channel. Natica (Natlcarlus) collma Strong & Hertlein, 1937 Plate 3, figure 11 ca nn]l«s Strong & Hertlein, 1937i174, pi. 35, figs. 12, 15716. 259 D e s c r i p t i o n i Color.--Shell with irregular axial stripes of altemat ing medium brown and white, interrupted below suture and at periphery by narrow spiral bands of whitei wider pale brown bands usually occur on shoulder and between base and periphery! base whitet nuclear whorls medium orange-browni early adult whorls often graylshi callus whitei interior streaked with violet and white, with white band at suture and base. Periostracum thin, pale yellowish brown. Size.--Averagei height 18 mm, diameter 15 rami larg est specimen! height 22.5 mm, diametert 19.9 nun CLACM 67-20, Mazatlah, Mexico!. Shell Form.--Shell somewhat elongate, spire moderately elevated, body whorl Inflatedi whorls evenly rounded, shoulder slightly flattened! shell thlm nuclear whorls 3, with weak, closely spaced, microscopic spiral costellaei postnuclear whorls 3i suture moderately impressed. Adult whorls smooth except for closely spaced retractive axial grooves extending from suture half-way to periphery on first two adult whorls, becoming indistinguishable from growth lines on final adult whorl. Parietal callus thin, transparent at its center, lightly filling posterior apertural anglei anterior lobe weak, extending over umbilicus. Umbilicus openi sulcus broad, shallow) channel narrows anteriorly nearly to a groove. Umbilical callus narrow, with gentle anterior taper, more abrupt posterior 260 taperi slightly concave anterlorlyt funicle low, broad. Inner lip thickened, basal lip thin. Operculum.--Calcareous, white, central area often brown from Incorporated sediment grainsi central area shallowly concave. Outer surface typically sculptured with 8 sharply incised spiral grooves (may have 7 to 10), separated by 7 flat-topped to slightly concave ribs, with a rounded outermost rib. Inner margin with fine irregular wrinkles along its posterior half, and microscopic longitudinal wrinkles along its anterior half. Outer margin with 1 to 3 (usually 2) microscopic rlblets, in distinctly beaded. Geographic Occurrence and Ecologyi Cedros Island, western Baja California, Mexico (28°N), throughout the Gulf of California, and south to Golfo de Nlcoya, Costa Rica (10°09*N). One specimen CCAS 17991-94] is from "Bahia Honda, Panama," which 1 cannot locate* Found alive in depths of 32 to 130 meters, most commonly in 45 to 65 meters. Type Localityi CAS locality 27574, lat. 18°33'N, 103°45*W, Punta San Juan de Lima, about 47 miles southwest of Man zanillo, Colima, Mexlcoi 52 fathoms (Strong and Hertlein, 1937). Type Materiali Holotype, CAS 6996. 261 Discussioni The color pattern Is often poorly developed* so that specimens may mostly be pale brown. The final 1/3 whorl of a large adult is usually axially streaked with light and medium brown* lacking spiral color pat terns entirely* no matter how strong the spiral pattern may have been on earlier whorls. Variation in the number of spiral ribs on the operculum, from 7 to 10* is seen in a single population and has no obvious environmental sig nificance. Natlea (Natlcarlua) scethra Dali, 1908 Plate 3, figure 12 Natica (Cochlls) scethra Dali* I908i333* pi. 11* fig. 5| M. Smith* 1944i12. fig. 132. Natica (Natica) scethra Dali* Keen* 197li475* fig. 866. Descriptioni Color.--Shell light to medium yellowish brown* usual ly axially streaked, commonly but not predominantly with narrow pale bands at the suture, on the shoulder, and below the peripheryt base and callus white. Nuclear whorls light to medium brown, becoming white on last half- whorl. Ferlostracum thin, pale yellowish brown. Size. - - Average i height 24 mm* diameter 21 mmi larg est speciment height 29.3 mm, diameter 26.1 mm fLACM* un- 262 cataloged, off northern Peru In 160 meters!. Shell Form.--Shell somewhat elongate, spire moderately elevated» body whorl Inflated* whorls evenly roundedi shell thinj nuclear whorls 3. smoothi postnuclear whorls 3i suture moderately Impressed. Spiral sculpture of minute, poorly developed, obscure costellaei axial sculpture of fine, retractive grooves that extend from suture halfway to periphery, better developed on early adult whorls. Parietal callus very thin, transparent, lightly filling posterior apertural angle, with weak to moderate anterior lobe overhanging umbilicus. Umbilicus open, narrow, with a shallow sulcus and a channel that tapers anteriorly but not to a pointi a low cord or angu lation bounds the anterior side of the channel near the inner lip. Umbilical callus narrow, very gently tapered anteriorly, more abruptly tapered posteriorly, slightly concave anteriorly. Basal lip not thickened. --Calcareous» white, central area often dark from incorporated sediment grainsi outer surface smooth except for two deep, sharply Incised grooves near outer margin that forms two undercut ribs, one between the grooves and one along the outer opercular margin. Outer most margin with one minute, weakly pustulate costella. Specimens Examined* 292. 263 Geographic Occurrence and Ecology» off La Faz, eastern Baja California, Mexico (24°15'N), south to near Islas Lobos de Tlerra, northern Peru (6°23*S, 80°56*W), Rare throughout nearly its whole range, and not known in the Gulf of California north of La Faz, All but about 40 of the specimens examined by me came from two dredge hauls in the Gulf of Tehuantepec, southernmost Mexico (16°N), and one off of northern Peru. Found alive in depths of 37 to 281 meters, most abundantly in depth of 160 meters off northern Peru, in muddy substrates. Type Localityi Gulf of Panama, in 153 fathomsi U.S.S. "Albatross" station 3391 (Dali, 1908), Type Material» Holotype, USNM 123048. Discussioni flgttfiP (Naticarius) scethra differs from (£.) othello Dali by having a lower, more globose form, an umbilicus of more uniform width, and a smaller umbilical sulcus. Natica (Naticarius) scethra has two grooves near the opercular outer margin, whereas £. (£.) othello has two ribs. (Naticarius) collma differs from g. (£.) scethra by its color pattern, prominent umbilical sulcus, and more abruptly tapered umbilical channel, plus having a multi-grooved operculum. The principal differences between f i . (£.) scethra and 264 £ j . (H.) caneloensla Hertleln & Strong are ahell color and opercular sculpture. Natica (Naticarius) caneloensla has three spiral rows of brown spots on each whorl, and the operculum has 6 to 9 ribs. However, these two species are essentially Indistinguishable In shell form. Natica (Naticarius) caneloensla has a more globose shell, with a lower spire, and an umbilical callus that Is slightly wider and has a slightly more abrupt posterior taper than seen In (£•) scethra. These species probably could not be distinguished If they were preserved as fossils lacking color and opercula. Possible remnants of microscopic spiral sculpture are present on some nuclear whorls of specimens at hand, but none Is preserved well enough to show the sculpture clear ly, If It exists. Natica (Naticarius) frrvmwUnga McLean, 1970 Plate 3, figures 13, 14 Natica brunneollnea McLean, 1970*313, pi. 46, figs. 9, 10. Nat1^^(Natica) brunneollnea McLean, Keen, 197li473, fig. Descriptioni Color.--Shell marked with narrow, alternating axial stripes of medium brown and yellowish white, from middle of shoulder to point Just below peripheryt color becomes 265 paler and fades out toward suture, with narrow band of orange or yellowish white at suturei base and umbilical area whitei Interior whitish, showing exterior color pat tern. Nuclear whorls dark brown, becoming whitish. Periostracum thin, pale yellowish white. Size.--Averagei height 22 mm, diameter 21 mm* larg est speclmeni height 45.8 mm, diameter 42.3 CLACM type no* 1283, Isla Santa Cruz, Galapagos Islands]. Shell Form.--Shell globose, spire moderately elevated, body whorl inflated* whorls evenly rounded, very slightly concave below suture* shell thlni nuclear whorls 3, slightly tabulate, with minute retractive axial grooves extending a short way from the suture* postnuclear whorls 3i suture slightly impressed. Axial sculpture of retrac tive grooves that extend from suture less than half-way to periphery, more sharply incised on early whorls* spiral sculpture of minute, poorly developed, obscure costellae. Parietal callus thin, transparent, lightly filling post erior apertural angle* anterior lobe well-developed. Umbilicus open, relatively narrow* sulcus distinct but broad and shallow, rounded* channel tapers anteriorly, but remains broadly open to inner lip* a low cord or angulation bounds the anterior side of channel near the inner lip. Umbilical callus narrow, very gently tapered anteriorly, more abruptly tapered posteriorly. Basal lip not thicken ed. 266 Operculum.--Calcareous. white, central area often dark from Incorporated sediment grainsi outer surface smooth except for two deep, sharply Incised grooves near outer margin that form two undercut ribs, one between the grooves and one along the outer opercular margin. Outer most margin with one minute, weakly pustulate costella. Specimens Examined» 14. Geographic Occurrence and Ecologyi The Galapagos Islands, Ecuador (0°10*S, 87°45'W), dredged in 37 to 292 meters depth. ✓ Type Localityi Academy Bay, Isla Santa Cruz, Galapagos Islands, Ecuador, 0°45'S, 90°20'W, 50-100 fathoms (McLean, 1970). Type Materiali Holotype, LACM 1282. Discussioni This species is most similar to f t , (ft.) scethra Dali, and the opercula of the two species are in distinguishable. The most obvious difference is in the color patterns, the strong axial banding of f t . (IS*) brunneollnea contrasting to the more evenly brown color of f t . (ft.) scethra. In shell form, f t . (ft.) brunneollnea differs from the other species by its lower shell and more inflated body whorl, with more weakly impressed sutures, slightly broader umbilical callus, and more 267 arcuate umbilical channel. Although the two species are not known to occur together* they are closely related and probably derived from a common ancestor. Natica CNaticarius) gray! Philippi, 1851 Plate 3, figures 15, 16 Natica depressa Gray, I839il36, pi. 36, fig. 2. not Natica depressa S o w e r b y , 1812, vol. Ii21, pi. 5, lower figs. Cfossll, EnglandJ. not Natica alaucinoldea var. £• depressa Grateloupe, 1847i Atlas, Natica. pi. 5, figs. llT l2 Lfossll, France!. Natica arayl Philippi, 1851*74-75, pi. 11, fig. 13 (plate 1850; Creplacement name for J J . depressa Gray, 1839, not Sowerby, 1812J| Hertleln & Strong, 1955t286-287, pi. 2, fig. 14. Natica (Natica) aravl Philippi, Keen, 1958»320, fig. 260j Parker7 1^64i1537 Keen, 1971i475, fig. 863. Natica catenate Philippi, 1853 (1849-53)*130, pi. 18, fig. 11 (plate 1851)i Philippi, 1853i233* Reeve, 1855ipi. 21, fig. 92a, not 92b,ct Sowerby, 1883*82, pi. 8, fig, 106, not 107i Tryon, 1886i22, pi. 4, figs. 71- 731 Steams, 189411951 Pilsbry A Lowe, 19321126 Css "N. catenatus* * 1 i Palmer & Hertleln, 1936178-79, pi. 19, figs. 2, 111 M. Smith, 1944i12, not. fig. 124 C*fi. chemnltz11 Pfeiffer, 1840D. Descriptioni Color.--Shell light to medium brown, commonly grayish, with 5 narrow spiral white bands marked with dark brown spotsi uppermost band near suture is most distinct on early whorls, another band is on shoulder, one at peri phery, and two on base* lowermost band is along border of 268 white beset there is narrow white band between suture and uppermost brown-spotted bandi callus whitet interior white* sometimes showing pale violet* showing exterior color pattern. Nuclear whorls medium brown. Periostracum very thin* pale yellowish white. Size.--Averagei height 14 mm* diameter 14 mmj larg est speciment height 20.4 mm* diameter 20.3 mm QLACM HH-112* Venado Island* Panama Bay* Panama}• Shell Form.--Shell somewhat elongate* spire moderately elevated* body whorl Inflatedi shoulder slightly flattened* with a very weak tabulation near the suturef shell thini nuclear whorls 3, with microscopic retractive grooves on shoulderi postnuclear whorls 2j suture slightly to moderately impressed. Spiral sculpture of inconspicuous* irregularly-spaced lineations. Axial sculpture of closely spaced and sharply Incised retractive grooves that extend from suture half-way to the periphery) additional axial sculpture of incremental growth lines only. Varietal callus thin* transparent* lightly filling posterior apertural angle) anterior lobe weak* extends over umbili cus. Umbilicus open, sulcus broad and deept channel narrows anteriorly* but not to a point or groove. Umbilical callus concave* strongly depressed anteriorlyi anterior end gently tapered* posterior end abruptly terminated, greatest width at posterior margin) funicle robust. Basal lip slightly thickened. 269 Operculum.--Calcareous. whitef with depressed central area and slightly thickened nucleusi smooth except for two sharp. closely set ribs along outer margini outer side of outermost rib is wrinkled) looks almost beaded. Specimens Examinedi 480. Geographic Occurrence and EcoIqrvi Cabo San Lucas, southernmost Baja California. Mexico (22°45*N). the southern end of the Gulf of California, and south to Paita. Peru (5°S>. and the Galapagos Islands. Hertleln & Strong (1955) indicate a northern range limit on the outer coast of Baja California at Magdalena Bay (24°30'N). The Galapagos Island occurrence is based on a single dead specimen. Live specimens are found in 2 to 58 meters depths* most often in 5 to 15 meters, and empty shells have been found Intertidally. Specimens have been found on substrates of mud, sand and shell debris. Stratlaraphic Occurrencei Pleistocene - Oaxaca, Mexico (Palmer & Hertleln, 1936). Type Localities* Natica depressa - Unknown. Natica catenata - Unknown. 270 Type Material! Natica depressa - BM(NH) (Dance, 1966), Natica catenate - Unknown* presumably In BM(NH) (Dance* 1966), Nomenclature! Cnrai>»nfarv. Ihis species* as £ * . catenate. was reported from Sicily by Reeve (1855) and Sowerby (1883) who* according to Tryon (1886), confused it with 1 3 * marochlensls Gmelin* 1791. Figures 92b and 92c of Reeve (1855) and figure 107 of Sowerby (1883) are clearly not of £ 4 . (£J.) aravl. whereas the other figures they give are of this species. The figure given for £ 4 . aravl by Smith (1944) is of £ 4 . (fi.) chemnltz11 Pfeiffer, 1840, which is similar in form and color to £ 4 . marochlensls. Discussioni Five spiral bands of white and dark brown are invariably present, although the uppermost band is sometimes obvious only on early adult whorls. Some authors beginning with Hertleln & Strong (1955) reported only four such bands on the shells. Natica (Naticarius) oosuncula Hanna & Hertleln, 1938 Plate 3, figure 17 Natica poauncula Hanna & Hertleln* 1938il07-108* pi. 21, F i g . oi Keen, 1943i37* Keen & Bentson, 1944i1/7) Stewart * 19461table 2. Natic^(Ng^lg|rlua7)^poymculg Hanna & Hertleln* Addlcott, 271 Descriptioni Size.--Averagei height 21 mm, diameter 19 mmi larg est specimeni height 28.1 mm, diameter (Incomplete) 24.6 mm EUCB 2713, Temblor Formation, Callente quadrangle, Kern River area, California, middle Miocene]* Shell Form.--Shell somewhat elongate to globosei spire moderately elevatedi body whorl greatly Inflated, slightly concave near suturei shell thlni whorls about 5) suture slightly impressed. Shell smooth except for minute Incremental growth lines and low, Irregular, closely spaced axial wrinkles that extend a short way from suture of some specimens, especially on early adult whorls. Parietal callus thin to average, lightly to moderately filling posterior apertural anglei anterior lobe very weak to Inconspicuous, slightly overhangs umbilical sulcus. Umbilicus openi sulcus broad but shallow) channel broad posteriorly, narrowing nearly to a groove anteriorly* Umbilical callus relatively small, with evenly rounded and slightly elevated margin) funicle strong, relatively narrow. Anterior inner lip thickened) basal lip thickened and raised into a low forward projection. Or>erculum.--Unknown. Specimens Examinedi 174. Stratiaraphlc Occurrence i Known only from Middle Miocene strata. Middle Miocenei Temblor Formation, lower member 272 (Vertloecten zone) at Reef Ridge, CoalInga area (Stewart, 1946), Caliente quadrangle (UCB), Callfomlai upper Olcese Sand and lower Round Mountain Silt, Kern River area, Cali fornia (Addlcott, 1970i USGSi UCB)t Saltos Shale Member of Monterey Shale of Hill, Carlson and Dlbblee (1958), Caliente Range, California (Vedder In Addlcott, 1970). Type Localityi CAS locality 65, hills Just north of Kern River and northeast of Barker's ranch house, Kern County, California. Temblor Formation, Miocene (Hanna & Hertleln, 1938). Type Materiali Holotype, CAS 7084. Momenclatural This species Is assigned to the subgenus Naticarius because some well preserved specimens have axial wrinkles that extend from the suture part way to the periphery, even though most specimens are too worn to show this. Addlcott (1970) noted one specimen with faint spiral rows of light brown spots, similar to the color pattern of £. (ft.) canrena Linnaeus, 1758, the type- specles of Naticarius. DiscussIoni Natica (Naticarius) poauncula is most similar to £. (Jg.) tea land 1 Hanna & Hertleln, but differs by having a broader umbilical callus with a more strongly curved margin, and a much more robust funlcle. The umbilicus of (£.) poauncula Is thus filled by the callus 273 and funlcle to a greater extent than in (£J.) tea land 1. and the umbilical channel is correspondingly more narrow. Natica (Naticarius) tealandl Hanna & Hertleln, 1938 Plate 3, figure 18 Natiga (Natica) dal11 Tegland, 1933i138, pi. 14, figs. not Tectonatica dalli Cosstaann, 19251121. Natica tealand1 Hanna & Hertleln, 1938i108 £new name for 1 2 . dal 11 T08land, 1933]. Natica (Tectonatica) teglandae Hanna & Hertleln, Weaver, 1*5421333, pi. 68, figs. 5-7 [emendation of namej • Natica (Natica) teglandae Hanna & Hertleln, Durham, 1944i 139i Addlcott,1970i65-66, pi. 5, figs. 1, 2, 20. Descriptioni Size.--Average (based on few specimens)! height 20 mm, diameter 18 mmj largest specimen! height 26.0 mm, diameter (incomplete) 25.3 mm [tJCB 32215, type Blakeley Formation, northwestern Washington, Oligocenei holotype}, Shell Form.--Shell somewhat elongate, spire moderately elevatedi body whorl moderately inflated, flattened to slightly concave near suturei shell thini whorls about 5, protoconch sculptured with minute, closely spaced spiral costellaei suture moderately impressed. Shell smooth ex cept for incremental growth lines and weak, irregularly spaced axial wrinkles on some specimens. Parietal callus thin to average, lightly to moderately filling posterior 274 apertural anglej anterior lobe very weak to inconspicuous, slightly overhanging umbilical sulcus. Umbilicus openi sulcus broad but shallowj channel broad throughout its length, not narrowing much anteriorly. Umbilical callus small, with evenly rounded and slightly elevated marginj funicle strong, relatively narrow. Anterior inner lip thickenedt basal lip thickened and raised into a low for ward projection. Operculum.--Unknown. Specimens Examinedi 15. Strati&raphlc Occurrencei Ranges from Upper Oligocene to Middle Miocene. Upper Ollaocenei Blakeley Formation of Weaver (1912), Puget Sound area, Washington (Tegland, 1933i Weaver, 1942i Durham, 1944). Lower Miocenei Echlno- ohorla apta zone, northwestern Washington (Durham, 1944). Middle Miocenei upper Olcese Sand and lower Round Mountain Silt, Kern River area, California (Addlcott, 1970i USGS). Tvne Localityi UCB locality 681, Restoration Point, Kit sap County, Washington, Blakeley Formation of Weaver (1912) (Addlcott, 1970). Type Materiali Holotype, UCB 32215. Nomenclatural Co— ~nrmry, The specific name of £. (£.) tealandl was proposed by Hanna & Hertleln (1938) to re 275 place that of Watlea (Natica) dalll Tegland, 1933, which is preoccupied by Tectonatica dalll Cossmann, 1925. Un fortunately, (££.) tea land 1 was erroneously latinized with a masculine ending, even though it was named in honor of Nellie May Tegland. However, emendation of the ending to the feminine gender by Weaver (1942) Is unjustified, because Article 32(a)(11) of the International Code of Zoological Nomenclature clearly rejects improper latiniza- tion as grounds for altering the wcorrect original spell ing** of a name. Diocussioni This species is similar to Natica (Naticarius) posuncula Hanna & Hertleln, from which it differs by hav ing a narrower umbilical callus with a more gently curved margin, a less distinct funlcle, and a broader umbilical channel. Subgenus Lunala Berry, 1964 Lunala Berry, 1964i148. Type-speciesi Lunala lunarla Berry, 1964, by original designation. Recent, tropical eastern Pacific. Figured herein. Diagnosisi Shell medium in size, globose, thlni spire low to moderately elevated. Axial sculpture of short, retractive axial grooves extending from suture part way 276 to periphery on early adult whorls, usually absent on later adult whorls* Umbilicus very narrow. Umbilical callus very slender, merely an elongate thickening along edge of inner lipi funicle distinct but low. Operculum with one spiral rib near outer margin. Discussioni This subgenus is characterized by its peculiar funicle, which is reduced to a narrow twisted vertical ridge along the inner lip. It is known only by the type-species. Natica (Lunala) lunarls (Berry, 1964) Plate 3, figures 19, 20 Lunala lunarls Berry, 1964i148-149. Natica (Lunala) lunarls (Berry), Keen, 197li477, fig. 869. Description! Color.--Shell light to medium brown, with grayish cast in Juvenllest base and umbilicus paler than rest of shell, usually white, narrow whitish band occurs below suturei may show white blotches on shoulderi white base sometimes bounded above and at aperture by narrow band of dark browni interior reflects exterior colors. Periostracum thin, light to medium yellowish brown, obscures shell colors in some individuals. Size.--Averagei height 22 mm, diameter 19 mmi larg- 277 eat specimeni height 27.7 mm* diameter 25.5 mm B>* S. Berry collection, paratype. Estero lastlota, Sonora, Mexico]. Shell Form.- -Shell relatively high, spire moderately elevated to lowj body whorl highly Inflated, slightly tabulate below suturei shell very thini nuclear whorls 3, smoothi postnuclear whorls 3| suture strongly impressed. Axial sculpture of fine, retractive grooves below suture that are best-developed on first 1% adult whorls, usually absent later on. Spiral sculpture of minute, closely spaced, slightly wavy costellae, three of which are more prominent at periphery of Juveniles. Parietal callus ex tremely thin, transparent. Inconspicuous, lacking anterior lobe, very lightly filling posterior apertural angle. Umbilicus open posteriorly, narrowi sulcus shallow) channel reduced to shallow groove alongside callus. Umbilical callus very narrow, merely a thickening of inner lip, fills anterior half of umbilicusi funicle low, in distinct. Basal lip thin. Operculum.— Calcareous, white, central area often brown from incorporated sediment grains. Outer surface sculptured with minute, weakly developed spiral costellae, and with one narrow rib near outer margin) rib is under cut on each side and separated by shallow, flattened groove from upturned outer margin of operculum. 278 Specimens Examinedi 32. Geographic Occurrence and Ecologyi Estero Tastiota, Sonorai Mexico (28°22iN)» to Palta, Peru (5°S). South of the Gulf of California, known only by one specimen from Panama and two from Paita, Peru [CAS, Frizzell collec tion]). Found on mud bottoms in depths of 13 to 46 meters. Type Localityi Off Estero Tastiota, Sonora, Mexico, in 14 to 25 fathoms depth (Berry, 1964). Type Materiali Holotype, S. S. Berry collection, Redlands, California. Discussion i The three heavier spiral costellae at the periphery, noted by Berry (1964), are only distinct on Juveniles. The operculum of this species has not previous ly been described. Subgenus Glvphep* rh*m* Rehder, 1943 Glvpheplthema Rehder, 1943i196. Type-speciesi Natica idlopoma Filsbry & Lowe, 1932. Recent, tropical eastern Pacific. Figured herein. Diagnosisi Shell medium in size, globose to slightly elongatei whorls slightly flattened below suture, distinct ly inflated, with short, sharply incised grooves that extend part way from the suture to the periphery and are 279 Indistinct on final adult whorl. Umbilicus broad* with cavernous sulcus and broad channel. Parietal callus thin. Umbilical callus small* semicircular* with a deep posterior notch that isolates the posterior margin as a low* sharp ribleti funicle robust. Operculum with several spiral ribsi outermost rib hollowi ribs have pustulate summits and 2 outermost ribs have pustules commonly elongated into rods that roof over the intervening grooves. Discussion i f l i vnhwni i-h»ma i8 characterized by having the posterior margin of its umbilical callus isolated as a low* sharp rlblet* and by its elaborately sculptured operculum with short rods joining the summits of adjacent spiral ribs. The name was applied by Rehder (1943) to two Carib bean and one eastern Pacific species with certainty. Natica (Glypheplthema) idlopoaa Pilsbry & Lowe, 1932 Plate 3* figure 21 Natica (Naticarius) idiopoma Pilsbry & Lowe* 1932>84* pi. 57 figs. 8-11. Natl^ Pilsbry & Lowe, M. Smith* 1944112* fig. Natica (Natica) ihinnnma Pilsbry & Lowe, Keen, 1958i320, rig. 2oI|Keen7 1971*475, fig. 864. fllYPlWPlthftMl ifllflftflf (Pilsbry & Lowe), Rehder, 19431196. 280 Description! Color.--Shell light to medium brown with 4 narrow spiral bands of white marked with medium brown spotsi uppermost band on shoulder* one at periphery! and two on basej one or both basal bands may be absenti shoulder be tween two upper bands is often darker brown than rest of shelli callus whitei interior white near outer lipt pale brown within. Periostracua thin, pale yellowish white• may obscure shell colors* Size■--Averagei height 15 an, diameter 14 mmi larg est speciment height 21*3 am, diameter 22.0 am CUSNM 4120, Cabo San Lucas, Baja California, Mexico]. Shell Fora.--Shell somewhat elongate, spire moderately elevated, body whorl inflated! whorls distinctly flattened at suturei shell thini nuclear whorls 2k, with weak, microscopic, closely spaced spiral costellaei post- nuclear whorls 3i suture strongly impressed. Adult whorls smooth except for retractive grooves extending from suture half-way to periphery, grooves are well-developed and closely spaced on early whorls, obscure and widely spaced on final whorli incremental axial growth lines are strongest below suture. Parietal callus thin, lightly filling posterior apertural anglei anterior lobe weak* Umbilicus widely open, deeply excavates posterior inner lip of aperturei sulcus broad, fairly deep, roundedi channel broadly open to inner lip. Umbilical callus 281 relatively snail* gently tapered anteriorly* with a deep posterior notch that Isolates the posterior margin as a low* sharp rlbleti funtcle robust* Anterior inner lip and basal lip slightly thickened. Operculum * --Calcareous. white* with 5 spiral ribs in adultsi innermost rib poorly developed* may appear as thickening of nucleus % second rib is largest* irregularly pustulatei third rib narrow* undercut on its outer sidei fourth rib narrow* set close to fifth rlbi a weak rlblet usually occurs between third and fourth ribs. Fifth rib hollow* contains narrow cavity formed by addition of thin wall to outer surfacei both walls of fifth rib have minutely pustulate summits* with pustules commonly elongated into rods that Join the two walls and roof-over intervening cavityi minute* Irregularly beaded costella runs along outer margin of operculum. Specimens Examinedi 129. Geographic Occurrence and Ecologyi Bahia San Quintin* western Baja California* Mexico (30°22'N) CAHF 488-36j, in the southern Gulf of California* and south to Puerto Utrla* Colombia (5°59’ N)* and the Galapagos Islands, Ecuador. Rare north of the southern tip of Baja California* or south of Panama Bay* Panama. Collected living In depths of 4 to 61 meters* most often in 18 to 35 meters, on bottoms of mud, sand and shell debris. 282 Type Locality i San Juan del Sur, Nicaragua (Pilsbry & Lowei 1932). Type Material! Holotype, ANSP 155437. Nomenc latural Commentary! (Glvpheoithema) Ulwgaa was designated the type-species of G1vphepithema by Rehder (1943), who also included Atlantic, Caribbean and Indo- Pacific species in the genus, was founded upon the opercular features of (G.) idlopoma. although Rehder did not describe the operculum in detail or con trast it to opercula of other species. Although (G.) Idlopoma has the most elaborately sculptured operculum of eastern Pacific natlcids, opercular sculpture alone is an insecure basis for generic distinctions in this family, because it may vary widely within one species, and because it shows no consistent relationship to shell features of accepted generic significance. Fortunately, as described here, there are shell features of (G.) idlopoma that distinguish it from other natlcids and place it in a separate subgenus. Discussioni This species is readily identified by the deep notch on the posterior side of the umbilical callus, its color pattern and tabulate whorls, and its highly sculptured operculum. The slender rlblet distinct from the main mass of the umbilical callus is unique among 283 eastern Pacific natlcids. On some Individuals, this rib- let Is not developed strongly, and the umbilical callus has a more gradual anterior termination, giving the shell an atypical appearance. There are more spiral ribs on opercula of Juveniles than on those of adults. In young shells, the massive second rib Is lacking and replaced by 4 or 5 narrower ribs, which later coalesce at their crests to form a single rib. Subgenus Stlamaulax Morch, 1852 Stiamaulax Morch, 1852*133. Type-species* Nerita sulcata Bom, 1778, by subsequent designation (Harris, 1897), Recent, West Indies. Diagnosis * Shell small to medium in size, globose to slightly elongate, body whorl distinctly inflated. Sculpture of sharply incised axial grooves that extend from suture to base, sometimes also with weaker spiral grooves to form a reticulate pattern. Umbilicus broad, with deep sulcus separating umbilical and parietal calli. Parietal callus thick. Umbilical callus semicircular, with concave anterior surfacei posterior portion of callus distinctly extended as a low swelling) funicle massive. Inner lip sinuous* Operculum with several small ribs on outer portion and more massive, elevated inner ridge form- 284 ed of coalesced ribs. Discus slcni Stlftmaulax Is characterized by Its axial or reticulate pattern of sharply Incised grooves, Its broadly open umbilicus with deep sulcus■ and Its operculum with a massive Inner ridge. It Is an exclusively tropical sub- Nat lea broderlplana Recluz. 1844i205, Philippi, 1853i134- l357pl7l97flg. 2, pi. 17, fig. 5 (plate 1851)) Reeve, I855ipl. 15, figs. 66a,bi Sowerby, 1883i80, pi. 7, fig. 91i Doll, 1909(235) Jordan, 1936tl63| Hertleln & Strong, 1955i284-285, pi. 3, fig. 8. Natica (Stiamaulax) broderlplana Recluz, M. Smith, 1944i 12, fia. 126 Tas "N. (S.) broderlplana Reeve"3i Keen, 1958*321, fig. 263) Parker, 19647l§3, pi. 4, fig. 3) Keen, 1971*477, fig. 870. "Natica alapapillonls (Rodlng)", Carpenter, 1864(624 fnot H. |^py)lllonls (Rodlng. 1798), Recent, Indo- Natica lostoma Menke, 1847(178, Tryon, 1886(21, pi. 4, rig* 67 Cas synonym of J j . alapapillonls var. broderl- EiSQaJ. genus. Natica (Stiamaulax) broderlplana Recluz, 1844 Plate 2, figures 22, 23 / Natica ala-paplllonls var. broderlplana l586i2l7 pi. 4. figs. 65-67. Recluz, Tryon, zonaria LansrcK. 1010. ■ n. aiaoaDLiranLS ikdoiiu plana var. lostoma Menke, Philippi, 1851i120 285 Watlea taslel Recluz, 1853i53-54, pi. 2, figs. 11, 12, Tryon, 1886t21, pi. 4, fig. 66 Fas synonym of f j . alapapillonls var. broderl p Una T. f is a s u c lB L is m * Color.--Shell buff to medium orange-brown, with three spiral bands of alternating white and purplish browni middle spiral band divided into wider upper portion and narrower lower portion) base whitej narrow white band occurs Just below suture in darker colored shellsi callus white) interior white. Ferlostracum very thin, pale yellowish white, inconspicuous. Size. - - Average i height 19 aim, diameter 17.5 mmi larg est specimen* height 21.4 mm, diameter 19,9 mm CAHF 945- 39, Secas Islands, Panama3. Shell Form.--Shell elongate, spire moderately elevated to high) body whorl inflated, aperture largei whorls even ly rounded* shell moderately thick) nuclear whorls 2\% smooth) postnuclear whorls 3) suture strongly impressed. Spiral sculpture of very weak, minute, closely spaced costellae that are slightly wavy. Axial sculpture of sharply incised grooves extending from suture to umbilicus, which are strongest on shoulder and base* low, rounded riblets formed between grooves at suturei axial Incremental growth lines also present. Parietal callus fairly heavy, thickly filling posterior apertural angle, acutely tapered anteriorly) anterior lobe inconspicuous. 286 Umbilicus broadly open, with large sulcus and a broad channel tapering to a groove anteriorly. Umbilical callus large, broad, semicircular in outline, with concave anterior portioni very low ridge on posterior side of callus is reflected as bump on callus margini funicle large, filling anterior half of umbilicus, with a sharply incised groove along its anterior margin. Anterior inner lip and basal lip thickened. 2BS££UlU!B»--Calcareous, white, usually with pale gray or brown along median rib. Central area depressed, slightly concave, smooth, with distinct blister-like swelling covering its earliest-formed portion. Outer sur face sculptured with numerous spiral ribs, several of which merge to form a relatively narrow, sharply crested median ridge with a pustulate surfacei three outer ribs flat-topped, undercut by Intervening groovesi outermost rib narrow, sharp, minutely beaded on inner side, often connected to adjacent rib by calcareous roofing of inter vening groove. Outer margin with 4 to 5 minute, finely beaded riblets. Specimens Examined* 239. Geographic Occurrence and Ecologyt Cabo San Lucas, southernmost Baja California, Mexico (22°45'N), throughout the Gulf of California, and south to Manglar Alto, Ecuador <1°5S'S). Hertleln & Strong (1955) give the northern 287 range limit at Cedros Island (28°N) and the southern limit at Lobltos, Peru (4°39*S). Found alive in depths of 2 to 70 meters, mostly between 15 and 45 meters, in sand and mud. Stratiaraphlc Occurrencei Pliocenei Costa Rica, Panama and Ecuador (Hertlein & Strong, 1955). Pleistocenet Magdalena Bay, Baja California (Jordan, 1936), Islas Ires Marias, Mexico, and Panama (Hertlein & Strong, 1955). Type Localitiesi Natlea broderiolana - "Xipixapi, West Columbia" Jipljapa, Ecuador , 29 meters, sandy mud (Recluz, 1844). / Nat lea loatfiM - Mazatlan, Mexico (Menke, 1847). Natlea taslei - Mazatlan, Mexico (Recluz, 1853). Type Materiali Natica broderiolana - Unknown, presumably in Museum d'Histoire Naturelle, Geneva, Switzerland (Dance, 1966). Natica ioatoma - Unknown, sold to a dealer (Zilch, 1958i53). Mli£§ taslei - Unknown, presumably in Museum d' Hlstoire Naturelle, Geneva, Switzerland (Dance, 1966). 288 Nomenclatural Commentaryi This species was often con fused with the Indo-Pacific species f ) . alapaoilionis (Roding* 1798) by early workers* because of similar color patterns. However* alapapilionis lacks the axial grooves from suture to umbilicus that places N. (£.) broderiolana in the subgenus StUmaulax. Discussioni There is a functional relationship between the sculpture of the operculum and the shape of the umbilicus and umbilical callus in this species. When the animal is extended from the shell* the spiral median opercular ridge fits closely into the curved umbilical opening and the blister-like swelling over the origin of the operculum fits exactly into the concave anterior portion of the umbilical callus. The greatest width of the opercular ridge is the same as the width of the umbilical sulcus where it notches the inner lipi the opercular ri4&e thus rests in this sulcal notch and fixes the operculum in one position when the animal is extended. This relationship is also clearly seen in (£.) elenae Recluz* which has a closely similar operculum and umbilicus* and has not been previously described. The matching opercular and umbilical features may be a defense mechanism that maintains the operculum In the best position for rapid closing in case of danger. However* this would be an elaborate method of achieving a condition that would 289 probably have been selected for anyhow tinder normal living conditions. Rapid closing response is probably beneficial to any operculate marine snail, especially to active ones such as the naticids, and would likely not be dependent upon matching opercular and umbilical morphology. Among eastern Pacific naticids, this is seen only in N. (£.) broderlplana and £. (£.) elenae. This relationship may somehow be beneficial when the animal is plowing beneath the substrate in search of prey. The interlocked operculum and shell may provide a point for leverage when the animal is extending itself from its shell and wrapping its body around a victim. Similarly, the locked operculum and shell may be of help when maneuvering within the substrate, providing leverage for twisting its body to and fro while seeking out prey. This relationship between shell and operculum is also true for the Caribbean Miocene species (£•} guppiana (Toula, 1909), Judging from illustrations (Woodring, 1957, pi• 20, figs. 11-18). Natica (Stlgmaulax) elenae Recluz, 1844 Plate 3, figures 24, 25 Natica elenae Recluz, 1844*205-206i Philippi, 1853i135- 136, plT 19, fig, 4 (plate 1851)i Reeve, 1855ipl. 21, figs. 94a,b| Sowerby, 1883*79, pi. 9, fig. 150) Tryon, 1886*28-29, pi. 8, figs. 55, 56f Dali, 1909* 235i M, Smith, 1944*12, fig. 128i Hertlein & Strong, 1955*285-286, pi. 3, fig. 31. 290 Stiamaulax elenae (Recluz). Abbott, 1954ipi. 5, fig. mi Woodring, 1957*87• Natica (Stiamaulax) elenae (Recluz), Keen, 1958i321, fig. 254) Parker, 1964*153) Keen, 1971*477, fig, 871, Natica hanetl Recluz, 1850*389-391, pi. 13, figs. 6, 7| Tryon, 1886*29, pi. 8, fig, 56 Cas junior synonym of £. elenae!. Natica sulculosa Philippi, 1851*76-77, pi. 11, fig. 15 (plate 1850) Cplate caption as "N. elenae Recluz"}) Hertlein A Strong, 1955*286 Cas probable junior synonym of N. elenae1. Natica excavata Carpenter, 1856*165i Carpenter, 1857*282i Tryon, 1886*29 Cas synonym of £. elenae 1* Steams, 1894*195i Palmer, 1963*345, pi. 67, figs. 12, 13. Description* Color.--Shell with alternating, thin axial lines of pale yellowish white and reddish brown, which are usually irregular and discontinuous between suture and base, some times with gray tint, especially on later adult whorlsi umbilicus, base and narrow band at the suture are pale yellowish whitei aperture white tinted with violet or browni callus whitei nuclear whorls pale orange-brown. Perlostracum very thin, pale yellowish white, inconspicuous. Size.--Average* height 29 mm, diameter 26 mmi largest specimen* height 34.3 mm, diameter 31.0 mm CLACM 66-18, Punta Gorda, Baja California, Mexico}. Shell Form.--Shell elongate, spire moderately elevated to hlghi body whorl inflated, aperture largei whorls even ly rounded to slightly tabulate) shell thick) nuclear whorls 2%, smoothi postnuclear whorls 3%i suture strongly 291 Impressed. Spiral sculpture of weak, minute, closely spaced costellae that are usually wavy. Axial sculpture of sharply Incised grooves running from suture to umbili cus, strongest at suture where they are separated by rounded rlblets, weakest at the periphery where they may be Inconspicuous, do not extend all the way to umbilicus on final V whorl of large adultst axial incremental growth lines also present. Parietal callus massive, thickly filling posterior apertural angle, acutely tapered anteri orly! anterior lobe inconspicuous. Umbilicus broadly open, with cavernous sulcus and a broad channel tapering to a sharp point anteriorly. Umbilical callus large, broad, semicircular in outline, with concave anterior portion! small bump at posterior end intrudes into umbilical sulcus! funicle massive, filling anterior half of umbilicus, with a shallow groove along its posterior side. Anterior inner lip and basal lip thickened. --Calcareous, white, sometimes with pale gray or brown at its center. Central area depressed, con cave. Outer surface sculptured with numerous spiral ribs, several of which merge to form a broad median ridge with a mostly pustulate surfacei central area bounded by this ridge may be smooth or bear up to 10 low rlbletsi outer ribs usually 3-5, of various sizes, two larger ribs may nearly roof-over groove between them. Outer opercular margin with 1 to 4 microscopic costellae that show some- 292 tines discontinuous beading. Inner margin serrated by ends of opercular ribs* with microscopic costellae along the inner edge and parallel to it. Specimens Examined! 96. Geographic Occurrence and Ecologyi Magdalena Bay. western Baja California. Mexico (24°30*N), to Santa Elena (2°10tS) / and the Galapagos Islands. Ecuador. There are few records from the Gulf of California, and the species is uncommon throughout its range. Rare lntertldally. mostly in depths of 15 to 70 meters, in substrates of mud and sand. Type Localitiesi Natica elenae - Santa Elena, Ecuador, in sandy mud at 6 fathoms (Recluz, 1844). Natica haneti - "Habit, la cdte de Bahia1 * (Recluz, 1850), probably Bahia, Ecuador (0°0.5'S), not Bahia, Brazil, as speculated by Hertlein & Strong, 1955. Natica sulculosa - "West Coast of Mexico or South America" (Philippi, 1851). Natica excavata - Panama (Carpenter, 1856). Type Materiali Natica elenae - Unknown, presumably in Museum de Hlstolre Naturelle, Geneva, Switzerland (Dance, 1966). Natica haneti - As above. 293 Wat lea sulculosa - Unknown, presumably in BM(NH) or Humboldt Museum, East Berlin (Dance, 1966). Watlea excavata - Holotype, BM(NH) reg. no. 196322 (Palmer, 1963). Womenclatural Commentaryi In his type description of g. / elenae. Recluz separated some specimens as MVar. B. ?,M and thought they might be a new species distinct from £ J , elenae. He may have had this in mind when he described the synonymous species £ 4 . haneti in 1850. Discussioni £ 4 . (£.) elenae is most similar in form to £ 4 . (^.) gupplana (Toula, 1909), which occurs in Middle Miocene to Lower Pliocene deposits of the Caribbean area (Woodring, 1957186-88). The latter species seems to differ from £ 4 . elenae by having a thicker shell, a slight ly more massive parietal callus, and by having an operculum with a more variable number of ribs. Considering the minor differences between some Natlclnae species, as in the HStlCA (Watlcarius) othello speclea-group herein, these differences may be enough to separate £ 4 . (£.) elenae and £ 1* (£.) gupplana. but further study, perhaps of a quantita tive nature, may place £ 4 . (£.) gupplana as a synonym of £ 4 . ( 5 .) elenae- Natica ( 3 ,) elenae is also related to £ 4 . rtnrwna Linnaeus, 1758, of the Recent Caribbean fauna, as well as to several Recent through Miocene Caribbean species. Along with other Recent eastern Pacific species 294 such as f i . (£,) broderlpiana Recluz, Pollnlces (Pollnlces) uber (Valenciennes), and £. (£.) pananaensls (Recluz), J J . Cg.) elenae provides a clear link to the Recent Caribbean natlcid fauna and to the ancestral Miocene fauna that gave rise to the Recent naticids of each region. In the Recent eastern Pacific fauna, (£.) elenae Is closest to £. (£.) broderiolana Recluz. The most ob vious difference between these species Is that J S . (£.) elenae has an axial color pattern, whereas (£•) broderiplana is decorated with spiral color bands. In form, £. (£.) broderi plana is distinct from £. (£.) elenae by having stronger axial grooves, a much less massive parietal callus, and a less cavernous umbilical sulcus, besides reaching only about half the adult height of £. (£.) elenae. Subgenus Tectonatlca Sacco, 1S90 Iectonatica Sacco, 1890i33. Type-speciesi Natica tectula Bonelll, 1826, by monotypy. Pliocene, Italy, Figured in Wenz (1941, p. 1041, fig. 2979). Diagnosisi Shell small to medium in size, globose to slightly elongatet whorls smooth, not greatly Inflated. Umbilicus narrowly open along margin of semicircular umbilical callus. Parietal callus thin. 295 Discussion* Tect(matlea la characterized by having a amooth shell and an umbilicus that Is narrowly open along the margin of a semicircular umbilical callus. The operculum of the extinct type-speclea is unknown, Tectonatlca differs from Crvotonatlca by having an open umbilicus, whereas that of the latter subgenus Is en tirely closed by the semicircular umbilical callus. Its geologic history extends into the Pleistocene of Italy and western North America, and it Is represented in Recent seas by two cool-water Japanese species. Natica (Tectonatlca) lanthoatoma Deshayes, 1841 Plate 3, figures 26, 27, 28 Natica Desha yes, 183913611 Philippi, 1852153- 54, pi. 8, fig, 8 (plate 1850)i Reeve, 1855tpl. 18, figs. 79a,b* Sowerby, 1883i82, pi. 4, fig, 52i Pils- bry, 1895i71t Kuroda & Habe, 1952t71. Natica (Cryptonatlca) lanthoatoma Deshayes, Dali, 1921i 164, pi, 147 fig. 12i Oldroyd, 1927*725, pi. 97, fig. 5* MacNef “ Cas "aff, 5* MacNeilet al., 1943*84, pi. 11, figs. 12, 14 mjT (Tectonatlca) frnthostoma Deshayes, Takl, 1937* 88-89, figs, 5, 6 Cflgs. ■ N. lanthostomoIdes Kuroda & Habe, 1949ji Burch, 1946*2* Kuroda & Habe, 1949*71, figs. la,b. Tectonatlca lanthoatoma (Deshayes), Habe, 1961*39, pi. 18, rig. 9f Habe, 19o4i60, pi. 18, fig. 9| Habe & Ito, 1965*32, pi. 8, figs. 10-11. Natlc^ c^auy^var^ lanth^toy^ Deshayes, Tryon, 1886*31, Natica aevera Gould, 1859*43, Gould, 1862*109* Kuroda & 296 Ha be, 19521711 Johnson, 19641149, pi. 16, fig. 14* Pollnlces (Natica) severa (Gould), Pllsbry, 1895i72. D e s c r i p t i o n i Color.--Shell light to medium pinkish brown, usually with narrow, vague whitish bands on shoulder, periphery and basei callus and adjacent part of base whitei nuclear whorls white on shoulder, medium brown belovi axial streaking of colors common, especially on early adult whorls. Interior nottled nedium brown and white. Perlo- stracum thin, pale yellowish white. Size.--Averagei height 53 ram, diameter 50 mmi larg est specimeni height 57.4 mm, diameter 50.0 ram (broken) CUSNM 210941, "Bering Island," north Pacific^* Shell Form.--Shell globose to slightly elevated* body whorl not strongly Inflatedi whorls evenly rounded except for narrow tabulation at suturei shell thlcki nuclear whorls 2%, smoothi postnuclear whorls 4%i suture moderate ly Impressed. Spiral sculpture of minute, weakly develop ed, closely spaced costellae that may be better developed Just below suturei axial sculpture of Incremental growth lines that are best-developed near suture and on base. Parietal callus thin, moderately fills posterior aper- tural anglei anterior lobe very weak* Umbilicus narrowly open at sulcus, closed anteriorlyt sulcus narrow, deepi channel reduced to a marginal groovei funicle large, fill- 297 tng anterior end of umbilicus. Basal lip thickened* Operculum. - -Calcareous. white, inner margin thickened) thickening greatest over nucleus) outer surface with minute, weakly developed spiral striations, usually strongest near outer margin. Specimens Examined* 173. Geographic Occurrence and Ecologyi Kamchatka to Hokkaido, northern Japan (44°N), and Vladivostok, U.S.S.R. (43°N). Reported from Korea by Habe (1950), The bathymetric range is not known. Stratlaraphlc Occurrencei Ranges from Lower Pliocene to Recent. Lower Pliocenei Empire Fm., Coos Bay, Oregon (CAS)) Montesano Fm., Grays Harbor County, Washington (CAS)) 7Montesano Fm., Montesano, Washington (CAS). Middle Pliocene to Pleistocenei Merced Fm., Wilson Ranch, Sonoma County, California (CAS). Lower Pleisto cene i Elk River Fm., Curry Co., Oregon (UCB). Tvoe Localitiesi Natica I f f f f l t h g f l t f f l l * " Kamchatka, U.S.S.R. (Deshayes, 1841). Natica severa - Hakodate Bay, Japan (Gould, 1859). 298 lyre Natica " Unknown, presumably in Ecole des Mines, Paris, France, or BM(NH) (Dance, 1966). Natica severa - Holotype, MCZ 169369. Discussioni Natica 1anthostoma has not previously been recognized as a fossil, but was in collections and labeled as "Natica cons or s Dali, 1909" Cnot Wood, 18481 renamed Tectonatlca dalli Cossmann, 1925, not Tegland, 1933i ■ clausa Broderip & Sowerby, 1829D. Typical fossil speci mens of f i . (X*) lanthoatoma have lower spires, a tendency toward smaller umbilical openings, and attain a smaller maximum size than typical Recent specimens, but are other wise Identical. The largest known fossil specimen CCAS 4, Empire Fm., Coos Bay, Oregon, lower Pliocene3 is 34,2 mm in height and 34.6 mm in diameter, compared to the largest Recent individual at 57.4 mm and 50.0 mm (broken), re spectively. No fossil opercula are known. Natica (Tectonatlca) lanthoatoma is known as an early Pliocene through early Pleistocene fossil in northern California, Oregon and Washington, but it does not occur living along those coastlines. Its modern distribution Is mainly in Japanese waters, but it is apparently not known as a fossil in Japan. It is possible that this species originated in the northeastern Pacific during the early Pliocene, and existed there through the early Pleistocene, 299 but ha a migrated westward in more recent times and reached Its greatest development In modem Japanese seas. Its absence In upper Pleistocene deposits of western North America may simply reflect the fact that most such deposits represent rocky nearshore habitats, in which (I.) lanthoatoma is unlikely to have existed. There is no analogous Arctic or northern Atlantic species. (!•) lanthoatomoldes Kuroda & Habe, 1949, of Japan, is closely related to £ J . (X.) lanthoatoma. The principal differences are that £. (X>) 1anthostomoldes has two shallow grooves near the outer margin of its operculum, and has a narrower umbilical callus, so that the umbilicus is open along most of the callus margin instead of only at its posterior end* The umbilical sulcus of (J.) lanthoatomoldes is much reduced in size compared to that (I*) lanthoatoma. Some specimens of (X*) Ian- thostoma have faint strlatlons along the outer margins of their opercula, but never distinct grooves. (X*) j£Q~ thoatomoldea is presumably a recently evolved variant of (I*) IllthffltTiri because the umbilical features of western American fossils are those of the latter species. Fossil and Recent specimens of £ ) . (X*) IflflthftBtftlf and Natica (Crvptonatlca) clausa are quite similar in general aspect, with equivalent sizes and nearly identical oper cula, although the closed umbilicus of f l . (£.) clausa clearly characterizes it. Given their similarities, and the fact that J J . (£.) clauaa la known In deposits of upper Miocene to Recent age, £ * . (I.) Ian t ho stoma Is almost certainly a descendant from early Pliocene stocks of f } . (£.) clausa, N a t i c a ( T e c t o n a t l c a ) s a t a o p e n s I s ( A d d i c o t t , 1 9 6 6 ) Plate 3, figure 29 Tectonatlca? satsooensls Addicott* 1966at639, pi. 77* figs. 8-9, Size.--Haight 20.7 mm, diameter 17,5 mm (holotype). Shell Form.--Shell globose* spire moderately elevatedt body whorl not greatly inflated) whorls evenly rounded except for slight flattening in narrow area below suturej shell of moderate thickness) whorls about 4i suture moderately Impressed. Spiral sculpture of minute costel lae i axial sculpture of Incremental growth lines. Parietal callus thin* moderately filling posterior apertural anglei anterior lobe weak. Umbilicus narrowly open along margin of broad* rounded umbilical callusi channel narrow, sulcus narrow but distinct) funicle robust. Anterior inner lip thickened. Operculum.-•Unknown. 301 StratUraphlc Occurrencei Upper (?) Pliocenei Montesano Formation of Weaver (1912), Wynoochee Valley quadrangle, Washington (Addicott, 1966a), Type Localityi USGS Cenozoic locality Ml545. Cut on south side of Still Creek logging road, 400 feet north and 2200 feet west of southeast corner section 5, T18N, R7W, Wynoochee Valley quadrangle, Washington (Addicott, 1966a), Type Materiali Holotype, USNM 649134. Discussioni This species is very similar to Natica (Tectonatlca) lanthoatoma Deshayes, which ranges from the Early Pliocene to Early Pleistocene of western America and is living in Japan. £fl£i££ (Tectonatlca) satsopensls ap parently differs by having a more widely open umbilicus along the margin of its umbilical callus. However, a larger series of specimens might show this species to Intergrade with £. (J.) 1anthostoma. Subgenus Crvptonatlca Dali, 1892 Crvptonatica Dali, 1892t362. Type-speciesi Natica clausa firoderip & Sowerby, 1829, by original designation. Recent and late Cenozoic, eastern Pacific. Figured herein. Diagnosisi Shell medium to large, globose to slightly 302 elongate) whorls smooth, not greatly inflated. Umbilicus completely filled by thick, smooth, semicircular callus. Operculum smooth or with single low rib at outer margin, outer surface distinctly concave. Discussioni CrvBtonatlca is characterized by its semi circular umbilical callus that completely fills the umbilicus. It is known by only two known species, both of which are treated here, and is a cool-temperate to Arctic group. Its geologic range extends through the Miocene. Natica (Crvptonatlea) clausa Broderip & Sowerby, 1829 Plate 3, figures 30, 31, 32, 33, 34, 35, 36 Natica clausa Broderip & Sowerby, 1829(372, Gray, 1839i 136, pi. 34, fig. 3, pi. 37, fig. 6( Lov&i, 1847i149i Middendorff, 1849i 419-421j Mlddendorff, 1851t208-210) Philippi, 1851(98-99, pi. 14, fig. 5| Reeve, 1855) pi. 20, fig. 88, pi. 25, fig. 113) Danlelssen, 1861i 31) Carpenter, 1864(661) Gould, 1841(238, fig. 167) Gould, 1870(342, fig. 612) Dali, 1874»251 Cas of BroderlpDi Sars, 1878i159-169, pi. 21, figs. 12a,b, 13, pi. 12, figs. la-ci Sowerby, 1883(96, pi. 4, fig. 48) Tryon, 1886(30-31, pi. 9, figs. 65, 67-69, 73) Dali, 1892(362) Pilsbry, 1895(72) Arnold, 1907(26, pi. 54, fig. 16) Arnold & Anderson, 1907(144, pi. 21, fig. 16) Arnold, 1908«353| Odhner, 1913(7, 14-23, pi. 3, figs. 1-3, 5-14, 16, 17, pi, 5, figs. 7-14) Arnold & Hannibal, 1913(590, 592, 594, 597, 598) Martin, 1916(229, 233, 239, 243, 257) Weaver, 1916i 22i Ha riser, 1921(672-674, pi. 56, figs. 1-5) Howe, 1922) sp. list) Keen, 1937(41) Keen & Bentson, 1944( 176) Smith & Gordon, 1948(198) Thorson, 1951(21-22) MacGinitie, 1959(90-91, pi. 1, fig. 10, pi. 12, fig. 8) Bousfield, 1960i17, pi. 2, fig. 24) MacFherson, 1971(56-58, pi. 3, fig. 9. 303 Natica (Cryptanatlea) clauaa Broderip & Sowerby, Arnold, l503.313-5l4, pi. 16, fig. 13* Moody, 1916*44i Dali, 1921*163, pi. 14, fU. Hi Oldroyd, 1924*161, pi. 3, fig. 3i Oldroyd, 1927*724, pi. 97, fig. 2i MacNell, 1957*109, pi. 13, figs. 12-13, pi. 15, fig. 19. Natica (Tectonatlca) clausa Broderip & Sowerby, Grant & Gale, 1931*797-798, text-fig. 11i Soper & Grant, 1932* 1063i Burch, 1946*26i Kotaka, 1962*134-135, pi. 33, fig. 17i Burch, 1963il52j Faustman, 1964*135-136. usa (Broderip & Sowerby), Woodring et al.. Crvptonatlca cla 1946T71i Add ddicott, 1966*4, pi. 1, fig. 13. Tectonatlca clausa (Broderip & Sowerby), Okutani, 1964* 395, pi. lV fig. 18, pi. 5, fig. /i Habe & Ito, 1965* 30, pi. 8, fig. 4f Okutani, 1966*17. Folvi^|ges clausa (Broderip & Sowerby), Weaver, 1916*216, Natica clausa forma normalis Middendorff, 1849*420. Natica clauaa forma elatlor Middendorff, 1849*420. Natica clauaa forma depreaaior Middendorff, 1851*209-210. Natlga clausa var. minor Philippi, 1851*99-100, pi. 14, Natica clausa forma tvolca Sars, 1878*159-160, pi. 21, figs7l2a,b, 13. ? Natica beverlii Leach, 1819*61 * Philippi, 1853*l46t Carpenter, 1864* 523• ? Hfltifia conaolldata Couthouy, 1838*89-90, pi. 3, fig. 14| Pnillppi, lwSivol. 1, p. 17, pi. 1, fig. 11. 7 Natica septentrionalis Beck in Moller, 1842*80. 7 septentrionalIs (Beck in Moller), Dali, 1874* Neverlta ruaaa Gould, 1859*43. Natica ruaaa (Gould), Gould, 1862*109i Dali, 1874*251i Dal1, 1886*2181 Tryon, 1886*531 Cooper, 1894*291 Keen, 1937*41. 304 Natica (Cryptonatlea) xyug (Gould), Dali, 1921 * 163» Oldroyd, 1927*725. Natica (Tectonatlca) ruaaa (Gould), Grant & Gale, 1931t 798* Burch, 194615/1 Kotaka, 19621135, pi. 33, fig. 18f Glen, 1959i183i Faustman, 1964i136, pi. 5, figs. 23-24. Tectonat^ga ^yaaa^(Gould)■ Habe, 1950i13* Habe, 1961i39, Natica operculata Jeffreys, 1885t34-35, pi. 4, figs. 7,7aj Tryon, 1886■ 31 Caa Junior synonym of f j . clausal. Natica (Cryptonatlea) conaors Dali, 1909*86-87, pi. 5, fig. 10, pi. 6, fig. 9i Grant & Gale, 1931*797 Cas Junior synonym of £. clauaa1. not Natica mlllepuncta var. consora Wood, 1848*148 Cfossll, u7 England. Tectonatlca dalli Cosamann, 1925*121 C new name of N. consora Dali, 1909, not Wood, 1848Jf Weaver, 1942* 350, pi. 67, fig. 30. not Tectonatlca dalli Tegland, 1933*138, pi. 14, figs. 8-12 COligocene, western U.S.A.D. Natica (Neverlta) convexa Nomland, 1916*86-87, pi. 7, figs. 1, 2i Grant & Gale, 1931*797 Cas Junior synonym of £ J . clausal* Keen & Bentson, 1944*176. Neverlta convexa (Nomland), Adegoke, 1969*169, Natica convexa Nomland, Nomland, 1917*221. Cryptonatlea aallmba Dali, 1919*351. WatiQldroydPt%S^1%5 (Dall)i Dal1' 1921'164< Natica sallmba (Dali), Keen, 1937*41. (Tectonatlca) aallmba (Dali), Burch, 1946*28. Euaolra acoamlta Dali, 1919*352. FoUnices (Eusplra) acoamltua (Dali), Dali, 1921*164| Oldroyd, 1927*724-7*71 Burch, 1946*29. 305 Cryptonatlea aleutlca Dali, 1919a1352 fnomen nudunli Dali, 1919b1251 fdescrlbedJ* Woodrlng & Bramlette, 1950163, 72, pi. 10, fig. li Winterer & Durham, I962i296. Natica (Cryptonatlea) aleutlca (Dali), Dali, 1921i164, pi. 14, fig. 10i Oldroyd, 1924i161, pi. 22, fig. 12* Oldroyd, 1927i726. aleutlca (Dali), Keen, 1937*41 Cas Junior synonym of U. clausal. (Tectonatlca) Burch, 1563*152. a) aleutlca (Dali), Burch, 1846i28i "Nerlta affinis Gmelln," Jeffreys, 1867*229 Cnot Natica afflnls (Gmelln. 1791)D. "Natica afflnls (Gmelln)** Cnot Natica afflnls (Gmelln, 1791)31 Philippi, 1851*86-87, pi. l3, fig. 3* Morch In Rink, 1857*80j Jeffreys, 1869*215-216, pi. 102, fig* 3i Jeffreys, 1885*35* Carpenter In Dawson, 1872* 392i Sars, 1878*160-161, pi. 21, figs. 14a,b, pi. V, fig. 16* Watson, 1886*430* Frlele & Grieg, 1901*70* Oldroyd, 1927*794. "Natica (Tectonatlca) afflnls (Gmelln)" Cnot Natica afflnls (Gmelln. 1791)J* Burch, 1946*28. Natica afflnls var. vlttata Jeffreys, 1877*318-319. not Nerlta vlttata Gmelln, 1791*3674 CRecent, northern ' s j . Atlantic. > c l a m _________ Cnot U. clausa var. occlusa Wood, 1848*146, pi. "Natica clausa var. occlusa Wood," Frlele A Grieg, 1901* 70Cnot f i . gjfluga ______ 12, figs. 4a,b* fossil, EnglandJ. Description* Color.--Shell exterior and callus white. Periostracum thin, pale yellowish white to dark orange-brown, most commonly pale yellowish brown* base commonly paler* pale band may occur on shoulder* axial banding of light and dark uncommonly seen on Juveniles. Interior white or pinkish to medium brown. 306 Size.--Average Arctic specimen! height 30 mm, diameter 28 mm. Average southern specimen i height 13 mm, diameter 11.5 mm. Largest speciment height 61.0 mm, diameter 54.0 mm (broken)CUSNM 635660, St. George Is land, Alaska}. Shell Form.--Shell globose, spire low to moderately elevatedt body whorl not greatly Inflated, flattened or slightly concave Just below suturei shell thin to thickj nuclear whorls 2, smoothi postnuclear whorls 3*fi suture slightly to moderately Impressed. Spiral sculpture of minute, very weak, slightly wave striaei axial sculpture of incremental growth lines. Parietal callus relatively thin at midpoint, thickens to moderately fill posterior apertural anglei anterior lobe lacking. Umbilicus closed. Umbilical callus smooth, semicircular, flat to slightly concavet margin blending smoothly into underly ing whorl or raised slightly above it. Anterior inner lip and basal lip thickened. Operculum.--Calcareous. white, shallowly concavei central area thickenedt margins smooth. Specimens Examinedi 2300. Geographic Occurrence and Ecologyi Clrcumboreal. In the eastern Pacific, south to San Diego, California (32°43*N)i in the western Pacific, south to Japan and Koreai in the western Atlantic,south to Cape Hatteras, North Carolina! 307 In the eastern Atlantic* south to Spain and perhaps the Mediterranean. Known on soft bottoms In depths from 9 to 970 meters, in progressively greater depths from north to south. Scratinraphlc Occurrencei Ranges from Lower Miocene to Recent. Middle Ollaocene to Lower Miocenei Poul Creek Formation. Yakataga area. Alaska (Clark. 1932t USGS). Lower Miocene (?)« lower Yakataga Formation, Yakataga and Malasplna areas, Alaska (USGS)i Big Sitkinak Island, Trinity Islands, Alaska (USGS). Middle Miocenei Astoria Formation, Newport, Oregon (CAS). Upper Miocenei Clerbo Formation, Pleasanton quadrangle, California (UCB)t un named formation, western Washington (Weaver, 1916). Upper Miocene to Lower Pliocene i Montesano Formation, Grays Harbor County (UCB), Humptulips quadrangle, Grisdale quadrangle, Aberdeen quadrangle (USGS), and Sylvia Creek (SU), Washington) Quillayute Formation, La Push quadrangle, Clallam County, Washington (USGS). Lower Pliocenei Empire Formation, Coos Bay (Dali, 1909i Arnold & Hannibal, 1913j Howe, 1922) Weaver, 1942, 1945) (UCB), Curry County (UCB), and Cape Blanco (Arnold & Hannibal, 1913), Oregon. Middle Pliocenei Merced Formation, Sonoma County (USGS) UCB), Santa Cruz area (Arnold, 1908), Bolinas Bay (Martin, 1916), Sebastopol quadrangle (CAS), Palo Alto quadrangle (USGS), and type Merced, San Francisco peninsula (Martin, 1916i Glen, 1959), Californiai Etchegoin Formation, CoalInga area, California (Nonland, 1916i Adegoke, 1969). Middle to Upper Pliocene» Rio Dell Formation, Humboldt County, California (Fauatman, 1964* UCB). Upper Pliocene Pico Formation, southeastern Ventura basin, California (Winterer A Durham, 1962)t San Diego Formation, San Diego County, California (LACMt UCB)t Fernando Formation, central Los Angeles basin (Moody, 1916i Soper A Grant, 1932f LACM) and Newport Bay (LACM), California. Pliocene Wildcat Formation, Humboldt County, California (Martin, 1916) UCB)i Purlsima Formation, Halfmoon Bay (Martin, 1916), Santa Cruz quadrangle (Arnold, 1908) UCB), Ano Nuevo quadrangle (SU), and San Mateo Mountains (SU), Californiai Quinault Formation, Taholah quadrangle, Washington (USGS)t Foxen Mudstone and Cebado Member of Careaga Sandstone, Santa Marla basin, California (Wood- ring & Bramlette, 1950). Pliocene to Pleistocenei Breldavic and TJomes sequences, Iceland (USGS). Lower Pleistocenei Timms Point Silt Member (Clark, 1931i Valentine, 1961t LACM), Lomlta Marl Member (Woodrlng et £l" 1946) LACM), and San Pedro Sand Member (Woodrlng ££ al., 1946| LACM) of San Pedro Formation, San Pedro, Cali fornia) Santa Barbara Formation, Santa Barbara, Cali fornia (LACMt UCB)t Elk River Formation, Cape Blanco, Oregon (USGSt CASi SU) UCB)j Anchor Silt, northwestern 309 Los Angeles basin, California (Rodda, 1957). Upper Pleistocenei Palos Verdes Sand, San Pedro, California (Arnold, 1903| Valentine, I96l| LACMf UCB)i Bahia San Quintin, western Baja California, Mexico (Valentine, 1961, as cf.). Pleistocene» Peard Bay, Alaska (Meek, 1923). Ire? LwaUUgg* Natica clausa - Unknown (Broderip A Sowerby, 1829). Natica clauaa forma normalla - "Ins. Sltcha" Cprobably Kruzof Island, northwest of Sitka, Alaska2 (Midden dorff, 1849), Natica clauaa forma elatlor - Novaya Zemlya Island, U.S.S.R. (Middendorff, 1849). Natica clausa forma depresslor - Unknown (Middendorff, 1851). Natica clausa var. minor - Ncoasts of the United States and Greenland" (Philippi, 1851). Natica clausa forma tvolca - Unknown (Sars, 1878). Natica beverlll - Baffin Bay, Canada (Leach, 1819). Natica conaolldata - "coast of New England" (Couthouy, 1838). Natica septentrionalla - Greenland (Moller, 1842). Neverlta ruaaa - Arctic Ocean (Gould, 1859). Natica ooerculata - North Japan (Jeffreys, 1885). Natica consora - Empire Formation, Coos Bay, Oregoni lower Pliocene (Dali, 1909). 310 Natica convexa - Etchegoin Formation, northeast of CoalInga, California} middle Pliocene (Nomland, 1916). Cryptonatlea aallmba - USFC station 4423, between Santa Barbara and San Nicolas Islands, southern California, 395 to 620 meters depth, sand (Dali, 1919). Eusolra acosmlta - USFC station 3128, off Monterey Bay, California, 1147 meters depth, mud (Dali, 1919). Cryptonatlea aleutlca - Unalaska, Aleutian Islands, Alaska (Dali, 1919). Natica af£lnla var. vlttata - Godhavn ^Greenland] (Jeffreys, 1877), Type Material! Natica clausa - Unknown, presumably in BM(NH) (Dance, 1966). Natica clausa forma normalls - Academy of Sciences, Leningrad (Dance, 1966). Natica clauaa forma elatlor - As above. Natica clausa forma deoresslor - As above. Natica clausa var. minor - Unknown, presumably in BM(NH) or Humboldt Museum, East Berlin (Dance, 1966). Natica clauaa forma tvolca - Unknown, presumably In Oslo Museum (Dance, 1966). 311 Natlca beverlll - Unknown, presumably in BM(NH) (Dance, 1966). Natlca consolldata - Unknown. Natlca septentrlonalls - Unknown, presumably In BM(NH) or Zoological Museum, Copenhagen, Denmark (Dance, 1966). Neverlta russa - Unknown (Johnson, 1964). Natlca operculata - Unknown, presumably In BM(NH) (Dance, 1966). Natlca consors - Syntypes, USNM 153917. Natlca convexa - Holotype, UCB 12064. Crvptonatlca sallmba - Holotype, USNM 209295. Eusplra acosmlta - Holotype, USNM 207218. Crvptonatlca aleutlca - Syntypes, USNM 217156 (Dali, 1919), also bear number 635660 on specimens and label. Natlca affinis var. vlttata - Unknown, presumably In BM(NH) (Dance, 1966). Nomenclatural Commentaryi The complex taxonomic history °f H* (£•) clausa has resulted from variations In size, shell thickness, spire height and umbilical callus morphology, plus broad geographic distribution. The sub species normalIs. elatlor and depresslor of Middendorff (1849, 1851), minor of Philippi (1851) and tvpica of Sars (1878) are based on Individuals with higher or lower 312 spires* or smaller size* than average specimens* A number of possible synonyms* such as beverlll. N. con sol idata and septentrlonalls. are poorly known in the literature and difficult to evaluate* but seem closer to clausa than to any other recognized species* Crvpto natlca sallmba and Euspira acosmita of Dali (1919) are based on specimens from off southern California that differ from typical northern specimens only by their smaller size* Pliocene specimens from California and Oregon have been referred to J j . convexa and £. consors. respectively. Natlca (Crvptonatlca) clausa may be a Junior synonym of Natlca afflnis of the northeastern Atlantic* but the problem is difficult to evaluate because descriptions of affinis differ among workers. The two species are similar in general shape* but afflnis is said to have an open umbilicus by some workers (Verrill* 1873* Tryon* 1886) and a closed umbilicus by others (Jeffreys, 1885i Harmer, 1921). Natlca afflnis was considered to be the southern white or semlpellucid form of f J * (£.) clausa by Odhner (1913)* and to be a nomen dublum by MacPherson (1971). Natlca clausa £££lufi£* an English Pliocene fossil, has a much more elevated spire than specimens of (£.) clausa I have seen, but the species are otherwise similar. A proper comparison of the two species would center on a study of shell form in £• clausa occlusa. to determine 313 whether or not the spire Is consistently elevated in large populations. The most recent taxonomic controversy surrounding | J . (£.) clausa has been its relationship to J £ . russa and £ £ . aleutlca. Natlca rusaa was described by Gould (1859) as having a thin shell and shouldered whorls. Natlca aleutlca was not originally differentiated from either russa or (£.) clausa, but was said to be larger than g. russa (Dali, 1919). More recent workers have most often considered J J . russa a thin-shelled species with a dark orange-brown perlostracum, £. aleutlca a large, thick- shelled species with a buff or pinkish color, and (£.) clausa a species of average shell thickness and pale yellowish brown color. Natlca russa was thought to be re stricted to the Bering Sea, £. aleutlca was recorded be tween Alaska and Puget Sound, Washington, and X J . (£.) clausa was recognized as far south as southern California. These forms are here considered to be conspeciflc because their characteristics intergrade to the extent that as signment to one specific name or the other is uncertain much of the time. Large, thick-shelled Recent specimens of £• (£•) clausa are known in the eastern Pacific from Alaska to Puget Sound, and the darker, thin-shelled form is known from the Bering Sea to Monterey, California. These extreme varieties of (£.) clausa seem to be developed to lesser degrees in other areas. 314 Discussioni Variations in size and shell morphology of (£.) clausa have been exhaustively documented by Odhner (1913) and confirmed in the present study* Specimens from the Bering Sea are larger than those from elswherei the largest Bering Sea specimen has a height of 61.0 mm and a diameter of 54.0 mm (broken), whereas the largest speci mens from elsewhere arei western Pacific, 29*7 mm X 27*1 mm USNM 205686, Sea of Okhotsk j western Atlantic, 36,2 mm X 31.1 mm USNM 600921, Labrador t eastern Atlantic,, height 30.0 mm Odhner, 1913j Berufjord, Norway . Compar ing Arctic and southern forms, the number of adult whorls is constant despite great differences in size, showing that the southern specimens are dwarfed and not merely Juveniles. In the eastern Pacific, an average Arctic specimen measures about 30 mm X 28 mm, whereas one from southern California measures about 13 mm X 11.5 ram. Southern forms also tend to be lighter in color than northern ones. The dark orange-brown specimens are known mainly from Arctic seas, with rare occurrences in the eastern Pacific south to Monterey, California. In addition to being much smaller, southern specimens of a. (£.) clausa live at progressively greater depths. Specimens live as shallowly as 9 meters in Arctic Alaska, but no shallower than about 150 meters off of southern California. Odhner (1913) noted that individuals from deeper water tend to have higher spires and shorter 315 apertures, although 1 have not seen this among eastern Pacific specimens. Odhner also noted that the radula was broader in shallow-water fonts and narrower in deeper- water forms, showing changes similar to those of the shell. However, his observation is based on only 5 speci mens, and may not be generally true* Northern specimens are also larger among fossils. In the abundant Pacific Coast Pliocene populations, the largest specimens (up to 47 mm X 46 mm) are from northern Washington, whereas the largest southern California Pliocene specimens measure about 15 mm X 13 mm. Pleistocene and Miocene specimens from southern and central California are equally small. The earliest known fossils of £. (£.) clausa are from the Middle Oligocene to Lower Miocene Poul Creek Forma tion, Yakataga area, south-central Alaska, which seems to confirm the anticipated northern origin of this species. The only other known lower Miocene fossils are also from Alaska, whereas Middle Miocene specimens are known in Oregon, and upper Miocene and later specimens in northern California. The species thus took a considerable time to extend its range southward, presumably in response to progressive marine cooling from Miocene through Pleisto cene time. The Pleistocene southern range-limit seems to have been at Bahia San Quintin, western Baja California, Mexico (Valentine & Meade, 1961), about 150 miles south of Its present southern limit, in response to late Pleistocene 316 cool-water conditions. Fossil occurrences In the western Pacific and Atlantic are not well-documented. The synonymy given above was initially intended to be exhaustive, but numerous European references have been omitted for lack of time to check them all. Odhner (1913) cites most of these. Natlca (Crvptonatlca) oreaonensls (Conrad, 1865) Plate 3, figures 37, 38, 39 Slaaretus scopulosus Conrad, 1849tapp. 727 tin part], pi. 19, figs. ob,c Lnot figs. 6,6a, ■£. scopulosus. or fig. 6d, -not determinable]. Lunatla oreaonensls (Conrad), Conrad, 1865i151. Natlca (Natlca) oreaonensls (Conrad), Dali, 1909i86, pi. 47 fig. 7. Natlca oreaonensls (Conrad), Arnold, 1908i349i Weaver, 19l2i2o, 22, Arnold & Hannibal, 1913i576, 583, 588i Weaver, 1916i172, 176. Natlca (Tectonatlca) oreaonensis (Conrad), Weaver, 1942i 331, pi. l05, fig. 27. Crvptonatlca oreaonensls (Conrad), Moore, 1963i27, pi. 2, figs. j , 4 [.not fig. 2, - J i * dark! Et he ring ton, 1931, or figs, 16, 17, « V J J Sowerby, 18293I Moore, 1971 fn by Ampu^ina^(Amurnn.l»^ oreaonensls Dali, I909i9l, pi. 3, Ammilllfta oreaonensls Dali, Weaver, 1912118. Amauropsis oreaonensls (Dali), Grant & Gale, 193li807| Weaver, 1942ij47-348, pi, 71, figs. 7, 11, 1931, or figs, 16, 17, * T J J . clausa Broderip & Sowerby, 18293I Moore, 1971i23i Addlcott, 1966i639, fnot pi, 77, fig, 10, 7- £, clausa Broderip & Sower- , 1829], 317 Descriptioni Size.— Averagei height 24 mm, diameter 19 nuns larg est apeelmeni heights 37.1 mm (Incomplete), diameter 29.6 mm (incomplete) CCAS 1, Coos Bay, Oregon. Empire Formation, Lower Pliocene3. Shell Form.--Shell elongate, spire elevatedi body whorl not greatly inflated, flattened to slightly concave near suturej shell thickness averagei whorls about 5i suture slightly to moderately impressed. Shell smooth except for minute incremental growth lines. Parietal callus average to thick, moderately filling posterior apertural angles anterior lobe weak. Umbilicus closed. Umbilical callus thick, semicircular, smooths margin evenly rounded, bounded by groove. Anterior inner lip slightly thickened. £B&E£UlUI*~-Calcareous, surface smooth, with a single low rib along outer margins central area slightly thicken ed. Specimens Examined> 65. Stratigraohlc Occurrence i Ranges from Middle Miocene to Lower Pliocene. Middle Miocenes Astoria Formation, Lincoln County (CASiSUsUCB), Curry County (UCB), Thurston County (UCB), Oregon, coastal Oregon (Moore, 1963), Seattle, Washington (Arnold & Hannibal, 1913)i Chehalis Formation, western Washington (Weaver, 1912). Lower 318 Pliocenei Empire Formation, Coos Bay, Oregon (CAS1UCB1 Dali, 1909). Iyge Lunatla oregonensls - Astoria, Oregon (Conrad. 1849) [Presumably from the Astoria Formation, Middle Miocene!• Ampul Una (Aaauropsls) oreaonensls - Coos Bay, Oregon. Empire Formation, Lower Pliocene (Dali, 1909). Type Materiali Lunatla oregonensls - Holotype, USNM 561551. Aapulllna (Aaauropsls) oreaonensls - Holotype, USNM 107780. Noaenclatural Commentaryt Natlca (Crvptonatlca) oreaonen- and Ampulllna oregonensls are considered synonymous here for the first time* The holotype of each species Is somewhat worn, but shows the elevated spire and simple callus covering the umbilicus that characterizes this species. Representative suites of specimens from the type areas of each species show similar ranges of morphologic variation. Discussioni Specimens are almost invariably decorticated, which has resulted in confusion as to the morphologic limits of the species. For example, Moore (1963) figured the holotype of (£.) oregonensls and also Included 319 figures of two other species under the sane name, and Addicott (1966) figured a probable specimen of £ 1 . (£.) clausa Broderip & Sowerby, 1829, as this species. Natlca (Crvptonatlca) oreRcnensls is the only species in the Neogene of the Pacific Northwest with an elevated spire and simple semicircular callus that covers the umbilicus. The operculum has not previously been found associated with the shell, as seen here on specimens from the Lower Pliocene Empire Formation of Coos Bay, Oregon* One speci men has the operculum perfectly in living position, another has the operculum slightly askew, and a third had the operculum lying loose in the shell. The operculum has a smooth outer surface, and looks much like that of (£•) clausa, except for the single low rib along the outer margin. Two opercula figured by Moore (1963) from the middle Miocene Astoria Formation are probably of this species, although their outer surfaces are concealed by matrix. If they lack a rib along their outer margins, they would then probably be of J j . clausa, which occurs un commonly in the Astoria Formation. The report of this species in the Ollgocene Blakeley Formation of western Washington (Weaver, 1916) is un reliable, considering the past difficulty in identifying this species and the considerable extension of stratl- graphic range that would otherwise be necessary. Natlca inezana Conrad, 1857 320 Plate 3, figure 40 Natlca Inezana Conrad, I857il95, pi. 10, figs, 5, 6* Anderson & Martin, 1914*43i Woodrlng, 1931t383i Moore, 1963129 £as "Natlca Inezana "1. Follnlces (Neverlta) Inezana (Conrad), Dali, 1909187-881 Weaver, 1942i342, pi. 70, fig. 15, pi. 101, figs. 7- 9, Hi Weaver & Klelnpell, 1963i101, 187, pi. 24, fig. 14. "7AiBPulllna (Ampul1Inopsla) mlaalaalpplensls (Conrad)," Dali, 1909bi90 Cnot A. (A.) mlsslsslpolensls (Conrad, 1842), Ollgocene, Mississippi]]. This species was briefly described by Conrad (1857) and Illustrated with a drawing of a very badly decorticated specimen completely lacking umbilical features. The type was collected at "Santa Inez," which Is taken to be Santa Ynez In Santa Barbara County, California, and was said to be "from Miocene beds." Dali (1909) reexamined Conrad's syntypes and concluded that the specimens, all poorly preserved, belonged to two genera, Neverlta and Ampullinop- sis. and that Conrad's (1857) figured specimen was a Neverlta. Dali also thought Inezana to be Identical with another Conrad species, ocovana Conrad, 1855, which 1 consider probably equal to £. anderaonl (Clark, 1918). Dali also reported £. Inezana as probably occurring In Miocene deposits at Astoria and Coos Bay, Oregon. Moore (1963) states that the name does not apply to any species 321 In the Astoria Formation. Weaver & Kleinpell (1963) designated Conrad's (1857) figured specimen as lectotype and recorded the species from the undifferentiated Gavlota Formation. of Oligocene age. in the Santa Barbara area. They agreed with Dali's (1909) opinion that two genera are represented by Conrad's type specimens, even though Woodring (1931) thought the supposed generic differences due to weathering. Miocene strata with shallow-water molluscan faunas do occur In the Santa Yhez Mountains (Dibblee. 1950). the type region for inezana. However, there are no Miocene species in the eastern Pacific with the low spire and inflated whorls of Conrad's figured specimen of £ 1 * Inezana. This species Is almost certainly a Paleogene one and probably an ampullinid natlcld. Dali (1909b) applied the name Ampulllna (AmpulllnoosIs) mlsslssiopiensls to specimens from Astoria. Oregon, the Santa Ynez Mountains. California. Port Blakeley. Washing ton. and Chichagof Bay. Alaska, in beds of supposed Oligocene and Miocene age. This species had previously been reported only from the southeastern U. S.. and its type locality is in Oligocene beds near Vicksburg. Missis sippi. According to Moore (i963t29), Dali's (1909b) specimen that supposedly came from Astoria, Oregon, is similar in matrix and preservation to specimens in the type lot of £• Inezana. Moore (i963) states that Dali's 322 specimen of (A.) mlsslflaiDpiensls supposedly from Astoria and the type specimens of £. inezana. from the Santa Ynez Mountains of California, bear the same lot number, USNM 3575. I have seen this number on Dali's specimen, but not on the types of £. Inezana. However, Conrad's specimens in the USNM are sometimes difficult to locate and identify, and 1 may have overlooked the Inezana types bearing the USNM 3575 number. Assuming Moore (1963) to be correct in identifying Dali's (1909b) specimen of (A.) misaissippiensls as coming from the type lot of f * . inezana. and given Dali's (1909b) vague description of the former species, Dali's specimen from the Santa Ynez Mountains and Astoria may be tentatively considered to be f t . inezana. Natlca saxea Conrad, 1849, nomen dublum Natlca saxea Conrad, 1849tappendix p. 727, pi. 19, figs. 7a,bj Dana, I909il55| Weaver, 19l6il72, 176* Moore, I963i28 fas nomen dublum1. Neverlta saxea (Conrad), Conrad, 1865«151. Polinices saxea (Conrad), Arnold & Hannibal, 1913i586, 588. Polvnlces (Neverlta) aaxea (Conrad), Reagan, 1909i171, 193- 194i Reagan, 1910i648. Natlca (Natlca) saxea Conrad, Etherington, 1931i93, not pi. 12, figs. 2, 3, 7, 14 Cflgs. are Polinlces vlctorianus Clark & Arnold, 1923]. 323 Natlca (Tec tonatlea) aaxea Conrad, Weaver, 1942*332, not pi. 68, fig. 4 Lfig. la Pollnlces vlctorlanua Clark & Arnold, 19233. As with other fossil natlclds authored by Conrad, U* saxea Is too briefly described and Is Illustrated with drawings of a poorly preserved specimen. The type speci men has apparently been lost. As noted by Moore (1963) 4 USNM specimens bear the number 3540 and are labeled as N. saxea. and another 4 specimens have USNM 3580 on the shells and USNM 3540 on a slip of paper In the vial. None of these 8 specimens matches the one figured by Con rad (1849). Call assigned specimens numbered USNM 3540 to Pollnlces (Eusolra) aallanoi Dali, 1909, and to £. (Neverlta) Inezana (Conrad, 1857). Moore's (1963) relegation of { 4 , saxea to the status of a nomen Is reasonable considering the poor original indication and missing type specimen. The figure given by Conrad (1849) entirely lacks diagnostic features. The only clues to correct identification of the figured specimen are its height (about 33 non) and a triangular remnant of callus at the posterior end of the umbilical callus area. Decorticated specimens of £. (£.) aallanol commonly have a similar triangular remnant of umbilical callus In the same position, and 1 have not ob served other species weathering In this way. The type locality of H. ***** 1* Astoria, Oregon, and £. (£.) 324 Ra Hanoi is common in the Astoria Formation there* so it Is stratigraphieslly possible for the two species to be synonymous* However* evidence for combining the species Is still debatable* and 1 therefore consider saxea to be a nomen dUfrlW* Rejected and Indeterminate Taxa Natlca Renlculata Conrad* 1855t18. A prior name for Bruclarkia barkeriana (Cooper* 1894)* In the family Neptuneldae* as noted by Clark (1937i387). Natlca glabella Reeve* 1855ipl. 27* figs. 124a*b. Reported from Central America (M. Smith* 1944i120, but illustrated as a sinistra1 shell with an unknown habitat by Reeve (1855), and unknown In the eastern Pacific. / Natlca mooulnlana Recluz, I853i154-156, pi. 5. figs. 9-10. Reported to live on the west coast of America or on one of the Pacific Islands. It is not known from the region covered by this study. Natlca sp. Ind. of Carpenter (1857t450), figured by Brann (1966ipl. 50, fig. 571), is indeterminate and may be a protoconch. Pollnicea rapulun llal Pilsbry, 1931i436-437* fig. 5. Apparently a Miocene fossil reworked from formations that crop out along the shores of the type locality of Panama Bay. Pllsbry's (1931) type specimen had been Identified earlier as £. stanlslasmeunlerl Maury* 1917* 325 by Li (1930i249), who stated of this and other specimens that Mthis collection is a mixture of Lower Miocene and Recent shells, for the collector was unable to keep them separate under the circumstances." It might be a variety of £. atanlaUgiiftnnlprl. which occurs in Miocene deposits of Central America and the Caribbean and is illustrated by Woodring (1957). Lunatla pallidoldes Carpenter, 1872*585, is mentioned from MArctic Ocean, near Behring Sts., mud, 30 fm." lack ing a description or figure, it is a nomen nudum. Lunatla tenulllrata Carpenter, 1857»45l, from Mazat- ✓ lan, Mexico, is apparently a Trlcolia (family Phaslanel- lldae) and is regarded as indeterminate (Keen, 1971i902). A figure is given by Brann (1966ipl. 50, fig. 572), and according to Keen (1971) the holotype in the British Museum is worn and broken, lacking part of the spire and much of the body whorl, and measures 1.4 mm in height. Lunatla ap. Ind. a, b, and c, of Carpenter (1857i 451), reported from Mazatlan, Mexico, are indeterminate and apparently based on protoconchs. Figures of these three forms are given by Brann (1966ipls. 49, 50). PLATES Fossil specimens have been whitened with am monium chloride. 326 PLATE 1 Figure 1. Aaauroaala ialandlca (Gmelin). USNM 108988, Recent, St. Michael Sound, Norton Sound, Alaska. Height 35.2 rnn. Holotype of &. purpurea Dali. 2. Pollnlces (Pollnlces) uber (Valenciennes). AM4H 73742, Recent, Santa Elena, Ecuador. Height 23.3 mm. 3. Pollnlces (Pollnlces) uber (Valenciennes). LACM 69-8, Recent, Bahia San Luis Gonzaga, Baja Cali fornia, Mexico, Height 11.4 mm. 4. Pollnlces (PqU nices) panamaensIs (R&luz). AMttH 154135, Recent, Farfan Beach, Canal Zone, Panama. Height 40.9 mm. Umbilical callus has atypical swollen appearance. 5. Pollnlces (Pollnlces) oanamaensls (Recluz). AhttH 154110, Recent, Viqui Point, Vacamonte, Panama. Height 36.8 mm. Umbilical callus typically develop ed. 6. Pollnlces (Pollnlces) intemeratus (Philippi). SU 52210, Recent, Panama Bay, Panama. Helsht 26.9 mm. 7« Pollnlces (Pollnlces) Otis (Broderip & Sowerby). USNM 46544, Recent, Panama. Height 30,4 mm. Holo type of ftaft subfusca Dali. 327 328 Figure 8. Polinices (Pollnlces) otis (Broderip & Sowerby). LACM B-345, Recent* Isla Vlveros* Panama. Height 36.2 mm. 9. Polinices (Polinices) rayldus (Souleyet). USNM 538002, Recent, Isla Lobos de Tlerra, Peru. Height 46.0 mm. 10. Polinices (Polinices) ravidus (Souleyet). Basal view of specimen in Figure 9, showing circular umbilicus. 11. Polinices (Polinices) blfasciatus (Griffith A Pldgeon). LACM 69-8* Recent* Bahia San Luis Gonzaga* Baja California* Mexico. Height 46. 1 mm. 12. Polinices (Euaplra) aa Hanoi Dali. USNM 153916* Early Pliocene* Empire Formation* Coos Bay* Oregon. Height 53.5 mm. Syntype* with decorticated shell surface and umbilical callus. 13. Polinices (Euaplra) aallanol Dali. CAS 4* Early Pliocene* Empire Formation* Coos Bay* Oregon. Height 34.6 mm. 14. Pollnlces (Eusplra) aaHanoi Dali. UCB 12058. Early Pliocene* NJacalitosM CEtchegoinJ Formation* Fresno County, California. Height 38.0 mm. Holotype of Ngtlca orbicularis Nomland. 329 Figure 15. Polinices (Euaplra) dlabloansla (Clark). UCB 11595, Late Miocene, upper San Pablo Formation, Contra Costa County, California. Height 49.0 mm. Holo type. 16. Pollnlces (Euaplra) lewis11 (Gould). USNM 3903, Recent, Puget Sound, Washington. Height 126.4 non. Probable holotypet supposed female form. 17. Pollnlces (Eusplra) lewis11 (Gould). LACM uncata loged, Recent, Redondo Beach, California. Height 118.8 mm. Supposed male form. 18. Polinices (Euaplra) draconla (Dali). USNM 172859, Recent, Monterey Bay, California. Height 51.5 mm. Holotype. 19. Polinices (Eusplra) draconls (Dali). Basal view of specimen in Figure 18, showing broadly open umbilicus. 20. Pollnlces (Euaplra) palIldus (Broderip A Sowerby). LACM A.375, Recent, Petersburg, Alaska. Height 41.5 mm. 21. Polinices (Eusplra) pallldus (Broderip & Sowerby). LACM A.375, Recent, Petersburg, Alaska. Height 21.6 mm. An average Juvenile specimen, with umbili cus more restricted than in adults. 330 Figure 22. Pollnlces (Eusplra) palIldus (Broderip f i t Sowerby). CAS 15, Early Pleistocene, Elk River Formation, Curry County, Oregon. Height 19.5 mm. 23. Polinices (Eusplra) victorianus Clark f i t Arnold. UCB 30203, Middle Miocene, Temblor Formation, Kern County, California. Height 38.3 mm. Paratype. 24. Pnllnlces (Eusplra) victorianus Clark f i t Arnold. CAS 582, Late Miocene (?), Sooke Formation, Vancouver Island, British Columbia, Canada. Height 29.8 mm. Holotype. 25. Pollnlces (Eusoira) agutonus Dali, USNM 110566, Recent, off Punta Aguja, Peru. Height 25.9 mm. Holotype. 26. Polinices (Eusplra) crawfordlanus Dali. USNM 123044, Recent, Gulf of Panama, Panama. Height 14.1 mm. Holotype. 27. fgllnUgg (Eusplra) lltorlnus Dali. USNM 96481, Recent, near Galapagos Islands, Ecuador. Height 7.7 mm. Holotype. 28. Polinices (Eusplra) oardoanus Dali. USNM 123046, Recent, Gulf of Panama, Panama. Height 12.8 mm. Holotype. 331 Figure 29. Polinices sookenals Clark & Arnold. SU 245, Early Miocene (?), Sooke Formation, Vancouver Ialand, British Columbia, Canada. Height 14.6 mm. Sup posed holotype. 30. Pollnlcea (Hvoterlta) hellcoldea (Gray). USNM 609- 343, Recent, Bah£a Salinas, Costa Rica. Height 13.0 nun. 31 < Pollnlcea (Hvoterlta) hellcoldea (Gray). Basal view of specimen In Figure 30, showing broadly open umbilicus and umbilical callus as a flattened lobe. 32. Pollnlcea (Mammilla) caprae (Philippi). AM4H 110384, Recent, Santa Cruz Island, Galapagos Is lands, Ecuador. Height 26.8 mm. 33. Neverlta (Neverlta) nana (Moller). USNM 214083, Recent, San Diego, California. Height 11.8 mm. 34. Neverlta (HsXSrilS) kiikensls (Clark). UCB 11590, Late Miocene, lower San Pablo Formation, Contra Costa County, California, Height 32.1 mm. Para- type. 35. Neverlta (Neverlta) klrkenals (Clark). UCB 11594, Late Miocene, upper San Pablo Formation, Contra Costa County, California. Height 38.7 mm. Holo type of Natlca amoldl Clark. 332 Figure 36. Neverlta (Neverlta) polltlana (Dali). USNM 205653, Recent, Petrel Bank, Bering Sea, Alaska. Height 15.0 on. Holotype. 37. Neverlta (Neverlta) new species. LACM uncataloged, Recent, off central Oregon. Height 18.5 nun. Holo type. 333 PLATE 2 Figure 1. Neverlta (Gloasaulax) anderaonl (Clark). UCB 11212, Early Miocene (?), San Ramon Formation, Contra Costa County* California. Height 15.3 mm. Holo type. 2. Neverlta (Gloaaaulax) anderaonl (Clark). SU 2763, Middle Miocene, Olcese Sand, Kern River area, Cali fornia. Height 36.8 mm. 3. Neverlta (Gloaaaulax) reclualana (Deshayes). LACM R-498, Recent, Alamltos Bay, California. Height 57.5 mm. 4. Neverlta (Gloaaaulax) reclualana (Deshayes). LACM uncataloged, Recent, Monterey Bay, California. Height 25.1 mm. 5. Neverlta (Gloaaaulax) reclualana (Deshayes). ANSP 147585, Late Pleistocene, Palos Verdes Sand, San Pedro, California. Height 30.0 mm. Neotype of Pollnlcea (Neverlta) reclualanua altito Arnold, designated by Pllsbry (1929). 6. Neverlta (Gloaaaulax) reclualana (Deshayes). USNM 36633, Recent, Mprobably San Diego" CCalifornia] on label. Height 30.0 mm. Syntype of Pollnlces Neverlta) reclualanua UwrfMlttH 334 335 Figure 7. Neverlta (Gloaaaulax) reclualana (Deshayes). LACM P.465.62, Recent, Anaheim Bay, California. Height 32.2 mm. A typical specimen of the so-called altus form of this species. 8* Neverlta (Gloasaulax) lamesae Moore. USNM 563129, Middle Miocene, Astoria Formation, Lincoln County, Oregon. Height 22.0 mm. Holotype. 9. Neverlta (Gloasaulax) lamesae Moore. UCB 15498, Middle Miocene, Round Mountain Silt, Kern River area, California. Height 22.7 mm. 10. Callnatlcina oldrovdll (Dali). SU 6447, Catalina Island, California. Height 35.0 mm. Holotype. 11. Buibus fraRilis (Leach). USNM 50319, Recent, St. Peter's Bank, north Atlantic. Height 22.6 mm. 12. Chorlstes carpenter! Dali. USNM 123039, Recent, Gulf of Panama, Panama. Height 20.8 mm. Holotype. 13. Choratea new species. LACM uncataloged, Recent, off central Oregon. Height 13.9 mm. Holotype. 14. sinum cvmba (Menke). AM4H 155878, Recent, Caldera, Chile. Height 54.5 mm. 15. sinum cvmba (Menke). Apical view of specimen in Figure 14. 16. s<ntttm cvmba (Menke). Basal view of specimen In Figure 14. 336 Figure 17* Slnum cvmba (Henke). AJf'lH 119534* Recent * Pucasana, Peru* Height 35.6 am. The whorls are typically shouldered. 18. Slnum Rravl (Deshayes). LACM uncataloged, Recent* off northern Peru. Height 25.7 mm. 19. Slnum Rravl (Deshayes). Apical view of specimen In Figure 18. 20. Slnum Rravl (Deshayes). Basal view of specimen in Figure 18. 21. Slnum Rravl (Deshayes). CAS 12601, Recent, near Mazatl£n, Mexico. Height 13.9 mm. Holotype of £. cortez1 Burch & Burch. 22. Slnum scooulosum (Conrad). USNM 3553, Astoria, Oregon, presumably from Astoria Formation, Middle Miocene. Height 22.0 mm. Lectotype designated by Moore (l963). 23. Slnum (Conrad). USNM 57375, Recent, San Pedro, California. Height 22.9 mm* 24. Slnum ftyQPMl^THI (Conrad). Apical view of specimen in Figure 23. 25. slnum Arr.ru]^nynfl| (Conrad). Basal view of specimen in Figure 23. 26. Slnum perrlni (Arnold). USNM 164979, Middle Miocene, Topanga Formation, Topanga Canyon, Los Angeles County, California. Height 19.6 mm. Holotype. 337 Figure 27. Slnum deblie (Gould). MCZ 169117, Recent, La Paz, Baja California, Mexico. Height 6.0 mm. Holotype. 28. Slnum deblle (Gould). Apical view of specimen In Figure 27. 29. slnum debile (Gould). Basal view of specimen in Figure 27. 30. Slnum noveail Dali. USNM 170298, Recent, Isla Gorgona, Colombia. Height 16.2 mm. Holotype. 31. Slnum novesll Dali. Apical view of specimen In Figure 30. 32. Slnum novesll Dali. Basal view of specimen In Figure 30. 33. Slnum aanctiJohannia (Pllsbry & Lowe). CAS 33281, Recent, San Felipe, Baja California, Mexico. Height 15.5 mm. 34. £Lou& aanctiJohannia (Pllsbry & Lowe). Apical view of specimen in Figure 33. 35. Slnum aancti Johannla (Pllsbry & Lowe). Basal view of specimen In Figure 33. 36. Eunatlclna inaculota (Carpenter), USNM 11841, Recent, Acapulco, Mexico. Height 6.9 mm. Holotype. 37. Eunatlclna Inaculota (Carpenter). CAS 5557, Pleistocene, Magdalena Bay, Baja California, Mexico. Height 9.6 mm. Holotype of £• helml Jordan. 338 Figure 38. Eunatlclna Insculpta (Carpenter). AHF 1725-49, Re cent, near Cabeza Ballena, Baja California, Mexico. Operculum, with central portion calcified and margins chltlnous. 39. flqUgfl (Matlca) clartel Etherlngton. UCB 31996, Middle Miocene, Astoria Formation, Grays Harbor County, Washington. Height 17.0 tun. Holotype. 40. Natlea (Natlea) Inexnectans 01ason. USNM 701161, Recent, near Cabo Mala, Panama. Height 23.1 mm. Holotype. 41. Natlca (fla.tUa) lnexpectana 01s a on. Operculum of specimen In Figure 40. _ 339 • f t a PLATE 3 Figure 1* Natica (Natlea) slgillata McLean. LACM 1284, / Recent, Isabel Island, Galapagos Islands, Ecuador. Height 10.0 nun. Holotype. 2. Natica (Natica) sl&lllata McLean. SU 49428, Recent, Carmen Island, Gulf of California, Mexico. Height 20,4 mm. 3. Natica (Natica) yokes1 Addlcott. USGS M1801, Middle Miocene, Astoria Formation, Lincoln County, Oregon. Height 15.0 mm. 4. Natica (Natica) yokes1 Addlcott. USGS M1802, Middle Miocene, Astoria Formation, Lincoln County, Oregon. Height 12.3 mm. 5. Natica (Natlcarlus) chemnltzll Pfeiffer. LACM 69-8, / Recent, Bahia san Luis Gonzaga, Baja California, Mexico. Height 27.2 mm. 6. Natica (Natlcarlus) unlfasclata Lamarck. LACM 65- 24, Recent, Farfan Beach, Canal Zone, Panama. Height 30.6 mm. 7. Natica (Natlcarlus) othello Dali. USNM 46446, Re cent, Panama Bay, Panama. Height 22.0 mm. Holotype. 8* (Natlcarlus) Dali. AWH 90853, Recent, Gorda Banks, Gulf of California, Mexico. Operculum of adult lnd^j.dual, enlarged. 341 Figure 9. Natica (Natlcarlua) caneloenale Hertleln & Strong. LACM uncatalogedf Recent* off northern Peru. Height 30.0 nun. 10. Natica (Natlcarlua) caneloenala Hertleln & Strong. Operculum of specimen In Figure 9, enlarged. 11. Natica (Natlcarlus) collma Strong & Hertleln. CAS 6996* Recent* near Manzanillo* Colima* Mexico. Height 21.0 mm. Holotype. 12. Natica (Natlcarlus) scethra Dali. USNM 123048* Recent* Gulf of Panama* Panama. Height. 15.1 mm. Holotype. 13. Natica (Natlcarlus) brunneollnea McLean. A*f*H 163358* Recent, Santa Cruz Island* Galapagos Is lands* Ecuador. Height 32.6 mm. 14. Natica (Natlcarlua) brunneollnea McLean. Operculum of specimen In Figure 13, enlarged. 15. Natica (Natlcarlus) aravl Philippi. USNM 517737, Recent, Venado Beach* Panama, Height 13.0 mm. 16. Natica (Natlcarlus) gray! Philippi. USNM 517737, Recent, Venado Beach* Panama, Height 13.0 mm. 17. Natica (Natlcarlus) oosuncula Hanna & Hertleln. CAS 7084, Miocene, Temblor Formation* Kern County, California. Height 22.4 mm. Holotype. 18. Natica (Natlcarlus) tealandi Hanna & Hertleln. UCB 342 Figure 32215, Late Ollgocene, Blakeley Formation, Restora tion Point, Washington. Height 25.5 mm. Holotype. 19. Natica (Lunala) lunarIs (Berry). SU 9501, Recent, off Estero Tastlota, Sonora, Mexico. Height 24.3 mm. Holotype. 20. Natica (Lunala) lunarIs (Berry). Berry collection 34003, Recent, off central Sinaloa, Mexico. Height 16.4 ram. 21. Natica (Glyphepithema) idlopoma Pllsbry & Lowe. ANSP 155437, Recent, San Juan del Sur, Nicaragua. Height 9.6 mm. Holotype. 22. Natica (StiUSaaiilax) broderlpiana Recluz. AHF 943- 37, Recent, Islas Secas, Panama. Height 21.4 mm. *■ 23. Natica (StlRmaulax) broderlpiana Recluz. Operculum of specimen in Figure 22, enlarged. 24. Natica (StlRmaulax) elanae Recluz. LACM 66-18, Recent, Punta Gorda, Baja California, Mexico. Height 34.5 mm. 25. Natica (StlRmaulax) elenae Recluz. Operculum of specimen in Figure 24, enlarged. 26. Nfltifift (Tectonatlca) JftnthOflLaaft Deshayes. USNM 210941, Pleistocene (7), "Bering Island," north Pacific. Height 57.4 mm. 343 Figure 27. Natica (Tectonatlca) lanthostoma Deshayes. Operculum of specimen in Figure 26, enlarged. 28. Natica (Tectonatlca) lanthoatoma Deshayes. CAS 4, Early Pliocene, Empire Formation, Coos Bay, Oregon. Height 34.9 mm. 29. Natica (Tectonatlca) aataopenais (Addlcott). USNM 649134, Early (?) Pliocene, Montesano Formation, Grays Harbor County, Washington. Height 20.7 mm. Holotype. Reproduction of original figure in Addl cott (1966a, plate 77, figure 8). 30. Natica (Cryptonatlca) clausa Broderlp & Sowerby. USNM 217156, Recent, St. George Island, Alaska, Height 44.9 mm. An example of the so-called aleutica form, with a large, thick shell. 31. Natica (Crvptonatlea) clausa Broderlp & Sowerby. USNM 212050, Recent, Unimak Island, Alaska. Height 23.3 mm. 32. Natica (Crvptonatlea) clausa Broderlp & Sowerby. USNM 209295, Recent, between Santa Barbara and San Nicolas islands, southern California. Height 14.0 mm. Holotype of Crvptonatlea sallmba Dali. 33. Natica (Cryptonatlca) Broderlp & Sowerby. USNM 75636, Recent, Halifax, Nova Scotia, Canada. Height 20.5 mm. 344 Figure 34. Natica (Cryptonatlca) clausa Broderlp 6 c Sowerby. USNM 207218, Recent, Monterey Bay, California. Height 16.0 mm. Holotype of Euspira acosmlta Dali. 35. Natica (Cryptonatlca) clausa Broderlp 6 c Sowerby, USNM 188401, Recent, Behring Strait, Alaska. Height 32.0 mm. An example of the so-called russa form of the species, with a thin shell. 36. Natica (Cryptonatlca) clausa Broderlp 6 c Sowerby. UCB 7106, Late Miocene, Clerbo Formation, Pleasanton quadrangle, California. Height 27.1 mm. 37. (Cryptonatlca) oreRonensls (Conrad). USNM 561551, Astoria, Oregon, presumably from the Astoria Formation, Middle Miocene. Height 14.1 mm. Holotype. 38. Natica (Cryptonatlca) 9CT6<fflgngtg (Conrad). USNM 107780, Early Pliocene, Empire Formation, Coos Bay, Oregon. Height 18.5 mm. Holotype of Ampullina (Amauropsls) orettonensls Dali. 39. Natica (Cryptonatlca) (Conrad). CAS 3, Early Pliocene, Empire Formation, Coos Bay, Oregon. Height 16.0 mm. Calcareous operculum is in life position. 345 Figure 40. Natica Inezana Conrad. USNM 12359, Miocene (?), Santa Ynez, Santa Barbara County, California. Height 38.4 ram. Lectotype designated by Weaver & Kleinpell (1963). REFERENCES CITED 347 REFERENCES CITED Abbott* R. T.t 1954, American seashells. Van Nostrand Co., Princeton, New Jersey, p. 1-54, pis, 1-40, figs. 1- 100. Adams, C, B., 1847, Catalog of the genera and species of recent shells In the collection of C. B. Adams. Mlddlebury, Vermont, p. 1-32. ______, 1952, Catalog of shells collected at Panama, with notes on their synonymy, station, and geographical distribution. R. Craighead, New York, p. 1-334. Adams, H. and Adams, A., 1853-58, The genera of Recent Mollusca, arranged according to their organization. London, vol. 1-3. Adanson, M., 1757, Hlstolre naturelie du Senegal, p. 1- 190 (Voyage au Senegal), p. i-xcvl, 1-275, pis. 1-19. Addlcott, W. O., 1965, Miocene macrofossils of the southeastern San Joaquin Valley, California. U. S. Geol. 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Creator Marincovich, Louie Nick, Jr. (author)

Core Title Neogene to recent Naticidae (Mollusca : Gastropoda) of the eastern Pacific

Degree Doctor of Philosophy

Degree Program Geological Sciences

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag OAI-PMH Harvest,paleontology

Language English

Contributor Digitized by ProQuest (provenance)

Advisor Easton, William H. (committee chair), Bandy, Orville L. (committee member), McLean, James H. (committee member)

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c18-875761

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