F- Paleonto: Americ E na 1997 raphıca Begun in 1916 | NUMBER 59 DECEMBER 20, 1996 Systematics and Evolution of Cenozoic American Turritellidae (Mollusca: Gastropoda) I: Paleocene and Eocene Coastal Plain Species Related to “Turritella mortoni Conrad" and “Turritella humerosa Conrad" by Warren D. Allmon PALEONTOLOGICAL RESEARCH INSTITUTION Officers БӘКНӘШУЕВИЕ ее rcc re LM CONSTANCE M. SOJA IRSE NAGE PRESIDEN ЕР а SA САС ЛА E E JAMES E. SORAUF SECOND: VICEPRESIDENT iss tc it UG din SHIRLEY K. EGAN SEGRE EAR Wa е E АШЫ ПИ M He HENRY W. THEISEN А У Врла S tonto: META UNT MER аа, HOWARD P. HARTNETT [Боле ое сука O OS r T Е WARREN D. ALLMON Trustees BRUCE M. BELL (to 6/30/99) EDWARD B. PICOU (to 6/30/98) CARLTON E. BRETT (to 6/30/98) GARY ROSENBERG (to 6/30/99) WILLIAM L. CREPET (to 6/30/97) MEGAN D. SHAY (to 6/30/99) J. THOMAS DUTRO, JR. (to 6/30/99) CONSTANCE M. SOJA (to 6/30/97) SHIRLEY K. EGAN (to 6/30/98) JAMES E. SORAUF (to 6/30/97) ANTON J. 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Lawrence, KS 66044 U.S.A. | | CONTENTS Page ADS trac ISIN EEE LATAS, mode НИН Жаа. Ge poo de e eed ce Kel ue EEE: 7 КОЛЕН РИТА ee ED EEE аа Ба ку шшш 7 ПОСТО НОО sas ЛЫ ES E E a ЖЕ O E V cuu aod ii Е зә ЕРИЋ 8 Difficulties vitbi/Türtitellids e ОО Eee A Gon qms T MMC Au net E re prp ecce жып 8 ESTO ЙС ӨШ на ам eR мы as sar эта IM ти FOU AI Mr dure Eu cde uo Ms 9 Introduction to Systematic Paleontology SPECIE СОЈ E E Н О оа о ER EN ee ыл бш 12 SupraspecitctBovelu A PUE sti Re e REX EEE нн TEEN EEE N ee 13 Phylogenetic Analysis Methods INTO TUCU о ra ER N ee E UI MET LSA Сеи ЕНИ: 14 @haracters МЕЕН КЕ Rese eas T eam AAR Ша NEM Те тат A E T ЕИ 14 Gcogiaphy asa TPaxonomio Characters A er КЗ ee Cn 16 MOHO Meu ARIS ТК ИИ eat ар АИО aO Eu eer EESTI eet күү ra 19 (пати TEEN od АЛИ INN NUM АСПА coUe такт та DIU meee 20 БОШОО ан поена за ИУ А 5 EN oue N N N 21 Gntogely:0f Spitak Scalp rire а ln e sc mer НТ и ПИ e c MC Se ыш ee 21 КОЛОМ И OSs nen алата ауа A oem TIC TREE N TORNA 28 WHOENPIONIEH Mw Eo c dC a N Tue e ПРИ nier EIE RM 31 "pical/atid: Pkhunl Angles and Shell Gieometty ra ies Soe eee TEE NE IS EN OT ас 32 SIN MEX I за ИВА ОЬ Sue LS Ie xem deri ПА me 32 Adult Scülptule A a Ne Фа EN cuero Тт 33 BIOgeOpraphicieonirolw eee ee toe CN UU а esie ААЛА SEL ION, ESO TO cmm cen topi ее 2005, 33 АТАТ ТОО а РАИ реве ren 2 ca RE ИКЕМ ЛЗ зо M Suc qi aM лын 36 ida А Ву ие солу ОМАН TA aa С а MERI EUER T NEC o cocer V TER M ENTRY an 36 ыер иелле ыо а сше: д on aaa ac blo as Е 36 Relationships among Species Groups: а Clagistic Approach r. a bees aaa ес ees 36 Relationships within species Groups: a Siratophenstc Approach ran 39 ooreo росе k eE тее Екен шарлы ОЛ ыны Mine nate etre ure 22 2224 40 A PA EON ы И ЕО ааа uL ыле ccc c 44 ADOT VAOS Or ROD SAO S E ts AE D LS л уы узур. ш da e 44 БУБЕШ СЕТ ТЕ ЛЕККЕ у here у A Qe eT Oe T 45 Па GLEN er A e тент ee гора AER Те о RR ee ORI. EO 91 exppendixel Procedures ТОЗСО МЄ ЖОШО Analysis ОРИСИ E I S 101 бан БО locality REER EE Wee a ИН O rr и 103 Ep Dump mae ЫН ОП Жик ER RE T E E EE E RE TQ ben T ETE QUE 111 LEN OE SU мекен ссуда аға Sere ааа сети Sp 128 LIST OF ILLUSTRATIONS Text-figure Page 1. Relative importance of the four principal shell characters used in the classification of the family Turritellidae since the description OMNI OTE ndun en е оо 9 2. Protoconchs of fossil and Recent turritellid species of the tribe Zeacolpini from New Zealand, showing range of form............ 19 РО Mieasuremtenls made ОВУ ec Ате Бресте v e odio ние asics E OS ony ee SNP 20 4. Apex of a turritellid shell, showing methods used for counting and measuring protoconch whorls. ............................. 22 5. Scanning electron micrographs of well preserved apices of two turritellid species, showing the two apical sculptural types most common among species from the Paleocene and Eocene of the U.S. Gulf and Atlantic Coastal Plains. ....................... 23 б. Notation systems used for describing spiral ribbing IE INA 24 7. Ontogenetic development of spiral sculpture in turritellid species from the Paleocene and Eocene of the Coastal Plains........... 25 S ИСОЕВ ОАО CUBES I BOE SOLIS eror rad tdm 28 9. Guillaume's classification of turritellid growth line types, and their stratigraphic distribution in the Cretaceous and Cenozoic of MEU ДАШ ОНДЫ пи ико Aras ME C NE II MM E M eH 28 10 Clossitieation: ОР Борик line traces used. ен paper. ee NE eA 29 11. Variation in basal and lateral aspects of growth line trace within the family ТиггнеШдае...................................... 29 12. Growth line traces of turritelline species from the Paleocene and Eocene of the Gulf and Atlantic Coastal Plains. ............... 31 13. Classitication of adult -whorlprofiles in turitelline gastropods; оаа ИН 32 14. Three alternative cladograms showing possible relationships of four of the principal species groups of turritellid gastropods from Paleocene and Eocene sediments. of the U.S. Gulf and Atlantic Coastal Plains. „н a па ни 40 Io Phylogenetic tree өрішшиеше species: discussed. mtis paper: al oO 41 16. Three possible supraspecific classifications of Paleocene and Eocene Coastal Plain іштіеШйбв.................................. 43 В. Besuliscoffaetdr analysis of specimens of Turritella terebra: a nenn etc aieia Tode Paha du мани d ies it 58 18. Results of factor analysis of specimens of Palmerella mortoni Conrad sensu lato. ............................................ 59 19. Four possible evolutionary scenarios for the forms most closely related to Palmerella mortoni (Conrad). ....................... 62 20. Results of factor analysis: of specimens of Hanstatorcanmata.k Lea). «ois eeepc е а, PAID Re 70 ЖЕ Results O lector analysis ol Specimens Of HMoustaror perdita (COMA ne ee ТК e SH 76 22. results of factor analysis of spectmens:of Turritella” praecincta Conrado. nn... eR ee ec ра ee 87 LIST OF TABLES Table Page 1. Character states for supraspeciic taxa im the A aa a наны 10 2. Confidence estimates for date of first appearance of turritellid species from the Paleocene and Eocene of the Coastal Plain........ 16 3. Protoconch size and inferred mode of larval development for fossil turritelline species from the U.S. Gulf and Atlantic Coastal Plans and the four living species for which these data are available... и rege а aha een wd vse да 18 4. Character states for Paleocene turritelline species from Nigeria, described by Adegoke (1977).................................. 34 5. Inferred apical spiral sculpture formulas for Caribbean Neogene turritellid species based on data presented by Macsotay and Scherer MI aac N зе MI WP ccc mn T 35 6. Character states for Cretaceous and Cenozoic turritelline species from California, described by Merriam (1941). ................ 36 7. Described species of turritellid gastropods from the Paleocene and Eocene of the U.S. and their taxonomic placement in the present DHD IR a I Qa qu I C EMI t qe M eu RE cp a deed i SIE 37 8. Geographic range and stratigraphic duration of Paleocene and Eocene turritelline species from the U.S. Gulf and Atlantic Coastal > НАТ ток ск La ne) ra fey meta car NEM er 3 9. Character states for Paleocene and Eocene turritellid species from the Gulf and Atlantic Coastal Plains discussed in this paper. (SOT ИКО rset ам e E ПАКИ ВУ КИН A СРЕ СИ ONE ы іы ы en 45 EE NIE а О ЕИО ОДО o a e. ee ы Eel c e ced en 46 СИЕ СХЕ ООА ЕТЕП ТООТ ЕО irs ee 47 аата a O САДО (NZ and TOn). e ЕСЕ OS 48 ТОЗО A clerelandia CHAIS) а а. 49 EBEN TER SITE ite Л СООО АЕ) е И 49 Ша А IC ASIEN AAN ced ci edhe Es RE ER MM ни oe PGT en 50 in e М АЯП TO SCR (ELAS) a put. e reU TE ROT 51 К bec orl A Karma a si Lee ast ceret CIA gero lp A O у. 52 ПЕВА ОМАНУ E O A ea e o o TUER erase um mL E QU S 53 21. Messurements of Paimerella lisbonensis БЕС ri a eed T P MU ee, 54 E IA A ee 59 23. Results of factor analysis of specimens of the Recent species Turritella terebra (Linnaeus): variance explained by first five factor К ADM sees en с Ча 24 Жо NE ie Lea dine en E M 5 24. Results of factor analysis of specimens of the Recent species Turritella terebra (Linnaeus): rotated factor loadings of each variable on each of the first three factor axes. | | | | - = . Results of factor analysis of specimens of Palmerella mortoni (Conrad): variance explained by first five factor axes. ............. 60 . Results of factor analysis of specimens of Palmerella mortoni (Conrad): rotated factor loadings of each variable on each of the first MTGE FACTOR CS СЕРИЕН ЕРО M кат E ERU ОО е БОИ 60 a Measurements Of pulmerella mortoni mediavid (BOWIES) 522-2222... 2... 63 . Measurements of Palmerella mortoni postmortoni (Натгів)..........................................................2.....2.. 63 К Measurements:of Palmerelia mortoni ssp. Cpremoriont Сомов) «ee la Неле ај шз 64 . Measurements of Palmerella pleboides (Vaughan). ........ eee e ehh hene 65 . Measurements of Palmerella potomacensis (Clark and Martin). ............................................ ene 65 7 Measurements Ob АННЫ Т 22222770 ДЕКА ые ee 66 9 Measurements орао нео ПЕРЕН ОИ en 66 -Measurements of Aastat alabamiensis ONEN Е ОИЕ УТЕ 67 ""Measurementis:of Haustator carinata Q. Led)... un ee nl esee et ыс ie wien ne db a tie 69 . Results of factor analysis of specimens of Haustator carinata (1. Lea): variance explained by first five factor axes. ............... 70 . Results of factor analysis of specimens of Haustator carinata (I. Lea): rotated factor loadings of each variable on each of the first ТИТОВА у аттау а ЧАТ ла тсе Чат 71 Боа о ОТО ОНО e EL Ll LES Sala RES Ta . Measurements of Haustator fischeri (Раітет).....................................................................2.2.2. 024 T2 . Measurements of Haustator gilberti (BOWleS). на ааа ei aG T3 . Measurements of Haustator infans (Stenzel and Тштпег).........................................2........2........4..4Ҙ4аа.. 73 = Measurements. о austaror аен а Eee EAS ee O Een s nae Weegee oe 74 < Measurements:of Haustatorperdita (Conrad). e a а ет en ee а een 73 . Results of factor analysis of specimens of Haustator perdita (Conrad): variance explained by first five factors.................... 76 . Results of factor analysis of specimens of Haustator perdita (Conrad): rotated factor loadings of each variable on each of the first a сы косса ы eU con ee Ned Se ea A ica acea dal 77 2 Measurements;oh Haustaton nina almene ain ersi qd de. ee mn мый. 02 2000 T? : Measurements ој Haustator.vivurbana (СооКе) к au: ae ee se ауға A ну 78 . Measurements of Haustator subrina (Palmer)... се «ыыт onen mens en O Re a eed iuh sere nen 79 MMeasiinementsrotel austators aba СШТШ) en u CAR elt estet т 222-2222. 79 К Measurements.of-Zizustaron.tennesseensisı Gab) car ec М2 5 edo а oot Gr 80 2 Measurements ob Haustator. yaughanu (BOWLES) aeons «s de o a a Деб a (o ete 80 fy MeasırementsoßTäurmitellasalänchiaBowlester кес an re ae TIA ee OR. 81 " Measurements.of--Turritellas biboraensis (Gardner) зе а rar teens ek eee en 82 - Measurements of “Zurritella”.claytonensis. BOWLES. ccc. u... A ҮЕ Апей саула канан ашын a лы. 82 . Measurements of “Turritella” eurynome УУһийеа......................................1..2.2..................2........... 83 - Measurements of “Turritella” gardnerae LeBlanc... 2... vu. ce eee ee een nen кай ne nee eee 2ш 84 “Measurements oft ТОО humerosa@onrade. ы zen rare ee ee 84 - MeasuremenistQob. Теа Ж ub PUB Т eat Ver een ee ee eier 85 - Measurementsio£ иеа руде тен рта о ава ки tesa wisi ne И erase Sata 2 86 . Results of factor analysis of specimens of “Turritella” praecincta (Conrad): variance explained by first five factor axes. .......... 88 . Results of factor analysis of specimens of “Turritella” praecincta (Conrad): rotated factor loadings of each variable on each of the ЕРИ ПОРАСТА РВИ Т аған аа RON ОА Ты 89 . Measurements of “Turritella” praecincta уіксіпіенбіб............................2...... ...... 2... Ҙ4Ҙ4ӘМ9РӘ-. ІШ.Ш Ц Ше nennt 90 "Measurements o£ Zurritelia sp (CDrehumerosa GOVODl) И. 90 pe Measurements ok ИИИЙ lid ТОППН ПАН КЕС a der bos UO LOCUM Sd e e cmm dea 90 . Summary of samples used for morphometric analysis. ............................................................... 102 | SYSTEMATICS AND EVOLUTION OF CENOZOIC AMERICAN | TURRITELLIDAE (MOLLUSCA: GASTROPODA) I: PALEOCENE AND EOCENE COASTAL PLAIN SPECIES RELATED TO “TURRITELLA MORTONI CONRAD” AND “TURRITELLA HUMEROSA CONRAD” WARREN D. ALLMON Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850 ABSTRACT Gastropods of the family Turritellidae are among the most important components of macrofossil assemblages in the Paleocene and Eocene of the U.S.Gulf and Atlantic Coastal Plains, and are of considerable biostratigraphic utility, yet their phylogenetic relationships have remained obscure. This paper treats 52 of these turritellid species, focussing on those related to the common and well-known species “Turritella mortoni Conrad” and “Turritella humerosa Conrad". Of the shell characters available in these species, the early ontogeny of spiral sculpture is believed to be the most reliable homology and so the best indicator of phylogenetic relationship. Because of the relatively high completeness of the Coastal Plain stratigraphic section, and the relative paucity of shell characters in these species, a two-step approach to phylogenetic analysis is used. Atemporal cladistic procedure, considering supraspecific turritellid taxa from other times and geographic regions, is used for analysis of relationships of species groups. Stratophenetic methods are used for analysis of relationships of species within these groups. The genus Turritella Lamarck has long been recognized as a form-genus and taxonomic wastebasket, but the taxonomic diversity and subtlety of morphological difference in turritellids, at least in the U.S. Coastal Plain, have hindered development of a coherent supraspecific taxonomy. This paper represents a first attempt at such a taxonomy. Three species groups are recognized: species most closely related to Turritella mortoni Conrad are placed in the new genus Palmerella; species most closely related to Turritella rina Palmer are placed in the genus Haustator Montfort; species most closely related to Turritella humerosa Conrad are placed (temporarily) in “Turritella” sensu lato. Multivariate morphometrics are used, apparently for the first time in turritellids, to discriminate species and subspecies. In part with the aid of these methods, “Turritella mortoni Conrad" is divided into four chronological/geographic subspecies: Palmerella mortoni mortoni (Conrad), P. m. mediavia (Bowles), P. m. postmortoni (Harris), and P. m. ssp. By similar methods, “Turritella” praecincta Conrad is divided into two geographic subspecies: “Turritella” praecincta praecincta Conrad and “7.” р. virginiensis n. ssp. Two species are recognized as new: “Turritella” toulmini, and Palmerella stenzeli. ACKNOWLEDGEMENTS I am especially grateful to D.T. Dockery, S.J. Gould, R.D. Turner, and L.W. Ward for all manner of sug- gestions, support, assistance and advice at various stages throughout this project. Thanks are also due to R.L. Aiello, R.C. Eng, F.C. d'Escrivan, R.M. Ross and J.R. Tegan for their patience, help and support at crucial times, to T. Rice for scanning electron microscopy, to S.A. Schellenberg and especially J.R. Tegan for assis- tance with morphometric analysis, to D. Campbell, W. Fallaw, C. Garvie, and V. Zullo for sharing unpub- lished information and to P. Gan for assistance with indexing. I owe a special debt to P.J. Morris, without whose patience, skill and computer wizardry the mor- рћоте с analyses presented here would never have been possible. Any errors or misinterpretations in these analyses are of course wholly my own. Thanks are due J. Allen (Alexandria, La.), E. Ben- amy (Academy of Natural Sciences of Philadelphia), W. Blow (U.S. National Museum), M. Carman (Field Museum of Natural History), K. Boss (Museum of Comparative Zoology), C. Durden (Texas Memorial Museum), P. Hoover (Paleontological Research Insti- tution), R. Portell (Florida Museum of Natural His- tory), C. Smith (Geological Survey of Alabama), and R. Stanton (Texas A&M University) for making pos- sible loans of specimens from collections under their care. D.T. Dockery, S.J. Gould, R.S. Houbrick, and R.D. Turner read earlier drafts and made many valuable comments. This work was supported by grants from the Geo- logical Society of America and Sigma Xi, by funds from the Department of Earth and Planetary Sciences of Harvard University and the Department of Inverte- brate Paleontology of the Museum of Comparative Zoology, and by a National Science Foundation Grad- uate Fellowship. Publication was made possible by the Trustees of the Paleontological Research Institution. This paper is dedicated to the memory of Joe Houb- rick and Wally Fallaw. INTRODUCTION In abundance of individuals and diversity of species, gastropods of the family Turritellidae are among the most important macrofossils in many Cretaceous and Cenozoic deposits worldwide (Allmon, 1988a). No- where is this more true than in Paleocene and Eocene strata exposed along the Gulf and Atlantic Coastal Plains of the United States, where turritellids are known from almost all marine units. Some beds contain vir- tually no macrofossils other than turritellids, varying in abundance from rare to densely packed; others con- tain a diverse mollusk fauna in which turritellids are of varying importance (Allmon and Dockery, 1992; Allmon and Knight, 1993). Although usually no more than one or two turritellid species are present in the same horizon, several beds contain as many as four or five. Here as elsewhere in the world, the abundance, wide geographic ranges, and relatively short strati- graphic duration of many turritellid species have led to their widespread use as guide fossils (e.g., Stenzel, 1940; Sohl, 1977; Toulmin, 1969, 1977); together with ostreid and venericard bivalves, they are the among the most biostratigraphically important macrofossils in the region. The nature of Turritellidae as a group suggests, fur- thermore, that it may be fertile ground for studies of considerable general evolutionary interest. Turritellids appear to be an example of prodigious taxonomic di- versity but relatively little morphological disparity. Turritellid evolution has been a story of both abundant speciation and repeated development, in both time and space, of a relatively small variety of forms (Kotaka, 1978; Allmon, 1987, 1994). In spite of this abundance, diversity and evident biostratigraphic and evolutionary importance, how- ever, Coastal Plain turritellids are poorly understood. Although considerable attention has been given to their description and classification, attempts at phylogenetic analysis have met with only limited success, and with- out coherent and reliable phylogenetic hypotheses, studies of evolutionary pattern and process in this pro- lific group have not been possible. This paper has three principle objectives: 1) to propose a preliminary, test- able phylogeny for a large group of turritellid species from the Paleocene and Eocene beds of the U.S. Gulf and Atlantic Coastal Plains, specifically those related to the common, conspicuous and well-known species Turritella mortoni Conrad 1830 and T. humerosa Con- PALAEONTOGRAPHICA AMERICANA, NUMBER 59 rad 1835b, which may serve as a basis for future evo- lutionary and biostratigraphic studies; 2) to present new formal descriptions of these taxa, most of which have not been systematically treated in more than half a century; 3) to place these species in the context of turritellids from other times and regions and begin the task of erecting a valid and useful classification and phylogeny of the entire family. Of the 69 species and subspecies in the family named from the Paleocene and Eocene of the coastal plain (excluding the genus Mesalia) (Allmon, 1988b), this paper treats in detail 25 DIFFICULTIES WITH TURRITELLIDS Phylogenetic and evolutionary analyses of turritel- lids are difficult for a number of reasons. Very little is known about either the ecology or non-shell characters (e.g., opercular and radular form, soft-part anatomy) of living members of the family (see Allmon, 19882; Allmon et al., 1992, 1994). Therefore, in this respect, adequate comparative data do not exist for studies of fossil species. More importantly for the purposes of this paper, the simplicity of the turritellid shell limits the number of characters available for phylogenetic analysis of fossils. As noted by many authors over the years, the turritellid shell consists essentially of little more than a high-spiralled tube of roughly circular cross-section, ornamented externally by spiral cords and occasionally some minor axial sculpture. This pau- city of characters has several implications: 1) Phylogenetic analysis. So few morphological char- acters are available in the shell of turritellids, and those available are so simple, that the possibility of homo- plasy is difficult to discount for almost any proposed set of relationships based on shell characters alone. For example, although Merriam (1941) and Marwick (1957a) have placed great emphasis on growth line form as an important taxonomic character, both au- thors also raise the possibility that growth lines can be susceptible to homoplasy. Several workers have em- phasized ontogeny of spiral sculpture, but MacNeil (in MacNeil and Dockery, 1984, pp.48—49) states that “it is difficult to imagine" that species of similar external form and sculpture are unrelated even though they differ in early sculpture. As a result of this ambiguity, other criteria usually must be used to derive phylo- genetic hypotheses. 2) Definition of supraspecific taxa. With probably 2000 named species and subspecies, fossil and Recent (Allmon, unpubl. compilation), the genus Turritella, sensu lato, long ago ceased to be useful in the sense of facilitating data retrieval, yet efforts toward a mean- ingful subdivision have been hindered by lack of phy- logenetic knowledge. At least 35 generic or subgeneric names have been proposed for fossil or living species | | | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 9 in the subfamily Turritellinae alone (see Table 1), but few have come into general use. Definition of supra- Specific taxa in the group has in fact tended to depend on unstated or ill-defined taxonomic philosophy (All- mon, 1992a), and as a result no serious supraspecific evolutionary hypotheses have been proposed. 3) Biogeographic control. Without reliable phylo- genetic interpretation, biogeographic history cannot be interpreted; yet biogeography can influence prelimi- nary phylogenetic hypotheses by suggesting which spe- Cies should be included in an analysis. I explore this relationship here by making use of both morphological and biogeographic data independently and together in a comparative fashion. Given the paucity of characters in fossil turritellid Shells, it could be argued that a more appropriate meth- od of analysis for this group of animals would begin with exhaustive study of living taxa; fossil forms could then be viewed in the context of preliminary phylo- genetic hypotheses on this basis. I have not proceeded in this way for two reasons. The first is that the present is not an adequate guide to the past for many aspects of turritellid biology. Not only have ecological param- eters such as substrate and temperature preference pos- sibly changed (Allmon, 1988a, 1992b; Allmon and Knight, 1993), but the former pan-Tethyan distribu- tion of the group, like that of many cerithioid gastro- Pods (Houbrick, 1981, 1984a; Jung, 1987), contracted in the late Cenozoic to a current center of diversity in the southwestern Pacific (see Allmon, 1992b for further discussion). Although it is probably true that data from fossils will seldom overturn a phylogenetic topology established on the basis of living taxa (Patterson, 1981), Only the fossil record can document the actual course of evolutionary events in a clade’s history. In a group such as Turritellidae, which clearly has so substantial a previous history, Recent taxa do not provide suffi- Cient information to answer many important evolu- tionary questions. The second reason is that one must start somewhere. Studies of living turritellids are few (e.g., Allmon, 1988a; Allmon et al., 1992, 1994; Lieberman et al., 1993), and we know little about the biology of the great ma- jority of species. In any study, it would of course be ideal to know the outcome at the beginning. This is especially true in systematic work, where the result is So often greatly affected by the starting point; the char- acters and taxa included in a systematic study, partic- ularly of a diverse and widespread group, will often be influenced in part by the experience and background Of the systematist. But where does one gain such ex- Perience? This paper discusses some of the turritellid Species from one region during one interval of time. Adequate understanding of the phylogeny of these spe- Cies, however, depends on detailed knowledge of older > 5 9 о = 2 E 2 > £ 8 > i 9 г) 2 5 © © 3 o a = E v - 1960 - 1920 1 E | - 1880 - 1840 i 4 \ Г 1800 Text-figure 1.—Subjective estimates of the relative importance of the four principal shell characters used in the classification of the family Turritellidae since the description of the genus Turritella by Lamack in 1799. and younger species from elsewhere, knowledge that does not yet exist. I would like to know the morphology and distribution of all other turritellid species, but I do not. The hypotheses I propose here can therefore be no more than tentative, constrained by the very limited data that are available at present. I hope that they may serve as a basis for future systematic work in this important but neglected group of gastropods. HISTORY OF STUDY The history of systematic study of the Turritellidae has been well summarized elsewhere (Merriam, 1941; Allison, 1965, 1967; Allison and Adegoke, 1969), and only the main points will be touched upon here, es- pecially as they are relevant to the classification of Coastal Plain species. Several studies of Paleogene Coastal Plain forms have in fact contributed to the classificatory practice for other turritellids, and so the two stories are to some degree interrelated. Classification of the family can be usefully thought of as having proceeded through three partially over- ' lapping phases (Allison, 1967), during which more em- phasis was placed on one or another of the four most conspicuous conchological characters of the turritellid shell: whorl profile, form of the growth line, ontogeny of spiral sculpture, and protoconch form (Text-figure 1). Lamarck erected the genus Turritella in 1799 to con- tain the Recent western Pacific species Turbo terebra Table 1.—Character states for supraspecific taxa in the family Turritellidae. Taxa listed include those recognized by Marwick (1957a) and all supraspecific taxa described since that time. Character state information derived from Marwick (1957a), original descriptions and illustrations, and observations of specimens in the MCZ collections. А growth line? protoconch apical strat. sculpture basal lateral sinus whorl diam- no. shell Taxon! range formula sinus type depth apex angle profile eter* whorls size Subfamily Turritellinae : Archimediella Sacco, 1895 Oligocene C2 Ві АЗ 3 3 5 E P C-J L 5 M-L A. (Torculoidella) Sacco, 1895 Pliocene 7? 3 3-4 S B ЈЕ D-J 7 dy M-L Callostracum Smith, 1909 Recent |: 2 Y da 2 У, С 2 ? м Coeloconica Eames, 1957 Eocene ? ? 2? M A OR C-D ? ? M-L Colposigma Finlay & Marwick, 1937 Danian C2 ВІ АЗ 3 1 M A 7 С 5 3 M Colpospira Donald, 1900 Eocene-Rec. d Cl b3 A2 2 2 D B OR C-D-J L 15-2 VS-M C. (Acutospira) Kotaka, 1959 Eocene C1 B3 A2/C1 A2 6? 2 D B OR X-W E 1.5-2 VS-M C. (Platycolpus) Donald, 1900 Recent C3 В! A2/C2 B1 A3 2 2 M B P-OR C-D Е 1.5-2 VS-M Colpospirella Powell, 1951 Recent 2 3 4 M B OR C-J ? ? M Costacolpus Marwick, 1966 U. Cretaceous 2 1 4 5 B VB C ? ? 5 Cristispira Allison, 1965 M.-U. Eocene d C2 ВІ a3 4? 1 M A P J-V S 3-4 M-L Ctenocolpus Iredale, 1924 Miocene-Rec. CIBA 2 3 M B P J-U E 1-5 VS-M Gazameda Iredale, 1924 U. Mio.-Rec. c2 Bl a3 4 2 M B-C P D-J T 1.5-2 M-L Hataiella Kotaka, 1959 Olig.-Mio. СІ B2 a3 3 1 M B OR C S 2-3 M Haustator Montfort, 1810 Eo.-Rec. C1 B2 a3 4 2 M B-C P-OR J S-L 2-3 S-L H. (Kurosioia) Kotaka, 1959 Pleist. C1 B2 A3 4 1 M B P C-D S 3-5 S-M Idaella Kotaka, 1959 Miocene C2 B1 A3 3 1 M A-B OR C-D S 2 M Maoricolpus Finlay, 1926 Olig.-Rec. C2 B1 A3 3 1 M B-C P J S 2.54 S-VL Neohaustator Ida, 1952 Plio.-Rec. C1 B2 a3 3 1 M A-B P-OR C-J ? 2 M Nipponocolpus Kotaka, 1959 Recent 2 2 1? М? А OR C ie 7 E Peyrotia Cossmann, 1912 Miocene ? 3 4 M B P C-J ? 2 M-L Reymentella Adegoke, 1977 U. ? Paleocene dcbar 3? 2? S? B P-OR L-H S 2.5-3 S Sechuritella Olsson, 1944 U. Cretaceous ? ? 4 M? B VP D-X ? 2 ? Spirocolpus Finlay, 1926 Eocene СІ B2 АЗ 3 3 р B OR-OP C-J 5 2-3 M Torcula Gray, 1847 U. Eoc.-Rec. C1B2a3 4 1 M B Р U B 1-2 M T. (Bactrospira) Cossmann, 1912 Pliocene ? 4 4 M B P J-U ? Т м Т. (Eurytorus) Gardner, 1947 Miocene ? 2 2 2 ? 2 J-U 7 ? M Torquesia Douvillé, 1929 Cret.-Eocene? C1 B2 A3? 3 2 M A P-OR X-Q 5 2-3 M-L Tropicolpus Marwick, 1931 U. Oligocene Cl Bl a2 6 2 м B-C P X-U 2 1-3 E T. (Amplicolpus) Marwick, 1971 U. Pal.-L. Mio. СІ Bl a2 2 27 M A-B B X-U 2? 2-2.5 L-VL Turritella Lamarck, 1799 7 C4 13 B1 A2 2 4 5 B P-VP С-Ј 57 2 E Wyatella Adegoke, 1967 M.-U. Eocene d CI Bi 2 1 м А OR X-Y de 2 M Zeacolpus Finlay, 1926 Plio.-Rec. EZBIA3 3 2 M B-C P D-J ? 2 M Z. (Leptocolpus) Finlay & Marwick, 1937 E: Bal. ? 3 1 5 В Р D-J 2 ? M Z. (Stiracolpus) Finlay, 1926 Plio.-Rec. C2 B1 A2/C2 Bl АЗ 3 1 M B Р С-У y 2 M Subfamily Protominae Marwick, 1957a Protoma Baird, 1870 Mio.-Rec. C2 B1 A3 5 1 M A P J 2 2 M P. (Protomella) Thiele, 1931 Recent 9 5 p M A P J ? Y M 01 65 UAGWON 'vNVOPIHWYV VOIHAVADOLNOAV ТУ 4 Table 1.—Continued. as. dei growth line? protoconch E Strat. sculpture basal lateral sinus whorl diam- no. shell e Taxon! range formula sinus type depth apex angle profile? eter? whorls size E Subfamily Pareorinae Finlay and Marwick, 1937 i. Craginia Stephenson, 1952 U. Cretacous 2 ? 3 M A P C-J ? p M á Mesalia Gray, 1847 Pal.-Rec. C1 Bl a2 1 3 S B P-OR С 5 2-3 M = Motyris Eames, 1952 Eocene ? ? de S 7 2 С D ? M © Neodiastoma Cotton, 1932 Recent 7 1 3 S B OR С 2 7 M Q Pareora Marwick, 1931 Oligocene C2 B1 A3 1 1 M A OR E S > 3% VS 4 Sigmesalia Finlay & Marwick, 1937 Eocene ? 1 4 M B OR С 2 % M 4 Tachyrinchella Titova, 1994b Oligocene СІ ВІ a3? 7 4 M A B (e 2 ? м = Tachyrhynchus Mörch, 1868 Recent C2 Bl a3 1 4 5 B OR € ? 2 5 ж Woodsalia Olsson, 1929 Eocene 2 2 7 de de ? J de ? M-L ш Zaria Gray, 1847 Recent C1 B2 a3 1 4 M B ME С-Ј ? 2 VL E Subfamily Turritellopsinae Marwick, 1957a y Glyptozaria Iredale, 1924 Recent C1 ВІ 1 4 5 2 2 С 2 155 VS (С)! Kimberia Cotton and Woods, 1935 Recent C2 Ві АЗ e 2 Y ? 7 C. ? 3-4 VS PA Orectospira Dall, 1925 Recent 7 2 ? 2 de 2 H de 7 S-M = Turritellopsis Sars, 1878 Recent СІ B1 A2 7 ? ? ? de С 7 2 VS-S С) Subfamily Vermiculariinae 3 Vermicularia Lamarck, 1799 Plio.-Rec. cl b2 a3 1 4 M B P-OR D-J 5 >37 M-L y ! Supraspecific taxa as listed by Marwick (1957a) or as described since then. > 2 Growth line traces as represented in Text-figure 10. E 3 Whorl profiles as represented in Text-figure 13. 5 4 Protoconch diameter (РІ + P2): Small (S) = < ~250 um; Large (L) = > ~250 um. 2 5 Size (maximum observed shell length) classification follows that of Marwick (1971): Very Large (VL) = > 100 mm; Large (L) = 50-100 mm; Medium (M) = 20-50 mm; Small (S) = 10-20 mm; Very Small (VS) = <10 mm. к. 12 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Linnaeus, 1758. Lamarck’s generic description (1799, p.74) reads in full as follows: Cog. turriculée; l’ouverture arrondie, entiere, mais ayant un sinus au bord droit. (Shell turriculate; aperture round, entire, possessing a sinus at the outer edge.) Between 1810 and 1912 several generic names were proposed for species similar to T. terebra; as Allison (1967) has pointed out, these names were based largely on differences in whorl profile, external sculpture of adult whorls, and overall shell form. These include Haustator Montfort, 1810, Smithia Maltzan, 1883, Ar- chimediella Sacco, 1895, Torculoidella Sacco, 1895, Altavillia de Gregorio, 1908, Callostracum Smith, 1909, Bactrospira Cossmann, 1912 and Peyrotia Cossmann, 1912. As Allison has also noted, however, several other generic names were proposed during this period based on more detailed morphological criteria, especially form of the growth line. These include Proto Defrance in Blainville, 1824, Tachyrhynchus Mörch, 1868, Pro- toma Baird, 1870, Turritellopsis Sars, 1878, and Col- pospira Donald, 1900. In addition, Gray (1847) de- scribed two genera in the group (Torcula and Zaria), but without descriptive comments of any kind. Cossmann (1912) reviewed all previous work, and recommended that the form of the growth line be em- ployed, together with whorl profile and external (adult) sculpture, in classification of the group. As pointed out by Merriam (1941, p.30), Cossmann was the first to attempt explicit definitions of what characters should be used to delimit supraspecific taxa in the family. Yet, although he suggested the use of growth line form, he did not use this character in his classification. Guil- laume (1924) criticized Cossmann on this point, noting that forms with similar growth lines had been placed by Cossmann in different taxa, whereas forms with very different growth lines had been placed in the same taxa, largely on the basis of whorl profile and overall shell form. Guillaume arranged European Tertiary spe- cies into five groups based exclusively on growth line form. Dollfus (1926) criticized this reliance on a single character. In the 1920’s, a number of New Zealand workers (Iredale, 1924, 1925; Finlay, 1926, 1930; Marwick, 1931; Finlay and Marwick, 1937) began to emphasize the importance of the ontogeny of spiral sculpture, i.e., the order of appearance and change in relative strengths of the spiral ribs, particularly on the earliest whorls, in grouping fossil and Recent species into supraspecific taxa. These authors described additional supraspecific taxa, relying mainly on spiral ontogeny, but also with consideration of growth line, whorl profile and pro- toconch and opercular form. Taxa introduced by these workers include Gazameda Iredale, 1924, Glyptozaria Iredale, 1924, Ctenocolpus Iredale, 1925, Maoricolpus Finlay, 1926, Spirocolpus Finlay, 1926, Stiracolpus Finlay, 1926, Zeacolpus Finlay, 1926, Pareora Mar- wick, 1931, Tropicolpus Marwick, 1931, Colposigma Finlay and Marwick, 1937, Leptocolpus Finlay and Marwick, 1937 and Sigmesalia Finlay and Marwick, 1937. In the following decades, workers beyond New Zea- land also began to emphasize early sculptural (“арі- cal") ontogeny in their turritellid classifications. Ida (1952) and Kotaka (1959) developed notation systems similar to and partly derived from that devised by Finlay (1930), and modified by Marwick (1957b), for describing sculptural ontogeny precisely. Merriam (1941), studying fossil species from the Pacific coast of North America, and Palmer (1937; Harris and Palmer, 1947) and Bowles (1939), studying Paleogene species from the U.S. Gulf and Atlantic Coastal Plains, also emphasized the early ontogeny of spiral sculpture. These authors did not employ a notational system such as Finlay's, using instead more ambiguous verbal de- scriptions based on the roots “costate” and “carinate”, together with prefixes (e.g., “uni-”, “multi-”), to in- dicate the number of spirals. Although their emphasis on spiral ontogeny laid valuable groundwork for later study, the lack of precision associated with these verbal descriptions probably discouraged more detailed com- parative work with taxa from elsewhere. Allison (1965, 1967), Adegoke (1967), and Allison and Adegoke (1969) continued the tradition of Palmer, Bowles and Merriam in emphasizing early ontogeny of spiral sculpture in their work on New World fossil forms, but improved dramatically upon these earlier efforts by applying the Finlay-type notational system to apical ontogenies. This not only allowed some sys- tematic clarification (e.g., distinguishing the genus- group Torcula from similar forms; Allison and Ade- goke, 1969) and identification and description of un- described taxa (Allison, 1965; Adegoke, 1967), but more importantly the hypothesizing of specific evo- lutionary patterns of sculptural change (e.g., Allison and Adegoke, 1969). INTRODUCTION TO SYSTEMATIC PALEONTOLOGY SPECIES LEVEL Because turritellids are abundant and diverse in modern seas as well as in the fossil record, it ideally should be possible to make comparisons between pa- leontological and neontological species concepts in the group. To my knowledge, however, no studies have been done on intra- or interspecific variation in non- PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 13 shell characters of Recent turritellid species and, with the exception of Charles (1977), neither do detailed Studies of variation in conchological characters appear to exist. Thus, asisthe case with most gastropod groups (e.g., Allmon, 1990), fossil species in turritellids are really no more or less equivocal than are living species. In the species descriptions given in this paper, I have analyzed patterns ofshell form variation quantitatively in several taxa that seem to be especially variable and for which a sufficient number of specimens was avail- able, and I compare these patterns to presumed pop- ulation samples of a living species (the type species of the genus Turritella) represented in museum collec- tions. Although such a procedure only compares ranges of shell form and not actual species status, it does represent an explicit attempt (of the only sort possible given currently available data) to link fossil and Recent Species concepts. It is worth noting here that many better-studied Recent non-turritellid cerithioid species seem to show relatively high intraspecific variability in shell form (e.g., Houbrick, 1974, pp.55-62), and indeed such high variation may be characteristic of Most cerithioid taxa (Houbrick, pers. comm.). Eldredge and Gould (1972, p.92) have characterized the voluminous literature on the definition and rec- ognition of species among fossil organisms as “а the- oretical debate unsurpassed in the annals of paleon- tology for its ponderous emptiness. . .”, because in the context of phyletic gradualism it amounts to little more than a bookkeeping problem of arbitrarily parsing up Continuous lineages. Eldredge and Gould got around the issue by suggesting that lineages seldom change in the phyletic mode; species tend to be more or less discrete and the problem seldom arises. This is clearly the case if ancestor and descendant overlap in time and space, resulting in a bimodal distribution of co- existing morphologies. If, however, apparent ancestor and descendant are disjunct or contiguous in time (im- plying a more or less wholesale transition from one to the other), then cladogenesis cannot be demonstrated conclusively, punctuated equilibrium is not applicable, and the issue of what to call various forms is important and cannot be avoided, if only for the largely utilitarian reason of recording biostratigraphically useful infor- mation. When these more problematic conditions ap- pear to apply to the taxa discussed here, I have adopted the following working definition (modified from Beer- bower, 1968, p.80-81; Waller, 1969, p.8; Gould, 1969, Dp.459ff. Raup and Stanley, 1978, pp.108ff): fossil spe- cies are groups of morphologically distinct populations within which variation is of the magnitude displayed by closely related, or presumably analogous, living species and their local populations, and between which the dif- ferences are of the kind and degree expected to result from reproductive isolation of populations in such re- lated or analogous species. In a similar fashion, subspecies may usefully be rec- ognized among fossils to designate populations sepa- rated in time and/or space and differing morphologi- cally to a degree less than that distinguishing two spe- cies in the group (cf, Newell, 1947; Gould, 1969). SUPRASPECIFIC LEVEL Turritellids represent a particularly conspicuous ex- ample of the problems of defining genera in fossils (Allmon, 1992a). The issues involved have been well summarized by Marwick: There has rarely been more reluctance to grant generic rank to sub- divisions of a broad Lamarckian genus than there has been with Turritella. The main causes for the desire to retain Turritella as a world-wide, Cretaceous, possibly Jurassic to Recent genus are, no doubt, the small range in shape, the relatively simple aperture, and the general absence of well-developed axial sculpture. One rightly hesitates to grant generic significance to differences that seem trivial. When, however, such differences, on close study, emerge as reliable guides to what appear to be genetically related groups, their useful- ness must not be ignored. Whether the subdivisions are accepted as genera, subgenera or sections, or neglected altogether, will depend on individual tastes, backgrounds and traditions. Much more in- formation is needed about the animals themselves before a really satisfactory classification can be widely accepted. (1957b, p.7) Turritellids show relatively low morphological (i.e., conchological) disparity but high species diversity; what variation there is does not indicate relationship un- ambiguously, but rather seems to have arisen again and again throughout the history of the group (Mar- wick, 1957a; Kotaka, 1978; Allmon, 1987), as indi- cated by the general lack of concordance among the most conspicuous morphological characters. Species do not appear to form homogeneous subgroups sepa- rated by conspicuous morphological gaps. Following the suggestions of previous authors (Mer- riam, 1941; Marwick, 1957a,b), Allison (1967; Allison and Adegoke, 1969) has argued that supraspecific taxa within Turritellidae should be based on differences in at least two characters, for example growth line and apical spiral ontogeny, rather than on only one. Su- praspecific taxa defined by as few as two characters are still very similar morphologically, and correspond more closely to a “phylogenetic” or “‘cladistic” definition (in which genera can be recognized as monophyletic clades defined by as little as a single synapomorphy) than to ' a “phenetic” or “gap” definition (in which genera must differ by a large number of characters) (see Allmon, 19922). In proposing supraspecific groupings for Lower Tertiary Coastal Plain turritellid species, I have fol- lowed Allison's suggestion and adopted a phylogenetic genus definition (see Allmon, 1992a for further dis- cussion). 14 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PHYLOGENETIC ANALYSIS METHODS Introduction The central problem in a phylogenetic analysis of fossil turritellids is the small number of available shell characters. Because in a purely cladistic analysis, at least n — 1 shared derived characters (synapomorphies) are required for 5 taxa, paucity of characters means that there will usually be too few characters to resolve dichotomous branching relationships fully (Eldredge and Cracraft, 1980, p.29). Rigorous cladistic analysis can be done on living and fossil gastropod shells (e.g., Houbrick, 1985, 1987a; Erwin, 1988; Michaux, 1989; Allmon, 1990). Such analysis is possible, however, only if characters are relatively numerous (i.e., at least ful- filling the above requirement) and abundant infor- mation is available on character states in other mem- bers of the group so that polarity determinations can be made (or confirmed) by either outgroup comparison or stratigraphic criteria. Neither of these conditions applies to the turritellid species considered here. Only six characters or character complexes are currently available for analysis (see below), and the states of several of these characters (especially apical ontogeny) are unknown for many other members of the group. Consequently, a purely atemporal cladistic analysis is not, for the present at least, the most productive meth- od of phylogenetic analysis for these species. Analysis of the species considered here has therefore proceeded at five levels: 1) species recognition, 2) grouping Coastal Plain species into morphological spe- cies groups, 3) hypothesizing and testing relationships of these species groups to each other, 4) hypothesizing and testing branching order among species within each group, and 5) hypothesizing and testing relationships among Coastal Plain species groups and other Recent and fossil turritellid supraspecific groups. In grouping species together, and in hypothesizing relationships among species groups (steps 2 and 3 above), an explicitly cladistic approach has been taken. Species and species groups are linked together on the basis of presumed shared derived (synapomorphic) rather than shared primitive (symplesiomorphic) char- acters (i.e., overall similarity). Given that perfect con- cordance among characters is not obtained, the small number of characters currently available for analysis makes it impossible to seek other characters to cor- roborate one hypothesis of relationship over another (cf, Eldredge and Cracraft, 1980; Wiley, 1981; Cairns, 1984; Sharkey, 1989). An explicit character weighting scheme has therefore been employed. Character Weighting Character weighting has been criticized by some au- thors (e.g., Eldredge and Cracraft, 1980; but see Wiley, 1981, p.141), who emphasize that all synapomorphic characters are useful in phylogeny reconstruction. А8 emphasized by Wheeler (1986), however, analytical errors in identifying synapomorphies at their appro- priate hierarchical levels are not uncommon. Weight- ing expresses one’s degree of confidence that a partic- ular character is a synapomorphy at the appropriate level. Viewed in this context, the issue is how best to go about determining character weights. Mayr (1969, pp.220ff) has suggested several criteria by which “characters with high weight” could be iden- tified, including complexity, constancy, function and correlation. Wheeler (1986, p.107) suggests that char- acters “not known to occur multiple times in related taxa” are of highest value for cladistic reasoning. Neff (1986) and Wheeler (1986) both express reservations about basing such judgements on a priori assumptions about some inherent quality of characters themselves. Wheeler favors a posteriori weighting, “after characters have been critically reexamined and parsimony has failed to yield an unambiguous solution to conflicting characters" (1986, p.107). Neffsuggests that a “rational basis" for a priori weighting is provided by asking “how much do we think we know" about a particular char- acter. All of these criteria are reducible to expressions of confidence in synapomorphy identification. Shar- key's (1989) recommendation for weighting by com- patibility (= concordance) of characters appears valid, but is of limited utility given a paucity of characters. Following the suggestion of Wheeler (1986, p.106), character weighting in this paper consists of “simply the statement that among the evidence in conflict some characters seem less likely to be misinterpreted than others", rather than of any sort of numerical scoring. Specifically, growth line form has been given greater weight at the level of all Coastal Plain species, and apical ontogeny at the level of species groups within the Coastal Plain. These decisions are based partly on an a priori subjective judgement of complexity; growth line and apical ontogeny both appear to be more com- plex than whorl profile, size or apical angle. They are also, however, based on the relationship between these two characters and geographic distribution: all of the Coastal Plain species discussed here have a generally similar growth line trace (see discussion below), sug- gesting that this is a shared derived character at the level of all Coastal Plain Paleogene species. Within the species groups resulting from this anal- ysis, too few characters are currently available to pro- duce fully resolved cladograms. The characters allow- PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 15 ing discrimination of species consist largely of contin- uously varying, poorly nested differences in size, whorl profile and adult sculpture, which are difficult to po- larize. The nature and paucity of these characters re- sults in a large number ( > 100) of equal-length clado- grams being generated in strict parsimony analyses us- ing PAUP (Phylogenetic Analysis Using Parsimony), version 3.0 (Swofford, 1990). Consensus trees based On these cladograms (e.g., Rohlf, 1982) include poly- chotomies containing most of the species. Thus to re- construct branching order a modified stratophenetic approach has been used. The logic and methods followed in this analysis are Consistent with those used in a previous study (Allmon, 1990), and also with the arguments of Lazarus and Prothero (1984). Phylogenetic inference based solely or largely on the stratigraphic position of known fossils has a probability of being correct proportional to the quality of the stratigraphic record (Fortey and Jeffries, 1982; Allmon, 1989). When the record is relatively incomplete, an atemporal, strictly morphological anal- ysis has a higher probability of yielding correct results than an analysis relying on stratigraphic data alone. The success of such a morphological analysis, however, will be limited if the organisms lack abundant, discrete, hierarchically nested characters not greatly prone to homoplasy. The degree to which cladistic and strato- Phenetic methods are used should be dependent upon the nature of the stratigraphic and morphological ev- idence available. The stratigraphic record here is of moderate to high completeness, and the morphology of the species is poorly suited for cladistic methods; I have therefore used a compromise" approach com- bining the two methods (see further discussion in For- tey and Jeffries, 1982; Allmon, 1989). With this in mind, the order of first appearance of Species within species groups in the stratigraphic record has been taken as the primary indicator oftheir branch- ing order. This still leaves, however, a substantial num- ber of uncertain branch points. These uncertainties cannot be resolved fully at present, but qualitative "confidence limits" can be placed on many of these Doints by making use ofthe distribution of fossiliferous beds within the Coastal Plain section. Observed stratigraphic ranges almost always under- estimate true durations of taxa (Marshall, 1990). It is reasonable to assume, therefore, that observed first ap- Dearances postdate actual evolutionary originations. Strauss and Sadler (1989) and Marshall (1990) have Presented methods for calculating confidence intervals for first and last appearances, but these techniques are applicable only if fossiliferous horizons are randomly distributed. This is not the case in the Coastal Plain Paleogene (Toulmin, 1977; Allmon, 1988b, 1989), and other methods must be sought. Most of the fossil mollusks from the Gulf Coastal Plain come from a small number of highly fossiliferous, vertically and laterally restricted, glauconitic sands and clayey sands, distributed irregularly both stratigraph- ically and geographically, and separated from each oth- er by thick, laterally extensive units of sparsely fossil- iferous sands and clays. These high-density, high-di- versity fossil beds represent a combination of the ef- fects of higher density and diversity of organisms, lower sedimentation rates, and a higher proportional pres- ervation potential of the organisms present (cf, Kid- well, 1986; Cummins et al., 1986). To the degree that the last factor applies, such horizons represent more reliable records of what species were present during their deposition than do less fossiliferous units. A cor- related effect is that substantially more paleontological attention has been given to these highly fossiliferous units and so knowledge of their faunas is probably more complete. The distribution of these fossil beds in the strati- graphic column can provide crude confidence levels for observed first appearances. Assuming that if a spe- cies was present, it would be more likely to be pre- served and found in a very fossiliferous than a less fossiliferous interval, degree of confidence in an ob- served first occurrence can be expressed as the interval between observed first appearance and the youngest highly fossiliferous bed below that appearance. The closer an apparent first appearance is to a highly fos- siliferous bed the smaller the uncertainty in time of actual first appearance because the fossil-rich bed is assumed to provide a maximum age. Results of such an analysis for Coastal Plain turritellid species are giv- en in Table 2. The high diversity, morphological simplicity, and imperfect fossil record of turritellids in the Coastal Plain may seem to amount to such a limitation of data that no firm phylogenetic and evolutionary conclusions can ever be drawn. My own response to this suggestion in the present context is admirably expressed by Whee- ler (1986): I find reaching “no conclusion" as uninspiring as any other inves- tigator, but we work in a less than perfect world. Rather than simply finding characters and recognizing their level of significance in the , hierarchy, we are relentlessly faced with data sets so small that per- turbations caused by convergent evolution mask the historical pat- tern we seek to discover. Such empirical constraints in the real world sometimes create situations wherein the researcher has to give up in the face of adversity or make less than idealistic methodological concessions. My position is that so long as the researcher— and the user community —recognize the inherent weaknesses in strict appli- cation of parsimony to complex data sets or the use of ad hoc weight- 16 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 2.— Confidence estimates for date of first appearance of turritellid species from the Paleocene and Eocene of the Coastal Plain. next oldest maximum error as observed first high-density possible percent of Species location! appearance?’ horizon? error (my)* total duration? alabamiensis CG CL — — — aldrichi CG, EG CL. - - — alveata сб MB GS 0 0 apita CG GS SCB/DBT ~7 2300 arenicola CG, EG MB GS 0 0 biboraensis WG KI — — - carinata CG, EG? UL SCB/DBT 0 0 claytonensis CG CE -- - - clevelandia WG, CG MB GS 0 0 cortezi WG CM SCB 0 0 creola WG WE BA ~8 267 creola CG MB GS 0 0 dumblei WG SCB WE 0 0 dutexata WG WE BA ~8 267 eurynome CG NF MLM 13 22 femina WG WE BA ~8 267 gilberti CG BA TSBL ~1 31 hilli WG KI == — — humerosa AT AQ BS 3.0 94 infans WG CM SCB 0 0 kincaidensis j WG KI — — - levicunea CG CL — — - lisbonensis CG LL BA ~8 133 mortoni mediavia CG CE - - - mortoni mortoni AT AQ BS 3.0 94 multilira CG NF MLM 1:9 50 perdita CG MB GS 0 0 pleboides WG CM SCB/DBT 0 0 mortoni postmortoni CG NF MLM 153 22 mortoni “ргетогіопі” AT BS - -- — praecincta praecincta CG NF MLM 199 22) praecincta virginiensis AT AQ BS 3.0 94 “prehumerosa” AT BS — — - rina CG, LL BA ~8 114 rivurbana CG, WG MB® GS 0 0 subrina CG LL BA ~8 114 tennesseensis CG CL - - - 1 Abbreviations: WG, western Gulf (Texas, Louisiana); CG, central Gulf (Mississippi, Alabama); EG, eastern Gulf (Georgia, western Florida); AT, Atlantic coast. 2 Abbreviations of lithostratigraphic units same as in Table 8. з Definition and origin of “high-density” fossil beds discussed in Toulmin (1977) and Allmon (1988b, 1989); stratigraphic distribution of these units shown in Text-figure 15. For taxa showing their first appearance in the Lower Paleocene Kincaid and Clayton Formations, no underlying bed is designated, since the sedimentology and taphonomy of coastal plain shell beds has not yet been investigated. It is important to note that there is as yet no positive evidence to support either continuity of discontinuity of particular turritellid lineages across the Cretaceous-Tertiary boundary in this region. For the purposes of this analysis it is assumed that all known Lower Paleocene species arose in the Lower Paleocene. 4 Таха are assumed to have existed throughout entire duration of any lithostratigraphic unit in which they occur. Biostratigraphic dates of boundaries of units discussed in Toulmin (1977) and Allmon (1988b). Absolute time scale that of Berggren et al. (1985). 5 Total observed durations of species given in Table 8. 6 Allison and Adegoke (1969) have reported a subspecies of T. rivurbana from “Undifferentiated strata of latest middle Eocene age" in Chiapas, Mexico. On the U.S. Coastal Plain the earliest occurrence is in the Moodys Branch Formation. ing assumptions to resolve homoplasy, then perhaps these maneu- Geography as a Taxonomic Character vers are preferable to giving up. After all, the cladogram does rep- WO На 3 ; 5 resent a testable hypothesis and, if the number of available characters Use of geographic distribution B phylogenetic anal- increases in the future, poor assumptions about weights will be shown ysis is problematic because there is no necessary link for what they are. (1986, pp.108-109) between recency of common ancestry and geographic | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 17 Proximity. The issue often arises in biogeographic Studies, in which the goal may be to reconstruct the geographic history of a group. To use geography as a measure ofrelatedness and then explain the geographic Pattern using the resulting phylogeny is clearly tauto- logical. Yet ifthe object is not biogeographic, but phy- logenetic reconstruction, the reasoning need not be cir- cular. Geographic distribution may in certain cases be a useful taxonomic character, because related taxa of- ten (though by no means always) occur in the same general area, a line of reasoning dating to Wallace’s "law which has regulated the introduction of new spe- cies" (1855). As Mayr (1969, p.141) puts it, "there is a high probability that related species in an area are descendants of a common ancestor and that in the majority of cases no other species of this group occur disjunctly at a far distant place". (Nelson and Platnick (1981, p.384) reach exactly the opposite conclusion, Stating that “Vicarious distributions are less common among distantly related taxa, such as genera within a family... Distantly related taxa tend to be sympatric 2°.) Mayr suggests that Geographical characters are among the most useful tools for clarifying a confused taxonomic picture and for testing taxonomic hypoth- eses” (1969, p.140; see also Simpson, 1961, p.74). Es- sentially the same reasoning underlies Hennig's (1966, p.172) “biogeographic method”, which uses geograph- lc range changes to assist in deriving minimum ages Of sister-group relationships. In Turritellidae, Marwick (19572) makes explicit use of geographic distribution to corroborate relationships based on morphological characters and to identify ho- moplasy: An important factor responsible for much of the confusion [in tur- ritellid classification], has been the failure of many, but not Iredale [1924, 1925] or of Merriam [1941], to realize the strict geographic limits of most of the ‘subgeneric’ groups [in the family]. (1957a, p.144) Similarity in shape and in adult sculpture is not sufficient to tran- Scend geographic regions. Close agreements in outer lip characters and in primary spiral ontogeny are essential for generic grouping, and even then the possibility of convergence of distant stocks with Simple characters must be considered. (1957a, p.158) Similarly, Allison (1967, p.252) suggests that the "finite and recognizable stratigraphic and geographic distribution" of many turritellid genus-groups contrib- utes to their recognizability. (Discussing bryozoans, Boardman et al. (1970) have argued along the same lines for taxonomic evaluation (and generic recogni- tion) of phenetic clusters of specimens based on their Occurrence in time and space.) Marwick (1957a,b) bases his use of geographic range on the belief that turritellids have relatively restricted dispersal ability. While noting that some species prob- ably have wide distributions, he cites Merriam (1941) as the authority for the claim that “Owing to their generally short free-swimming stage and their restricted bathymetric distribution . . ., the turritellas, on the whole, must have a comparatively limited colonizing ability. Therefore, in the long course of geological time, populations must have been par- ticularly liable to isolation through water changes in the distribution of land, the depth of the sea, and also the temperature of the water.” (1957b, p.8) “The restricted dispersal ability of many of the Turritellidae and the long period of time represented since they first appeared in the stratigraphic record . . . should mean that many distinct genera now inhabit different parts of the world.” (1957b, p.9). Direct information on the larval biology of Recent turritellids is scanty (Allmon, 1988a; Bieler and Had- field, 1990; Allmon et al., 1992; Lieberman et al., 1993). Observations are available for only four living species of the traditionally recognized subfamily Turritellinae, of which one is a long-term brooder releasing crawl- away young, two have short (7-21 day) planktonic phases, and one has either a very short or no planktonic phase. It is not known whether the larvae of any of these species feed while in the plankton. No living species is known to be teleplanic (sensu Scheltema, 1971), i.e., to have a very long, truly planktotrophic larval phase. Since turritellids appear generally to con- form to Thorson's “apex theory” (Thorson, 1946, 1950; see also Shuto, 1974b; Jablonski and Lutz, 1983), their mode of larval development can be inferred from form and size of the protoconchs of fossil as well as living species (see Allmon, 19882), and so the potential exists to test the basis of Marwick's generalization through time. Available data on protoconch size and form of fossil and living turritellid species are summarized in Table 3. Based on these data, it would seem that turritellids have shown a variety of larval ecologies throughout their history. All Cretaceous species for which data are available (all of which occur in the Gulf Coastal Plain), as well as several species from the Neogene of the Atlantic Coastal Plain, had small, multispiral proto- conchs, indicative of an extended planktonic (even if not teleplanic) phase implying high dispersal potential. Patterns are less clear for species from the Coastal Plain Paleogene. All species for which data are available have protoconchs with relatively small P1 (protoconch I), indicative of a small, yolk-poor egg and so probably : planktotrophic development, but also with relatively few whorls, indicative of a relatively short stay in the plankton (Fretter and Graham, 1962, p.472; Sohl, 1977, pp.520-521; see Jablonski and Lutz, 1983 for further discussion). Observed geographic ranges of these spe- cies vary from very large to very small (Table 8, p. ). A variety of protoconch sizes and forms is shown by 18 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 3.—Protoconch size and inferred mode of larval development for fossil turritelline species from the U.S. Gulf and Atlantic Coastal Plains and the four living species for which these data are available. Modes of Larval Ecology: N = nonplanktotrophic, planktic or nonplanktic; SP = short term planktic, probably nonplanktotrophic; P = planktotrophic; BR = brooding. References: (1) Chapter II; (2) Lebour (1933); Fretter and Manly (1979); (3) Jablonski (1979); (4) Spiller (1977); (5) this study. р eo inferred/ no. of whorls (um) observed in protoconch larval refer- Species age (Е-Е Е2) BI РІ + P2 (N) ecology ence variegata Linnaeus, 1758 Rec 2.25 214 387 1 SP (1) gonostoma Valenciennes, 1832 Rec 25 370 465 5 SP (1) communis Risso, 1826 Rec 160 450 SP (2) gunni Reeve, 1849 Rec 3.0 382 1080 1 BR (1) quadrilira Johnson, 1898 Leret. 30 80 P (3) trilira Conrad, 1860 L Cret. 3:25 80 P (3) bilira Stephenson, 1941 L Cret. 3.0 80 P (3) howelli Harbison, 1945 L Cret. 3.5 90 P (3) tippana Conrad, 1858 2. Стег. 30 75 Р (3) vertebroides Morton, 1834 L Cret. 3.25 85 Р (3) aequistriata Conrad, 1863 E Mio. 3-4 P (4) | duplinensis Gardner & Aldrich, 1919 E PHO. 3—4 P (4) | terstriata Rogers & Rogers, 1837 M Mio.-L Plio. 3-4 Р (4) variabilis Conrad, 1830 E Mio.-L Plio. 3-5 200 Р (4) | alcida Dall, 1896 M Mio. DES N? (4) alticostata Conrad, 1834 · E-L Plio. LS 110 N? (4) burdeni Tuomey & Holmes, 1857 E Plio. 2-3 N? (4) | chipolana Dall, 1892 E Mio. 2-3 N? (4) etiwanensis Tuomey & Holmes, 1857 E-L Plio. 1-2 N? (4) \ fuerta Kellum, 1926 E Mio. 2 N? (4) | indenta Conrad, 1841 E Mio. 122 N? (4) | mixta Gardner, 1947 E РПО. 2-3 N? (4) pilsbryi Gardner, 1928 M Mio. 2 N? (4) | secta Conrad, 1855 M Mio. 1-2 N? (4) subannulata Heilprin, 1887 E-L Plio. 2-3 N? (4) subgrundifera Dall, 1892 E-L Plio. 2 N? (4) alabamiensis Whitfield, 1865 E-M Paleo. 2.0 94 226 2 SP (5) aldrichi Bowles, 1939 E-M Paleo. 1-1.5 90 199 5 SP (5) alveata Conrad in Wailes, 1854 L Eoc. 0275 194 247 1 SP (5) apita de Gregorio, 1890 L Eoc. 2.0 N? (5) arenicola Conrad, 1865b L Eoc. 2.0 М? (5) carinata 1. Lea, 1833 M-L Eoc. 0.75 112 SP (5) chirena Stenzel & Turner, 1940 M Eoc. 2-2.5 112 240 1 SP (5) clevelandia Harris, 1896 L Eoc. 1.5 261 1 SP (5) creola Palmer, 1947 L Еос. 1.0 182 250 5) SP (5) | danvillensis Stenzel & Turner, 1940 L Eoc. 2.0 N? (5) | dumblei Harris, 1896 M Eoc. 179 121 230 1 SP (5) | dutexata Harris, 1896 M Eoc. 1.5 154 286 1 SP (5) 1 femina Stenzel, 1931 M Eoc. 1.73 126 246 1 SP (5) [ gilberti Bowles, 1939 L Pal.-E Eoc. 1.75-2.0 106 238 2 SP (5) humerosa Conrad, 1835b E-L Paleo. 3-47 p? (4) | mortoni Conrad, 1830 E-L Paleo. 1575 97 280 5 SP (5) | perdita Conrad, 1865 L Eoc. 1.75 97 223 3 SP (5) | pleboides Vaughan, 1895 M Eoc. 1.5-1.75 109 248 2 SP (5) “prehumerosa” Govoni, 1983 E Paleo. 2.0 88 250 1 SP (5) | “premortoni” Govoni, 1983 E Paleo. 2-2.5 88 291 1 SP (5) | rina Palmer, 1937 M Eoc. 1.75-2.0 106 242 2 SP (5) tennesseensis Gabb, 1860 E Paleo. 225 85 267 | SP (5) and Spirocolpus), however, all have multispiral pro- | toconchs. If turritellid species have varied in their larval ecol- ogy and geographic range throughout their history then, fossil and living turritellids from the Australia-New Zealand region (Text-figure 2) (Marwick, 1957b, p.8); the only fossil or Recent turritellid taxa common to Australia and New Zealand (Maoricolpus, Colposigma, PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 19 RAEBRERE Text-figure 2. — Protoconchs of fossil and Recent turritellid species of the tribe Zeacolpini from New Zealand, showing range of form (redrawn from Marwick, 1971). as pointed out by Sohl (1977, p.521), “It is not suffi- cient ... to depend upon knowledge of the length of larval life of living species to estimate the dispersal capabilities of related fossil forms". Because proto- conch size and form seem to be adequate predictors of geographic range, it is in theory possible to know in advance to what degree the species of interest were geographically restricted. Many Cretaceous and some Neogene species were widely dispersed, and Marwick's expectation of isolation and localization of related taxa would probably not apply. Most other species for which data are available (including the species under consid- eration in this paper), however, are consistent with the idea that turritellids did not have very high dispersal Capabilities. It is therefore reasonable to assume that descendant taxa tended to inhabit regions close to those of their immediate ancestors, and that to this extent, geographic range can be used as an indicator of pro- Pinquity of descent. Consideration of regional biogeography is also useful as an indicator of which taxa (i.e., which areas) should be given more attention as potential close relatives of the taxa of interest. In the present case we may con- Sider, for example, the biogeographic relationships of three areas with important fossil turritellid faunas that have received some degree of study: North America, western Europe, and Australia-New Zealand. Although earlier workers (e.g., Finlay, 1926) have suggested that the marine invertebrate faunas of New Zealand reflect- ed almost complete isolation since the Cretaceous, Fleming (1962, 1967, 1979) has showed that consid- erable faunal interchange occurred periodically throughout the Cenozoic. If these episodes could have involved turritellids then one might question the monophyly of most New Zealand turritellid supra- Specific taxa. For several reasons, however, this does Not appear to be the case. First, the majority of benthic Marine invertebrate immigration into New Zealand during the Cenozoic was from the tropical Indo-Pacific realm. Although turritellids do occur in the latter re- gion today and as fossils (e.g., Shuto, 1974a), asa group they are not thermophilic (Allmon, 19882), and prob- ably never comprised an important component of the warm-water fauna of the region (although further data are required to test this hypothesis). Secondly, docu- mented cases of immigration into the region from out- Side the southwestern Pacific (e.g., pectinid bivalves from the Mediterranean region during the Late Ter- tiary, discussed by Fleming (1957) involve taxa with easily dispersed planktonic larvae. On this basis it would seem that Europe or Africa or other areas of the New World would be more likely areas to investigate for relatives of North American and Caribbean turritellids than New Zealand-Austra- lia. These possibilities are considered in a later section. Morphometric Analysis Species- and subspecies-level nomenclature have been highly unstable in a number of cases among Coastal Plain Paleocene and Eocene turritellids. Such instances suggest that patterns of morphological vari- ation are complex, and that careful morphometric analysis might be useful for sorting them out. Large samples of specimens were therefore measured for four of these problematic taxa, using a video image analysis system. Measurements made on each taxon are illus- trated in Text-figure 3. In most gastropods it 1s fairly easy to identify ho- mologous points for aligning whorls prior to morpho- metric analysis. The end of the protoconch or a varix produced in the adult shell can be used as a landmark from which whorls can be counted. In high-spired shells with indeterminate growth, such as turritellids, how- ever, whorls differ only in size and shape, and lack such discrete ontogenetic landmarks. Incomplete spec- imens, which are particularly common as fossils, are therefore difficult or impossible to align, homologize, and analyze morphometrically. This situation is un- doubtedly chiefly responsible for the complete lack of any previous multivariate morphometric study of tur- ritellids, fossil or Recent, for systematic purposes, de- spite the relatively high level of interest in the bio- stratigraphy and evolution of the group. Fossil turri- tellid shells are often abundant, but truly complete adult (i.e., maximum size) specimens, with both pro- toconch and unbroken apertural lip, are extremely rare. When juvenile whorls are missing it is therefore dif- ficult or impossible to identify similar whorl number and so to specify homologous points on shells among specimens ofa given species. Construction of data ma- trices for morphometric analysis of a large number of turritellid specimens has thus heretofore been impos- sible. For these reasons, in this study I have utilized a new | computer program, “HISPIRE” (Morris and Allmon, 1994), to align specimens of single morphospecies pri- or to morphometric analysis. HISPIRE homologizes incomplete specimens of high-spired gastropod shells by matching whorls of similar size. It thus aligns spec- imens in a new data matrix, filling the remainder (miss- ing whorls) with zeroes (— missing values for each mea- surement). The assumptions underlying HISPIRE are 8 э 9 10 5 n CD G H Text-figure 3. — Measurements made on five turritellid species. Each diagram is a single whorl seen from the side. Numbered dots represent points captured as X-Y coordinates by a digitizer, from which linear measurements were calculated. Measurements are de- fined here as pairs of these points. Subscript p refers to point on succeeding whorl. A. Turritella terebra (Linnaeus): whorl height (WH) = 1-1,; suture width (SW) = 2-5; whorl width (WW) = 3-6; B. Turritella terebra (Linnaeus): inflation (IN), as indicated; C. Pal- merella mortoni (Conrad): whorl height (WH) = 1-1 p; Carina height 1 (CH1) = 5-1,; carina height 2 (CH2) = WH-CH1; suture width (SW) = 2-6; middle width (MW) = 3-7; carina width (CW) = 1-8; D. Palmerella mortoni (Conrad); carina depth (CD), carina angle (CAN), as indicated; E. Haustator carinata (I.Lea): whorl height (WH) = 1-1,; carina height 1 (CH1) = 7-1; carina height 2 (CH2) = WH-CH1; suture width (SW)= 2-9; width 1 (W1) = 3-10; width PALAEONTOGRAPHICA AMERICANA, NUMBER 59 discussed in greater detail in Morris and Allmon (1994). The details of the analyses performed here are given in Appendix 1. Multivariate analysis (factor analysis) was carried out on one Recent and four fossil taxa using BMDP (1992, Program 4M). CHARACTERS Some indication of the utility and reliability of shell characters in phylogenetic analysis of turritellids may be obtained from the many studies that have been carried out on other cerithioid families, principally by R.S.Houbrick. In general, Houbrick (e.g., 1988) con- tends that, at familial and higher levels, shell characters are usually unreliable indicators of phylogenetic rela- tionships, in any case much poorer than anatomical (“soft part") characters. In the phylogenies he proposes for various cerithioid taxa at lower levels, however, shell characters often appear reasonably consistent with non-shell characters, differing chiefly in being fewer in number and so more limited in their resolving power. For example, although he states (1984b, p.18) that “Polarities of shell characters . . . are either difficult to determine or suspect because of possible parallelisms or reversals," Houbrick bases recognition of three sub- genera of Cerithidea Swainson, 1840 (Potamididae) principally on shell characters (e.g., sculpture, form of aperture and apertural lip). Similarly, the species “Сеу- ithiopsis crystallina Dall” is placed in the genus Var- icopeza Grundel, 1982 (Cerithiidae) largely on the ba- sis of shell characters (Houbrick, 1987b). Species of the genus Clypeomorus Jousseaume, 1888 (Cerithi- idae), are stated to “differ from other cerithiids by their low-spired, frequently beaded shells" (Houbrick, 1985, p.1); shell characters corroborate a hypothesis of monophyly for the genus based on arrangement of the pallial oviducts and habitat requirements (pp. 7-8), and are consistent with its division into three species- groups (pp. 11-14). Although it is possible that in the future other useful shell characters (e.g., ultrastructure, developmental data on shell geometry) will become available, at present only six conchological characters or character com- plexes are of any real utility in phylogenetic analyses of turritellids: 1) protoconch size and form, 2) growth — 2 (W2) = 4-11; width 3 (W3) = 5-12; carina width (CW); 6-13; Е. Haustator carinata (1.Lea): carina depth (CD), as indicated С. “Tur- ritella” praecincia Conrad: whorl height (WH) = 1—1,; suture width (SW) = 2-7; carina width (CW) 3-8; width 1 (W1) = 4-9; width 2 (W2) = 5-10; width 3 (W3) = 6-11; H. “Turritella” praecincta Conrad: carina depth (CD), as indicated; carina ratio (CR) = CW/ W1; I. Haustator perdita (Conrad): whorl height (WH) = 1—1,; suture width (SW) = 2-7; width 1 (W1) = 3-8; width 2 (W2) = 4-9; width 3 (W3) = 5-10; carina height 1 (СНІ) = 6-1. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 21 line form, 3) ontogeny of spiral sculpture, 4) character of adult sculpture, 5) adult whorl profile, and 6) size. The nature of variability, apparent taxonomic value, and means of description and polarity determination for each of these characters are considered below. Protoconch Merriam (1941, p.9) states that the turritellid pro- toconch “is essentially the same in all species exam- ined, and is similar to that of other genera—for ex- ample some of the Cerithiidae," and concludes that “The structure is too simple and generalized to be of value for purposes of taxonomy." Allison (1967) sim- ilarly has placed little taxonomic value on protoconch form, stating that “much additional study will be re- quired before the [taxonomic] value of this structure is understood.” These statements now appear to be Oversimplifications. Although turritellid protoconchs Share many features, they also show substantial vari- ability which, in at least some cases, seems to have taxonomic significance. All known turritellid protoconchs consist of between One and four or rarely five smooth, unsculptured whorls (Text-figure 2, Table 3). The protoconch I—proto- conch II (P1-P2) transition (if present) is not conspic- uous and the boundary with the teleoconch is marked only by the development of spiral sculpture, which may be quite faint initially. Associated with differences in whorl number and diameter are often differences in Overall form, which may vary from having a first whorl well immersed within the others to an erect, high-spired cone (see Text-figure 2). Variation in protoconch form within supraspecific groups of turritellids occurs but seems to be limited in extent. Garrard (1972, p.268) has noted that the pro- toconch “тау vary to a small extent within a genus." Marwick (1971, p.8) has observed that in New Zea- land, “Oligocene and Miocene generic groups and stocks Show considerably more protoconch differences within the group than do Pliocene to Recent ones." Although Garrard states that protoconch form “remains fairly constant within species,” variation within species has been reported. In the Recent Australian species Ga- zameda declivis (Adams and Reeve, 1848), the first Protoconch whorl varies from depressed to erect (Gar- tard, 1972, p.311). One species from the Oligocene (Tropicolpus (Amplicolpus) gittosinus (Powell and Bar- trum, 1929)) and two species from the Miocene (Zea- colpus fyfei Marwick, 1931 and Z. woodhouseae Mar- wick, 1971) of New Zealand each show two protoconch forms, one large, bulbous and paucispiral, the other erect and multispiral (Marwick, 1971). All protoconchs of Paleogene Coastal Plain species Observed in the present study are similar in form (small P1, paucispiral). (Spiller [1977] has reported that Tur- ritella humerosa Conrad has a multispiral protoconch, but I have not been able to confirm this.) Available sample sizes of well-preserved protoconchs of Coastal Plain species vary (Table 3). Protoconchs are unknown for a number of species, and I have been able to find only single specimens for a number of others. These sampling problems aside, no appreciable intraspecific variation in protoconch form or size has been ob- served. Planktotrophy is probably primitive in mesogastro- pods (Shuto, 1974b; Jablonski and Lutz, 1983, p.41), although some planktotrophic species may be second- arily derived from nonplanktotrophic forms that re- tained larval feeding structures (Scheltema, 1978). The presence of only planktotrophic turritellids in the Late Cretaceous of the Coastal Plain (Jablonski, 1979; Table 3) and the lack of long-term planktotrophic species in the family today are generally consistent with the planktotrophic habit being primitive in Turritellidae, although the variation in the Lower Tertiary and throughout the Cenozoic suggest that the pattern may not be a simple one (see Lieberman et al., 1993, for further discussion). Ontogeny of Spiral Sculpture Even casual examination ofa turritellid shell reveals that the form of external sculpture is not constant throughout ontogeny. As noted by Palmer (1937, p.279), this variation commonly makes it difficult to identify whorls from different ontogenetic stages as be- longing to the same species. Ontogenetic changes are particularly conspicuous on the earliest teleoconch whorls, when spiral sculpture is just beginning (Text- figures 4,5). The adequate description and taxonomic usage of this variation, however, have proven difficult. Finlay (1927, 1930) was the first to call attention to the importance of the ontogeny of early spiral sculpture (= apical ontogeny), and he has devised a notation system to describe it. Before discussing this system and its modifications, however, it is necessary to consider the alternative verbal system used by several American workers, two of whom studied species discussed in this paper. Palmer (1937) uses the terms “unicarinate,” ““bicar- inate” and “tricarinate” to describe the number of spi- ral ribs observed both on the earliest teleoconch whorls : (= her “nepionic whorls”) and on later juvenile whorls. Thus a species could be ““bicarinate-tricarinate,” meaning that whorls with two spirals were followed by whorls with three. Merriam (1941) and Bowles (1939) uses the terms “unicostate” for species with a single spiral rib on the earliest teleoconch whorl, “cingulate” for those with two ribs appearing roughly simulta- 22 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 W 1 4 X |diameter P1 SS 1 | x protoconch teleoconch А diameter Р1+Р2 кү initiation of spiral sculpture Text-figure 4.—Apex of a turritellid shell, showing methods used for counting and measuring protoconch whorls. A. Because turritellids generally lack any conspicuous boundary marking the transition between Protoconch I (P1) and Protoconch II (P2), the diameter of Pl is somewhat arbitrarily measured along a line perpendicular to the suture at its origin (point X). Whorls in the protoconch (P1 + P2) are counted following the technique described by Marwick (1957b; taken from B. Smith, 1945): beginning at the origin of the suture line (X), one-half whorl is counted when the point Y is reached, and one full whorl at the point labeled W. B. The end of the protoconch (= beginning of teleoconch) is defined by the initiation of spiral sculpture (usually visible only with SEM). The protoconch illustrated consists of approximately 1.25 whorls. neously, ““bicostate” for those with two ribs more closely spaced, “tricostate” for those with three, and “multi- costate" for those with “three to six subequal revolving lirae originating almost simultaneously on the early apical whorls" (Bowles, 1939, p.270). Merriam intro- duces the terms “primary,” “secondary” and “terti- ary" to refer to the relative strengths of the spirals. Palmer (in Harris and Palmer, 1947) has pointed out that her terms had priority, but as discussed in detail by Allison (1965, 1967), “carinate” and “costate” are used in different ways by these authors. Palmer con- siders only the strongest rib present in making her designations whereas Bowles and Merriam considers all ribs present. Thus a shell with three ribs appearing on the first teleoconch whorl simultaneously, but with one stronger than the other two, would be “unicari- nate" in Palmer's system and “tricostate” in Merriam's and Bowles’ (Allison, 1965, pp.670-671). For this rea- son these verbal descriptions are of little or no use in describing turritellid sculptural ontogeny (Allison, 1965), and they should be abandoned. Finlay's (1930) system of notation is based on the observation that the vertical position of the first-ap- pearing, or "original keels," on the whorls appears to be constant with respect to the sutures. That is, the rib appearing in the center of the whorl of one individual ofa species is presumed to be homologous to the center rib in another individual of the same species and, by implication, to individuals in other species. Finlay la- bels the four original spirals A, B, C, and D from pos- terior (adapical or abapertural end, “‘top’’) to anterior (abapical or adapertural end, “‘bottom’’) of the whorl. Ida (1952) adopts Merriam's terms primary, sec- ondary and tertiary, but instead of using them to refer to relative strength, he used them to refer solely to order of appearance. Spiral ribs appearing first he called primary, those appearing after the primaries he called secondary, and those appearing after secondaries he called tertiary. Ida then develops a complex notation system capable of describing accurately every spiral thread on the turritellid shell. As Marwick (1957a, p.148) notes, this system is well-suited for detailed specific description, but inconvenient for brief generic differentiation. Largely independently of Ida's work, Marwick (1957b) has proposed a modification of Finlay's no- tation system. Instead of four primaries between upper and lower sutures, Marwick recognizes three: B at about midwhorl, A the first to appear above, C the first to appear below. The D spiral marked the base of the whorl, just above the suture. Marwick further suggests that “secondaries” (i.e., in order of appearance, not PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 23 Text-figure 5.—Scanning electron micrographs of well preserved apices of two turritellid species, showing the two apical sculptural types Most common among species from the Paleocene and Eocene of the U.S. Gulf and Atlantic Coastal Plains. a. Turritella mortoni Conrad from the Paleocene Aquia Formation of Virginia (160 x). b. Sketch made from photomicrograph (a) indicating primary spiral ribs. Notation follows that described in Text-figure 6 and text. The B and C ribs appear approximately simultaneously, followed by the A rib, and so the apical sculpture formula for this species is C, B, A;. с. Turritella carinata I.Lea from the Eocene Gosport Sand of Alabama (160x). d. Sketch of (c) indicating primary spiral ribs. The C rib appears first, followed by the B rib, followed by the A rib, giving an apical sculpture formula of C, В, As. strength) be labeled with lower-case letters: r,s,t, and u, posterior to anterior. “Tertiaries” are “denoted by numbers according to which secondary they flank, as rl, r2, sl, s2, and so on; but generally they do not need individual designation” (Marwick, 1957b, p.13). As indicated in Figure 4, Finlay’s A is equivalent to Mar- wick’s r, B to A, C to B, and D to C; Marwick’s D has no equivalent in Finlay’s system. Kotaka (1959) modifies Marwick’s notation by rep- resenting primaries with lower case letters (a, b, c, d) if they were not strongly developed and secondaries by upper case letters (R, S, T, U) if they were more 24 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 =A u о с а <= 7C в— ШШ d Ил.» C uml d suture Text-figure 6. — Notation systems used for describing spiral ribbing of turritellids. On the left is the system used here, essentially equiv- alent to that of Marwick, Kotaka and Allison, as described in text. A, B, and C are primary spirals, r, s, t and u secondary spirals. the d spiral is usually simply the angulation associated with the basal (adapertural) suture, but may be more pronounced. On the right is the system proposed by Finlay (1930); his A is Marwick's r, his B Marwick's A, his C Marwick's B, and his D Marwick's C. See text for further discussion. strongly developed than the primaries. More impor- tantly, Kotaka adds numerical subscripts to all rib no- tations to indicate the order of appearance of each individual rib. (Marwick [1957a] previously indicated order of appearance by the order in which he listed the primaries; e.g., B C A or C BA.) These notations can then be combined into a single formula (referred to hereafter as an “apical sculpture formula"), written from the anterior to the posterior of the whorl (e.g., C; t4 B, 5; a4), to represent the entire spiral ontogeny. (A minor problem with Kotaka's discussion, as Allison (1967) points out, is that he appears to have reversed the terms “adapical’” and “abapical” in describing the position of A, B, C and D, etc. As Allison suggests, this simple error can be disregarded and the position of the lettered ribs considered following Marwick (1957b).) Allison (1965) modifies Kotaka’s system by rec- ommending 1) that D be included in the sculptural formula when present, but that it be given no numerical subscript due to the difficulty in determining its point of origin, and 2) that D be capitalized when it forms a prominent carina on the side of the body whorl but designated in lower case when it consists only of the angulation at the whorl base. Allison also departs from previous practice by writing apical sculpture formulas in alphabetical order from the top down. I have not adopted this last change in convention here and list the ribs from the bottom up (Text-figure 6). Finally, Adegoke (1967) and Allison and Adegoke (1969) uses series of Kotaka-type formulas to represent not only order of introduction but sequence of changes in carina strength. For example, in the “Turritella rina group" the B spiral characteristically becomes obso- lete, and this can be represented as follows (Allison, 1967; see Allison and Adegoke, 1969): a, В C; > A, В, C, d— A, b; С d— A, С d Although these descriptive systems allow the relative order of appearance of all spiral sculptural elements on an individual shell (or of a single species) to be described, they cannot present symbolically the timing of all changes in sculpture during ontogeny. The closest approach to such is Ida’s (1952) system, which can describe every individual whorl, but which is also rath- er unwieldy (see, for example, Titova, 1994a). For the present work, I have tried to extract from this notational and nomenclatural history a series of designations that communicate the greatest amount of information in the least complex way. For individual species, I use apical sculpture formulas listing as pri- maries the three first-appearing spirals from the bot- tom, or adapertural end, of the whorl below the C spiral (e.g., C,B,A). The D spiral is at the base of the whorl. Secondaries appear after and are intercalated among the primaries. Well-developed spirals are given in up- per case, weakly developed spirals in lower case (see Text-figure 6). Rather than listing sculptural formulas for individ- ual whorls, I summarize the sculptural ontogenies of turritellid species graphically. I have modified the type of diagram used first in a very simple form by Kotaka > Text-figure 7.— Schematic diagrams illustrating ontogenetic development of spiral sculpture in turritellid species from the Paleocene and Eocene of the Coastal Plains. These are herein called “Marwick Diagrams” in recognition of the use of a similar technique for illustrating turritellid ontogeny by Marwick (1971). Numbers across the top represent whorl number, the protoconch and youngest whorls being on the right. Succeeding whorls are standardized to the same height. Horizontal lines represent the spiral ribs as they appear and change in strength throughout ontogeny. Curved vertical lines at intervals represent whorl profiles. Diagrams do not represent single individuals, but were constructed by counting whorl number from protoconch-teleoconch boundaries, recognized in scanning electron micrographs, then constructing “composite individuals” by assuming that whorls of similar size were the same number, and examining incomplete specimens of increasing size. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 25 25 24 23 22 2120 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 EA / | Lao chirena == 1 NENNEN Ee / zen arenicola ЕЕЕ / pe 5 | > ү b | 1 — T y: EA 74 = alveata E Е - = = ——— == с | Em 7 EEA | bw = 7 Bea pleboides — | а | = Я =ч { E 2 74 =. | сгео/а (= 7 7 | есі 2 i mg — \ \ ee e | =e ЈЕ 7 TA 7 / [ч] ] 7 j | dumblei IL " 7 " zm I | E : ER | Ve 7 7 ESTEE clevelandia E + + ТЕСТЕРІ | — 2 j— L. di | Хав \ ) \ | | dutexata 4 у ЈН | | ғаәағаш | femina == ] ] | | М < — | | - е f 7 z | ea | mortoni тогіопі е е Е ; е Ј am nn = 26 25 24 23 22 2120 19 18 17 16 15 14 13 1211100 8 7 6 5 4 3 2 A noc c Meere eei pec qus e ciel M pu ce са НЫШ PALAEONTOGRAPHICA AMERICANA, NUMBER 59 1 Ё Torcula exoleta a ғ? У sauren ETE) —— К perdita jacksonensis 1——— LL Z P El Y C infans rina rina carinata =: f Ena 8 2 Га j— 7 gilberti ЈЕ ] E P (СЕ JE 22 ЕГУ 7 T | 7% 7 OVE alabamiensis F A == 4 = Í LL — 3 ( а EEE tennesseensis E Ji Z Dal) | dei =: \ i ger тш Тк nd r Text-figure 7. — Continued. (1959, p.48) and developed further by Marwick (1971, p.63) for this purpose (herein referred to as a “Marwick diagram") (Text-figure 7). At supraspecific levels, how- ever, such graphical methods are not practical (because interspecific differences within species groups are too great), and the recommendations of Allison and Ad- egoke (Allison, 1967; Adegoke, 1967; Allison and Ad- egoke, 1969) are more appropriate: supraspecific apical sculpture formulae need only summarize the order of appearance and relative strength of the primary (i.e., first-appearing) spirals. For example, the generic for- mula for Haustator Montfort, 1810 is C, B; a; (Mar- wick, 1957a, p.154). The phylogenetic importance granted here to early apical spiral sculpture is based chiefly on its relative complexity, which I believe increases chances of cor- rect identification of homology, but also on corrobo- ration by geographic and stratigraphic criteria. The work of Palmer, Merriam, Bowles, Marwick, Kotaka and Allison has demonstrated that coherent lineages can | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 27 25 24 23 22 21 20 19 18 17 16 15 14 13 12 1110 9 8 7 6 5 4 3 2 1 | =} " . eee praecincta praecincta ) ) ] ] humerosa === + m 1 и t 7. TA ] di E } ) #59) 7 = aldrichi E А! ) СЕЗДІ ) 1 У U Text-figure 7.— Continued. be traced mainly on the basis of apical ontogeny within a given geographic and stratigraphic context. The re- Sults presented below, furthermore, suggest substantial concordance, at least within the Coastal Plain Paleo- cene and Eocene, among apical ontogeny, growth line form and whorl profile. (As already mentioned, Mac- Neil (in MacNeil and Dockery, 1984) maintains that Convergence is equally likely in early spiral ontogeny as in any other character, and implies that sculpture and whorl profile of later whorls may be of more im- Portance. Yet he presents no specific evidence for this conclusion.) It is important to note that all previous systematic Study of turritellid apices has been done solely by means of light microscopy. Although careful examination of а large number of well preserved apices of some species by this method can reveal their earliest apical ontog- eny, extensive examination of Coastal Plain species With scanning electron microscopy (SEM) in the pres- ent study has shown that this is not consistently the Case. Spiral ribs may be quite faint at their inception and difficult to discern, even with SEM. As a result, Species that appear to have an apical spiral formula of, for example, C, B; A; under the light microscope may be seen to have C, B, A, under SEM (e.g.,“ Turritella Mortoni Conrad”; see Text-figure 5). As noted by Merriam (1941, p.35), published figures and descriptions, particularly prior to the last 30 years, are usually inadequate for determination of apical on- togeny of a species with this precision. Furthermore, apical ontogenies reported by more recent workers (e.8., Marwick, 1957a,b, 1971; Allison, 1965; Allison and Adegoke, 1969) based on examination with the light microscope also may prove to be incorrect. This is one of the principal factors limiting both the systematic results of the present study and the development of what Merriam has called a “universal classification of Turritellidae". Well preserved apices are rare, especial- ly in museum material, and they usually must be ex- amined with SEM to determine their apical ontogeny accurately. Because spiral sculptural patterns in turritellids ap- pear to be prone to heterochronic change in a variety of modes (Allmon, 1994), polarity of apical ontogenies is best determined by outgroup comparison, rather than ontogenetic criteria. As discussed below, choice of an outgroup for lower Tertiary Coastal Plain species is difficult. Reference to all recognized turritellid supra- specific taxa, however, may make choosing a specific outgroup unnecessary, at least for this character. Ex- amination of apical ontogenies of other turritellids in Table 1 shows that only two groups outside the U.S. Coastal Plain (Tropicolpus Marwick, 1931 from the Oligocene of New Zealand and Hataiella Kotaka, 1959 from the Miocene of Japan) have an apical sculpture formula of C, B, A,. Thus, unless one of these two is the sister taxon to Coastal Plain species, which seems unlikely on geographical grounds, this apical formula cannot be primitive. Only two other apical formulae are known to be represented on the Coastal Plain: C; B, A; by Turritella turneri Plummer, 1933 and T. dobyensis Dockery, 1980, · and C, B; A, by all other species. Assuming the latter to be primitive in this case is not based so much on the “commonality principle," which is of dubious va- lidity (Watrous and Wheeler, 1981), but on stratigraph- ic criteria; C; B, A, species are known only from the Middle Eocene on the Coastal Plain, whereas C, B, A; species are known from throughout the Paleocene and Eocene. Pleural angle apical angle lateral growth line trace spiral (lateral) sinus antispiral sinus suture basal sinus outer lip Text-figure 8.—Morphological features of adult turritellid shells used in classification and species descriptions. Circle to the right represents the base of the last whorl, looking toward the apex of the shell parallel to the axis of coiling. PALAEONTOGRAPHICA AMER’ CANA, NUMBER 59 More direct application of stratophenetic criteria, by recourse to the apical ontogenies of Cretaceous species, is not possible at present, as SEM studies of most Gulf Coastal Plain species are not yet available. Sohl (1960, p.71) has divided Upper Cretaceous species he de- scribes from the Gulf Coast into “bicostate,” “trico- state," “multilirate”” and “placement uncertain." As discussed above, however, these verbal descriptions are of little use in comparisons with the data required here. It still remains to connect Cretaceous species to Paleogene species on the Coastal Plain. Growth Lines Although the apertures of most turritellid shells, even while the animal is alive, are broken, the form of the original aperture is reflected by the incremental col- labral growth lines that are often visible on the external surface of the shell. The growth line has two aspects, which can be termed lateral and basal (Text-figure 8). The lateral is visible on the side of any whorl, because the growth lines run between the bounding sutures. The basal is only visible on the bottom of the last whorl of the shell, where the growth lines run from the whorl periphery to the columella at the center. Guillaume’s (1924) landmark paper on turritellid classification, in which he emphasizes the importance of examining growth lines, discusses only the lateral aspect (Text- figure 9), a point for which he is criticized by Dollfus (1926). The turritellid growth line is a sinuous line bent into a third dimension. As such it has several aspects and GROUPS итда \subenauage yeretrabs | јетра шт Bash en PLIOCENE | MIOCENE $ OLIGOCENE | EOCENE CRETACEOUS == | | | En | Text-figure 9.—Guillaume’s classification of turritellid growth line types, and their stratigraphic distribution in the Cretaceous and Cenozoic of Europe and North America. Only the lateral aspect of the growth line was considered by Guillaume, shown here schematically with inflection points beneath its stratigraphic distribution. (From Merriam, 1941, modified from Guillaume, 1924.) | | | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 29 BASAL LATERAL SINUS SINUS Inflection Points Apex Upper Half Middle Lower Half A B с ASA ЕРУ €B- C» 1 1 on bottom len; 2 ee double a MN) DO E MA er ee 4 277% Text-figure 10.—Classification of growth line traces used in this paper (see Tables 1,9). Can be described in several ways. The most important Of these are as follows (see Text-figures 9-11): 1) Lateral and basal sinuses (cf, Marwick, 19572). These are curves in the growth line visible on the lateral and basal aspects of the shell, respectively. The lateral 1s always concave in the “spiral” direction (Merriam, 1941); that is, toward the aperture, if only so slighly that it appears virtually straight. The basal sinus may be absent, and the growth line straight across the whorl base, but if present it is concave in the spiral or, more rarely, the *antispiral" direction (away from the ap- ег те). 2) A third sinus, concave in the antispiral di- rection, may be present on the angulation at the an- Text-figure 11.—Variation in basal and lateral aspects of growth line trace within the family Turritellidae. (Modified from Marwick, 19572.) All except d,u, and aa based on type species. a. Turritella, b. Zaria, с. Archimediella, d. Torculoidella bicarinata, e. Kurosioia, f. Leptocolpus, g. Ctenocolpus, h. Gazameda, i. Torculoidella, j. Stiracolpus, k. Peyrotia, 1. Maoricolpus, m. Zeacolpus, n. Haustator, 9. Tropicolpus, p. Torcula, q. Bactrospira, x. Platycolpus, s. Colpos- Dira, t. Spirocolpus, u. Neohaustator, v. Turritella hybrida [referred to Torquesia by Marwick], w. Colposigma, x. Colpospirella, y. Par- еота, z. Tachyrhynchus, aa. Protoma, bb. Protoma (Protomella) knysnaensis, c. Mesalia, dd. Neodiastoma, ee. Sigmesalia, ff. Glyp- tozaria. d 5 212900002 FERRO os С DA с dd 30 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 terior end of the whorl, in which case the basal sinus is concave in the spiral direction and the lateral sinus is relatively deep; this may be called the antispiral sinus (cf, Merriam, 1941). 3) The growth line angle is the angle “between the axial plane of the shell and a line connecting the anterior and posterior terminations of the growth line as observed on the exposed surface" (Merriam, 1941, p.59). When this angle is greater than zero, it can be called “prosocline”, when less than zero “opisthocline”, and when equal to zero “orthocline” (cf, Cox, 1960). 4) Inflection points of the growth line are emphasized as important taxonomic characters by Guillaume (1924). As pointed out by Marwick (19572), however, Guillaume's ‘points of inflection’ suffer from the family weakness, variability, and the distance of the abapical one from the suture is, in many shells, due as much to the height of the suture on the whorl as to the shape of the outer lip. .. . Nevertheless, since the strength and distance apart of the inflection points influence the shape of the sinus, they can be highly significant.“ (1957a, p.147) 5) The apex of the lateral sinus falls at varying heights above the anterior suture: above, below, or approxi- mately at the midpoint of the whorl (see Text-figure 10). 6) Growth lines and spiral ribs. In his critique of Guillaume, Dollfus (1926) states that Guillaume ig- nores the effects of spiral ribs on the form of the lateral aspect of the growth lines. Guillaume (1926) responds that these effects were negligible, an opinion with which Allison (1967) and I agree. 7) Ontogeny and gerontism. Ontogenetic changes in growth line form can be sig- nificant, especially in the very largest whorls in which apparently gerontic effects are manifest (cf, Marwick, 1957a; Allison, 1967; Garrard, 1972, p.268). Geron- tism is conspicuous, and important from a taxonomic point of view, only in the larger species; ontogenetic changes, however, appear to occur throughout the life- times of even smaller species (Allmon, 1994). Of the taxonomic value of the growth line trace, Merriam (1941) writes: Many species may be identified with a fair degree of certainty by means of this character alone. Nevertheless, it cannot well be used as the sole basis for the foundation of subgenera or major divisions oflesser value. The curvature of the growth line is, after all, relatively simple, and not comparable to the sutures of ammonites or even nautiloids. The possibility that similar traces have originated in a number of distantly related or distinct phyla is a strong one. (1941, pp.34-35) Despite his opinion that the growth line trace is of great importance in turritellid classification (see Text- figure 11), Marwick (1957a, p.154) also suggests that the simplest types (e.g., that of the genus Haustator) may be prone to homoplasy. Examination of juvenile and adult growth line traces for Paleocene and Eocene Coastal Plain turritellid spe- cies (Text-figure 12) shows some interspecific varia- tion, but all traces appear to be of a basically similar type: the majority show three points of inflection on the lateral aspect (adapical, medial and basal, although the presence of adapical and basal points is often vari- able within species), have an orthocline or slightly pro- socline lateral sinus of moderate depth, a shallow to moderate spiral basal sinus, and display a spectrum of apical positions from above to around the middle of the whorl; no species (with the possible exception of “Turritella” (herein Palmerella) mortoni ssp. (“рге- mortoni" Govoni, 1983; Govoni and Hansen, in press) consistently shows a lateral sinus apex below the whorl middle. They would all thus seem to agree with Guil- laume's (1924) hybrida group, as has been suggested by many previous authors (e.g. Woodring, 1928; Palmer, 1937; Bowles, 1939) (Text-figures 9,12). Marwick (19572, p.147) suggests that “Guillaume’s figures do not do justice to the difference between the sinuses of the T. hybrida group and the T. imbricataria group," and observes that Guillaume's figures (1924, p.286, figs. 7,8, and 9) “have the apex of the hybrida sinus too nearly medial. They should be distinctly adapical as plainly shown by the photographs of his plate 10” (Marwick, 1957a, p.47) (see Text-figure 9). Even while emphasizing the differences between the hybrida and imbricataria lateral sinuses, however, Marwick (19572) also seems to deemphasize them when he places in the genus Torquesia, erected by Douvillé (1929) for species in the genus Haustator Montfort, 1810 (= Guillaume's 7. imbricataria group), many Cretaceous species of Guillaume's Aybrida group (1957a, p.160). Indeed Guillaume himself (1924, 1926) states that the differences between his hybrida and im- bricataria groups are not profound. Although Guil- laume's (1924) plates do show the subtle difference in height of the apex of the lateral sinus mentioned by Marwick, my own examination of specimens of these species (Turritella hybrida and T. imbricataria) suggest that this difference is variable (Text-figure 12-ll,mm). In summary, it is not clear that the Aybrida and imbricataria lateral growth line groups of Guillaume are distinct. All lower Tertiary Coastal Plain species show a similar lateral growth line aspect which they may share with many Eurasian species from both the Cretaceous (genus Torquesia Douvillé, 1929) and low- er Tertiary (genus Haustator Montfort, 1810, and one or more probably unnamed groups; see discussion be- low). The growth line form of these Coastal Plain spe- cies would therefore appear to be a shared primitive character, not useful in sorting out their relationships with each other or with their closest relatives in Eur- asia. | | | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 31 о Y e. а. BEB "Уд uA I DD MU AB IBM IM am Mae SB ge “ре 300 SOB SB c LB doe 299 We рее P Эле ж z A RED dua Spe awe Ф Ф о о > = о о Text-figure 12.—Growth line traces of turritelline species from the Paleocene and Eocene of the Gulf and Atlantic Coastal Plains. а. 7. mortoni Conrad, b. T. hilli Gardner, c. T. “premortoni” Govoni, d. T. levicunea Harris, e. T. kincaidensis Plummer, f. T. dumblei Harris, g. T. femina Stenzel, h. T. dutexata Harris, i. T. chirena Stenzel and Turner, j. T. apita de Gregorio, k. T. lisbonensis Bowles, l, m. T. pleboides (Vaughan), n. T. creola Palmer, o. T. alveata Conrad, p. T. clevelandia Harris, q, r. T. arenicola (Conrad), s. T. danvillensis Stenzel and Turner, t. T. tennesseensis Gabb, u. T. alabamiensis Whitfield, v. T. gilberti Bowles, w. Т. rina Palmer, x. T. subrina Palmer, у. T. carinata I.Lea, z, aa, bb. T. perdita Conrad, cc. T. infans Stenzel and Turner, dd. T. rivurbana Cooke, ee. T. *prehumerosa" Govoni, ff. T. aldrichi Bowles, gg, hh. 7. humerosa Conrad, ii. T. multilira Whitfield, jj. T. eurynome Aldrich, kk. T. praecincta Conrad. Non-coastal plain species presented for comparison: ll. T. hybrida Deshayes, Eocene, Europe, mm. T. imbricataria Lamarck, Eocene, Europe, nn. T. terebra (Linnaeus), Recent, Pacific, oo. T. robusta Gryzbowski, Miocene, South America. Whorl Profile Whorl profile here refers to the overall shape of the whorl independent of the effects of the spiral ribs. AI- though in practice this can sometimes be a difficult distinction to make by eye, it is readily accomplished via x-radiographs, camera lucida or analytically from morphometric data. Ida (1952) has proposed a set of whorl profile types designated with letters. Marwick (1971) adds names to the types. I have added two more types for the system employed here (Text-figure 13). These “types” are not intended to be interpreted as discrete character states, but only as end members of a multidimensional morphological continuum useful for identification and description. Although it is one of the most conspicuous aspects ofa turritellid species, and was long the most important character in diagnosing species and supraspecific taxa, whorl shape is of uncertain status in phylogenetic anal- yses of the group because ofthe problem of homoplasy. A priori, it is reasonable to suppose that of the char- acters considered here, whorl profile might be among the most prone to homoplasy. To the degree that shell morphology is functional in gastropods generally (e.g., Signor, 1982; Hickman, 1985; Stanley, 1988) the over- all shape of the whorl might be expected to be among the first shell characters to respond to selection pressure (cf, Ponder, 1973). Also of importance is the difficulty of homologizing whorl shape, the step that must precede the identifi- cation of homoplasy. Without more developmental in- formation that is currently available, it is difficult or 32 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 ipei convex subquadrate flat sided X telescoped W telescoped - acute frustate straight sided imbricate concave keeled = campanulate == hypercampanulate Text-figure 13.—Classification of adult whorl profiles in turritelline gastropods. Lettered types C-Y first distinguished by Ida (1952), to which descriptive terms were added by Marwick (1971). Types Q, W and Z are distinguished here for the first time, as variations on types X and Y. impossible to say whether a similar whorl profile in one turritellid species is the “same” as that in another. One technique that has been successful in this regard has been use of presumably homologous primary spiral ribs on the late whorls. For example, the two species Turritella (herein Palmerella) mortoni Conrad and T. (herein Haustator) rina Palmer are both basally cari- nate in whorl profile. Yet in mortoni the carina is com- posed of the C spiral, whereas in rina it is composed of the D spiral (Allmon, 1987, 1994). Without more knowledge of the direct Cretaceous ancestors of the lower Tertiary Coastal Plain species, or of their closest relatives in Eurasia or elsewhere, it is difficult to specify the polarity of whorl profile change in these species. The possibilities can be generalized to three, which are evaluated in the phylogenetic anal- yses below; the ancestor of the Coastal Plain species could have had 1) a round, 2) an adapically carinate, or 3) a basally carinate adult whorl profile. Few data are currently available to support one or another of these alternatives. The generally non-carinate whorl shape of most Cretaceous species from the Gulf Coastal Plain (e.g., Sohl, 1960) may support primitive status for a round profile. Ifthe adapically carinate “Turritella humerosa group” is not very closely related to the other lower Tertiary Coastal Plain species, it could have in- herited its whorl shape from a Cretaceous ancestor (perhaps the adapically carinate Cretaceous species of Torquesia discussed by Douvillé (1929) and Marwick (1957a)). Apical and Pleural Angles and Shell Geometry Although formal generative parameters, such as those of Raup (1966) or Schindel (1990), represent the most thorough method of representing the geometric form of the gastropod shell, apical and pleural angles (Text- figure 8) provide a close proxy for several of the most important geometric aspects, especially in a relatively simple shell such as a turritellid. It is possible that a comprehensive study of formal geometric parameters in these or other turritellids would yield systematically useful data at the species level or above. No such anal- ysis has yet been attempted. Use of apical and pleural angles in the present study has corroborated Merriam’s (1941, p.34) opinion that these characters are useful only at the species level. Size Since size in many organisms is apparently highly responsive to local selective conditions (Calder, 1984; Stanley, 1985; LaBarbera, 1986), it is of limited value as a taxonomic character. Turritellids exhibit a range of adult sizes from 15 mm total height to more than 300 mm. Modal values for all species, fossil and Re- cent, however, seem to be between 50 and 100 mm. | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 33 Whether size increase predominates over size decrease in the phylogeny of the group generally (“Cope’s Rule") is not known. Among Coastal Plain species, larger spe- cies tend to be more heavily sculptured than smaller species and on this basis could be seen as more derived. Stanley (1973) has argued that larger species tend to be more specialized than smaller relatives, and tend not to give rise to smaller descendants but rather to become extinct without issue. See Allmon (1994) for a further discussion of the significance of size in Coastal Plain turritellids. Adult Sculpture Compared to many other gastropod groups, external shell sculpture in Turritellidae is not well developed. It consists chiefly of spiral cords, ribs, or carinae of various number and strength. Axial elements are rare and of minor importance, but they do occur over the history of the group. The Venezuelan Miocene species Turritella zuliana Hodson, 1926, for example, shows pronounced scalloping on its flared anterior carina. A number of species, notably including Cretaceous forms placed in the genus Torquesia Douvillé, 1929 as well as the Recent western Atlantic species Torcula exoleta (Linnaeus, 1758) show faint to pronounced noding or beading on the spiral ribs of late whorls. There does not seem to be an overall polarity or trend in sculpture development for Turritellidae as a group over its entire history, although the extremely carinate morphologies typical of many of the larger species in the lower Tertiary of the Coastal Plain are absent from among Recent turritellids (Allmon et al., 1990). Individual lineages, however, do seem to show minor patterns. The beaded Cretaceous species placed by Douvillé (1929) in his genus Torquesia, for example, may have been ancestral to unbeaded but similarly adapically carinate Lower Tertiary species in Eurasia as well as the western hemisphere (Rutsch, 1943; Mar- wick, 1957a). BIOGEOGRAPHIC CONTROL A major problem in any phylogenetic analysis is distinguishing between evolutionary and purely bio- geographic events (e.g., Eldredge, 1977; Schankler, 1981). Taxa showing first appearances in a region may have recently arisen from in situ ancestors, or may have immigrated from elsewhere. This is a particular prob- lem in analyses of a portion of a diverse, widespread and long-lived group such as Turritellidae. A good way to deal with the possibility of such im- migration events would be to carry out phylogenetic studies on related species in likely source areas in order to judge the likelihood that they could have contrib- uted immigrants. Such work has simply not been car- ried out for turritellids which, despite receiving sub- stantial attention from many authors over many years, remain poorly- to completly unknown in most areas of the world. Even in those areas that have been studied more intensively, the detail is usually insufficient to judge accurately the possibility of immigration to North America at particular. For example, of the three most likely source areas for immigrants into the U.S. Gulf and Atlantic Coastal Plain region in the lower Terti- агу — western Europe and the Mediterranean, northern South America and the Caribbean, and the Pacific coast of North America—only in the last region have lower Tertiary turritellids been studied in detail in the last 70 years. The most recent work on European lower Tertiary species (Cossmann, 1912; Glibert, 1973) does not de- scribe sculptural ontogeny in any detail, and so despite the likelihood (based on overall biogeographic pat- terns) that Europe and southeastern North America exchanged benthic marine taxa during the early Ter- tiary, data presently available are insufficient to in- vestigate this possibility for turritellids.! Adegoke (1977) has reported on a probably Late Paleocene macrofauna from Nigeria, discussing tur- ritellids at some length, and describing a new genus, Reymentella. His data are summarized in Table 4. Even in this case, when information is available on the apical ontogeny of some species, data are insufficient to judge possible affinities with U.S. Coastal Plain taxa. Tor- quesia adabionensis (Oppenheim, 1915) from Nigeria, for example, is very similar in whorl profile and adult sculpture to “Turritella” humerosa Conrad from beds of probably similar age in Maryland, but the apical ontogeny of neither species is definitely known. Growth lines of the African species generally agree with those of American species. Of the Paleocene African species with known apical ontogenies, Haustator nigeriensis Adegoke, 1977 is somewhat similar to both Turritella (herein Palmerella) alveata Conrad and 7. (herein Haustator) perdita Conrad from the U.S. Coastal Plain, but these species are both of Late Eocene age. (It is interesting to note that protoconchs of the three Ni- gerian species for which they are available appear to be small and multispiral, suggestive of a long plank- tonic phase and potentailly high dispersal capability.) The most detailed works on South American Ter- ! Preliminary examination of collections in the Institut Royale des Sciences Naturelle de Belgique in Brussels in early 1992 revealed a number of forms from the French, German and Belgian Tertiary that bear considerable outward similarity to Paleogene Coastal Plain species. Detailed study of these forms, particularly SEM examination of their apices, will be required before these very cursory impressions can be verified or rejected. 34 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 4.—Character states for Paleocene turritelline species from Nigeria, described by Adegoke (1977). apical spiral growth line whorl no. whorls in Species formula type profile protoconch Torquesia adabionensis (Oppenheim, 1915) ? 2A AC y Torquesia oppenheimi Adegoke, 1977 C, ВуАз7 2В АС % Haustator furoni Адероке, 1977 Či B; А, 2A R 4-5 Haustator nigeriensis Adegoke, 1977 C, B, A, 2A BC 4-5 Haustator oyawoyei Adegoke, 1977 Ci В; А»? 2А R-BC А Reymentella olaniyani Adegoke, 1977 D C BAR; 2B BC 3 * “All three primary spirals develop in close succession on the 4th to 5th whorls and remain subequal throughout ontogeny; order of appearance of spirals not determined.” (Adegoke, 1977, p. 95). ж “Five spirals appear almost simultaneously on the third spire whorl, they represent г, a, b, с, and d (?)” (Adegoke, 1977, p. 98). tiary turritellids are those of Spieker (1922) and Hod- son (1926), but in neither paper are apical sculpture patterns discussed in any detail. Numerous scattered references have been made in the literature to Carib- bean and South American turritellid species that seem to be closely related to Paleogene species from the Gulf Coastal Plain. These include forms from the Paleocene Maria Farinha Formation in Pernambuco, Brazil (Gardner, 1931, 1935; Woodring, 1972), and the Pa- leocene Soldado Formation of Trinidad (Maury, 1912, 1929; Rutsch, 1943; Woodring, 1972). Macsotay and Scherer (1972) have summarized morphological data on Oligocene-Recent turritellids from the Caribbean and northern South America, and the results of a numerical taxonomic (i.e., phenetic) analysis. Although they present information on “sculp- ture of the first post-nuclear whorls" in the form of presence/absence of “medial,” “posterior,” “anterior” or “double” spiral keels, there is insufficient evidence that what is herein referred to as apical spiral ontogeny was in fact observed in all cases. Nevertheless, their study represents a major attempt at a phylogenetic analysis of turritellids, and provides numerous hy- potheses for testing. Apical ontogenetic formulae for species as given by Macsotay and Scherer are sum- marized in Table 5. Based on these data, it appears possible that some lineages of Gulf and Atlantic Coast- al Plain turritellids experienced exchange with some or all of the proto-Caribbean basin in the early Ter- tiary. The paucity of possible relatives outside the Coastal Plain, however, suggests that this interchange may have been in the form of emmigration to the south, rather than immigration into the U.S. Coastal Plain. Merriam (1941) describes Cretaceous to Recent tur- ritellid species from the Pacific coast in detail, but his information on apical ontogeny is not very useful be- cause, as discussed above, he uses the ambiguous de- scriptive “uni-, bi- and tricarinate" system. Table 6 summarizes the principal shell characters of the 12 “stocks” identified by Merriam, and his suggestions as to possible affinities with species on the Gulf Coastal Plain. This information is crude, but suggests that members of the “Turritella” humerosa group on the Gulf and Atlantic Coastal Plains, and perhaps mem- bers of the Turritella (herein Palmerella) creola group on the Gulf Coastal Plain (e.g., T. alveata Conrad; see discussion of lineages below), could have been Pacific immigrants rather than evolved in situ. As discussed in more detail below, these conclusions are consistent with other arguments that the humerosa group may not be closely related to other Coastal Plain species. Short of the ideal specific phylogenetic information that one would like to have, some very general paleo- geographic and paleobiogeographic information is available that may give rough indications of times of high immigration into the southeastern U.S. As sum- marized in Allmon (1990), in the early Tertiary, North America (including Greenland) and Europe were still essentially continuous, allowing not only land mam- mals to be freely exchanged (McKenna, 1975), but also probably benthic marine invertebrates with no or only short planktonic phases (Berggren and Hollister, 1974, p.150). Molluscan faunas from the Danian of West Greenland, for example, contain several bivalve spe- cies comparable to species from the Paleocene Aquia Formation of Maryland and the Thanetian of the Paris Basin (Rosenkrantz, 1970, p.447). According to Koll- mann and Peel (1983,. pp.43-44), however, West Greenland Paleocene turritellids show closer affinities to European than to American forms. Davies et al. (1975, p.128) have suggested that the Paleocene Midwayan Stage faunas of the Gulf coast show greater similarities to the faunas of northwestern Europe than to those in lower latitudes. Gardner (1931, 1935), on the other hand, emphasizes the Tethyan af- finiites of the Midwayan faunas. Middle Eocene mol- luscan faunas on the two sides of the North Atlantic show both similarities and differences. Assessing the magnitude of each is hindered by lack of reliable sys- PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 35 Table 5.—Inferred apical spiral sculpture formulas for Caribbean Neogene turritellid species based on data presented by Macsotay and Scherer (1972). It is unlikely that all of these inferred formulae actually represent earliest sculpture ontogeny, but most probably represent the condition of at least relatively early teleoconch sculpture. inferred apical inferred apical sculpture sculpture Species formula Species formula abrupta Spieker, 1922 ABC, forresti Brown, 1913 ABC acropora Dall, 1889 AB,C galvesia Olsson, 1931 AB,C alowensis F. Hodson, 1926 ABC gatunensis taratarana F. Hodson, 1926 AB,C, altilira Conrad, 1857a ABC, g. gatunensis Conrad, 1985b AB,C, a. guppyi Cossmann, 1909 AB,C g. caronensis Mansfield, 1925 AB,C, a. mirandana F. Hodson, 1926 ABC, gaweaveri F. Hodson, 1926 AB,C a. urumacoensis F. Hodson, 1926 ABC, gilbertharrisi F. Hodson, 1926 ABC, alturana Spieker, 1922 AB,C hubbardi F. Hodson, 1926 ABC amaras Woodring, 1957 A,BC, illesca Olsson, 1931 ABC andreasi Williston, in Hodson, 1926 ABC infracarinata Grzybowski, 1899 A,BC, anguillana Cooke, 1919 AB,C larensis F. Hodson, 1926 AB,C bayovarensis Olsson, 1932 AB,C limonensis Olsson, 1922 AB,C berjadinensis F. Hodson, 1926 AB,C lloydsmithi Pilsbry and Brown, 1917 AB,C, bifastigata Nelson, 1870 AB,C machapoorensis Maury, 1925 AB,C b. cartagenensis Brown and Pilsbry, 1917 AB,C m. wiedenmayeri F. Hodson, 1926 AB,C b. oreodoxa Olsson, 1922 AB,C m. guarirensis F. Hodson, 1926 AB,C buchivacoana F. Hodson, 1926 AB,C, m. paraguanensis F. Hodson, 1926 AB,C b. canonensis F. Hodson, 1926 AB,C, maiquetiana Weisbord, 1962 AB,C b. colinensis F. Hodson, 1926 AB,C masasensis Marks, 1951 ABC b. warfieldi F. Hodson, 1926 AB,C matarucana F. Hodson, 1926 ABC b. socorroensis F. Hodson, 1926 AB,C mauryae F. Hodson, 1926 AB,C b. cocoditana F. Hodson, 1926 AB,C meroensis Olsson, 1931 A,BC, caleta Olsson, 1931 A,BC, mimetes Brown and Pilsbry, 1917 AB,C caparonis Maury, 1925 AB,C, montanitensis F. Hodson, 1926 AB,C cauredalitoensis F. Hodson, 1926 ABC perattenuata Heilprin, 1887 AB,C chipolana Dall, 1982 A,BC, planigyrata Guppy, 1867 АВС chiriquiensis Olsson, 1922 AB,C praecellens Brown and Pilsbry, 1917 ABC, columbiana Weisbord, 1929 AB,C, prenuncia Spieker, 1922 ABC conquistadorana Hanna & Israelsky, 1925 ABC subgrundifera Dall, 1892 ABC cornellana F. Hodson, 1926 AB,C systiolata Dall, 1915 ABC costaricensis Olsson, 1922 AB,C varicosta Spieker, 1922 AB,C crocus Cooke, 1919 ABC, variegata Linnaeus, 1758 AB,C cruziana Olsson, 1932 ABC variegata paraguanensis F. Hodson, 1926 AB,C curamichatensis F. Hodson, 1926 AB,C venezuelana F. Hodson, 1926 AB,C domingensis Brown and Pilsbry, 1917 AB,C v. quirosana F. Hodson, 1926 ABC elemensis F. Hodson, 1926 AB,C, vistana F. Hodson, 1926 AB,C, falconensis F. Hodson, 1926 ABC, y. nicholsi F. Hodson, 1926 AB,C, fica Olsson, 1932 A, BC, zuliana F. Hodson, 1926 AB,C filicarmenensis F. Hodson, 1926 AB,C tematic revisions of even the more important groups. The introduction of molluscan species from Europe to the U.S. Gulf coast may have increased through the Late Eocene, peaking at or around the Eocene-Oligo- cene boundary (Dockery, 1984). The U.S Gulf coast may well have had more in common with northwestern Europe than with the Tethys-proto Caribbean realm, and only intermittent contact with the Pacific coast of North America (Palmer, 1957, 1967; Davies et al., 1975, p.130; Keen, 1976; Allmon, 1990, pp. 118-119; Nicol, 1991; Givens, 1989). For the purposes of the following analyses I have assumed that most of the lower Tertiary Coastal Plain species discussed here are not themselves immigrants, nor are they the immediate descendants of such im- migrants. Indications of Pacific coast affinities for some taxa are consistent with other data that are independent | of biogeographic evidence, as discussed below. Sug- gestions of relationships to species from the Caribbean and South America (e.g., Macsotay and Scherer, 1972) and Europe (e.g., Glibert, 1973; pers. obs.) also need to be studied in greater detail. This is clearly not an optimal situation. As I have emphasized, data on tur- ritellids from beyond North America are severely lim- 36 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 6.—Character states for Cretaceous and Cenozoic turritelline species from California, described by Merriam (1941). possible lateral apical whorl gulf coast Stock age sinus! sculpture? profile? relative(s)* broderipiana d’Orbigny, 1840 M Mio-Rec 4b mesocostate Q cooperi Carpenter, 1864 L Mio-Rec 2c cingulate U ocoyana Conrad, 1857a M-L Mio 4b tricostate Lo altilira Conrad, 1857b E Mio-Rec 2b tricostate U Torcula spp. uvasana Conrad, 1855 E Eoc-Rec 2b bicostate (B, C, ?) С Р. alveata (Conrad) buwaldana Dickerson, 1916 E-L Eoc Y tricostate (> andersoni Dickerson, 1916 E-L Eoc 2b tricostate H-L merriami Dickerson, 1913 E-M Eoc 2b bicostate (B, C, ?) x “T.” praecincta Conrad 7 reversa Waring, 1917 Pal-M Eoc 2b tricostate X “T.” humerosa Conrad pachecoensis Stanton, 1896 L Cret-Pal 2a tricostate J-Y chicoensis Gabb, 1864 L- Cret lb bicostate? (B, C, ?) (© T. vertebroides Morton tolenasensis Merriam, 1941 L Cret la tricostate J ' Lateral sinus type designations after lettered types in Text-figure 10. ? “Bicostate” and “tricostate” as explained in text. **mesocostate"': “in early ontogeny a more or less median primary, which exists as a carina or angulation in adolescent stages, often later becoming obsolete” (Merriam, 1941, p. 6). 3 Whorl profile type designations after lettered types in Text-figure 13. 4 Suggested relationships are Merriam’s (1941, pp. 38-43). ited, and as a result these biogeographic conclusions can serve as no more than hypotheses for future testing. ANALYSIS OF COASTAL PLAIN SPECIES Taxa Considered The following discussion concentrates on those tur- ritellid species most closely related to two common, conspicuous and historically important forms, “Тиу- ritella mortoni Conrad, 1830” and “Turritella hume- rosa Conrad, 18355", and is restricted to Paleocene and Eocene occurrences (Tables 7-9). Recognition of Species Groups Species related to “Turritella mortoni Conrad” and “Turritella humerosa Conrad” are recognized by their possessing characters shown by these two forms and lacking in all or most other species. Principal characters and character states for the Paleocene and Eocene Coastal Plain turritellid species that share such char- acters are given in Table 9. Based on these data and the foregoing discussion of character analysis, four spe- cies groups or lineages can be recognized on the basis of shared derived characters. These are referred to be- low as the “mortoni group", the “creola group”, the "rina group”, and the “humerosa group". Relationships Among Species Groups: a Cladistic Approach In attempting a cladistic analysis of these species, the first and in many ways most difficult decision is the choice of an outgroup. Eldredge (1979, p.171) has emphasized that there is no hard and fast rule govern- ing the selection of the correct outgroup for comparison with the taxa of interest. The method of outgroup com- parison should be viewed rather as a continuous pro- cedure of pair-wise comparison. Potential sister groups are proposed on the basis of previous or independent phylogenetic information and analysis, and alterna- tives are tested by how well they accord with available data. At this point in the study of Turritellidae, so little detailed morphological documentation and phyloge- netic analysis have been done that there is no guarantee that any named group chosen as an outgroup (e.g., a supraspecific taxon from Europe or the Pacific coast of North America) will, when combined with the Coastal Plain taxa of interest, comprise a monophy- letic, or at least not polyphyletic, group. 2) There are no corroborated hypotheses of possible evolutionary relationships between these lower Tertiary species and particular Cretaceous forms from the Coastal Plain or elsewhere. Those links that have been proposed (e.g., Merriam, 1941; Marwick, 1957a; Sohl, 1960, p.71) are based on little more than similar whorl profile. For these reasons, I have chosen to use hypothetical outgroups in the following analyses. This avoids hav- ing to wait for taxa in other areas and times to be examined in detail before Coastal Plain species can be considered at all. The number of relevant characters is low and can easily be specified for a variety of out- groups; these character lists can then be used at a later date to test relationships between actual monophyletic taxa and these coastal plain species. Three alternative cladograms for the relationship PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 57 Table 7.— Described species of turritellid gastropods (excluding species assigned to the genus Mesalia Gray, 1842) from the Paleocene and Eocene of the U.S. and their taxonomic placement in the present paper. * indicates species for which well-preserved shell apices are known. Names in brackets are not recognized as separate taxa in the present paper. “Turritella mortoni Group” (placed in Palmerella n. gen.) alabamiensis Whitfield, 1865* alveata Conrad in Wailes, 1854* apita de Gregorio, 1890* arenicola (Conrad, 1865)* [a. branneri Harris, 1891] [a. danvillensis Stenzel and Turner, 1940*] chirena Stenzel and Turner, 1940* clevelandia Harris, 1892* creola Palmer in Harris and Palmer, 1947* dumblei Harris, 1895a* dutexata Harris, 1895a* femina Stenzel in Renick and Stenzel, 1931* ГЕ oligoploka Stenzel in Renick and Stenzel, 1931] hilli Gardner, 1935 levicunea Harris, 1896 lisbonensis Bowles, 1939 mortoni mediavia Bowles, 1939 mortoni mortoni Conrad, 1830* pleboides (Vaughan, 1895)* mortoni postmortoni Harris, 1894a potomacensis Clark and Martin, 1901 mortoni “premortoni” Govoni, 1983 (Govoni and Hansen, in press)* stenzeli Allmon, n. sp. “Turritella rina Group” (placed in Haustator Montfort, 1810) carinata I.Lea, 1833* [carina palmerae Bowles, 1939] cortezi Bowles, 1939 fischeri Palmer in Richards and Palmer, 1953 gilberti Bowles, 1939* infans Stenzel and Turner, 1940* martinensis Dall, 1892 perdita Conrad, 1865* [perdita jacksonensis Cooke, 1926*] [perdita lowei Cooke, 1926*] rina Palmer, 1937* [rina carolina Palmer, 1937] [rina wechesensis Bowles, 1939] rivurbana Cooke, 1926 subrina Palmer, 1937 subtilis Kellum, 1926 tennesseensis Gabb, 1860* vaughani Bowles, 1939 “Turritella humerosa Group” (placed in “Turritella”) aldrichi Bowles, 1939* biboracesis Gardner, 1945 claytonensis Bowles, 1939 eurynome Whitfield, 1865 gardnerae LeBlanc, 1942 humerosa Conrad, 1835b* multilira Whitfield, 1865 praecincta praecincta Conrad, 1864* praecincta virginiensis Allmon, n. ssp. “prehumerosa” Govoni, 1983 (Govoni and Hansen, in press)* toulmini Allmon, n. sp. Incertae Sedis bunkerhillensis Palmer in Harris and Palmer, 1947 dobyensis Dockery, 1980* kincaidensis Plummer, 1933 mcbeanensis Bowles, 1939 mingoensis Bowles, 1939 nasuta Gabb, 1860* [n. brazita Stenzel and Turner, 1940] [n. felli Bowles, 1939] [n. houstonia Harris, 1895] [n. smithvillensis Bowles, 1939] nerinexa Harris, 1895a obruta Conrad, 1833 ola Plummer, 1933 plummeri Stenzel and Turner, 1940 polysticha Stenzel and Turner, 1940 saffordi Gabb, 1860 turneri Plummer, 1933* among species groups in these Paleogene Coastal Plain species are shown in Text-figure 14. Several points concerning these cladograms are important: 1) Outgroups in the three cladograms differ only in their whorl profile (i.e., “round,” “basally carinate" and “adapically carinate"). This is consistent with the foregoing discussions and the data summarized in Ta- bles 1 and 9 that suggest that the closest relatives of the Coastal Plain species had an apical sculpture for- mula of C, B, Аз, a hybrida-imbricataria-type lateral growth line aspect, a moderately deep basal sinus, and a moderate size and apical angle. These character states are therefore presumed to be plesiomorphic in the Coastal Plain species. 2) These cladograms are based on the weighting scheme described above, without which many other arrangements would be equally plausible. For example, if apical ontogeny and whorl profile were weighted equally and an outgroup with a round whorl profile were used, the creola group would be primitive rather than derived, and the С, В, A; apical sculpture formula of the rina group would be derived (and homoplastic) rather than primitive. А 3) Тһе only synapomorphic character possibly unit- ing all Coastal Plain species in a monophyletic group is the lack of beaded sculpture. This character change is proposed very tentatively and only because many Cretaceous species, particularly in Eurasia, bear such sculpture prominently (e.g., Douvillé, 1929) and some of the lower Tertiary Coastal Plain species vestigially. Given the uncertainty of the Cretaceous ancestry of Coastal Plain species, however, this is a very difficult hypothesis to support at the present time. Even if the ancestry were more definite, loss characters are gen- 38 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 8.—Geographic range and stratigraphic duration of Paleocene and Eocene turritelline species from the U.S. Gulf and Atlantic Coastal Plains. Ranges were calculated by connecting the geographic centers of counties containing one or more reported occurrences of the species with straight lines, forming maximum area convex polygons, and measuring the enclosed areas with an electronic digitizing tablet. Exceptions were made to allow for the known shape of the Lower Tertiary shoreline and continental shelf edge (i.e., ranges were not calculated over area presumed to be land or continental slope; using the paleogeographic interpretations of Murray [1961], Rainwater [1964], Toulmin [1977], Breard [1978], Siesser [1984b], Hansen [1987] and Dockery [1988]). Species known from only two localities are given ranges equal to the distance between the two in km?. Species known from single localities, or from multiple localities within a single county, are arbitrarily given ranges of 10 km?. Ranges are given for time slices defined by correlative lithostratigraphic units across the Coastal Plain; for species known from more than one such temporal unit, ranges are given for each unit individually. Duration is the estimated total lifetime of a species over the entire interval of deposition of all stratigraphic units in which it occurs, and so are maximum estimates given the known stratigraphic distribution. Values were calculated based on best available biostratigraphic dates for formational boundaries, using the absolute timescale of Berggren et al. (1985). Abbreviations of lithostratigraphic units: AQ, Aquia Formation; BA, Bashi Marl; BM, Black Mingo Group; BS, Brightseat Formation; CH, Castle Hayne Limestone; CL, Clayton Formation; CM, Cook Mountain Formation; DBT, Doby’s Bluff Tongue; GS, Gosport Sand; KI, Kincaid Formation; LL, Lower Lisbon Formation; LO, “Logansport Formation" (=Lime Hill); MB, Moodys Branch Formation; MCB, McBean Formation; MLM, Matthews Landing Marl Member of Porters Creek Formation; MR, Marthaville Formation; NA, Naheola Formation; NF, Nanafalia Formation; PC, Porters Creek Formation; PE, Pendleton Formation; QC, Queen City Formation; RK, Reklaw Formation; SCB, Stone City Beds; SR, Shark River Formation; TS, Tuscahoma Formation; TSBL, Bells Landing Marl Member of Tuscahoma; TSGL, Greggs Landing Marl Member of Tuscahoma; UL, Upper Lisbon Formation; WB, White Bluff Formation; WE, Weches Formation; WP, Wills Point Formation; YZ, Yazoo Formation. range | duration range | duration Species time unit (km?) (my) Species time unit (km?) (my) alabamiensis NA 10 7.6 levicunea NA 10 7.6 "WP-PC 66 Бе 10 KI-CL 404,054 lisbonensis CM-UL-MCB 248,239 6.0 aldrichi LO-NA 679 EG mcbeanensis MCB 10 3.0 WP-PC 66 mingoensis BM 10 3.0 KI-CL 9,122 mortoni mortoni AQ 2,092 3:2 alveata MB-WB 63,965 (0:5 multilira MR-NF 4,338 2.6 apita GS 10 0.3 nasuta CM-UL-MCB 206,987 13 arenicola YZ 205 у nerinexa WE 57,086 4.0 MB-WB 146,785 obruta GS 10 0.3 carinata GS 50 5.0 ola KI 10 4.0 CM-UL 74,160 perdita MB 37,526 0.5 chirena WE 10 3.0 pleboides CM 64 3.0 claytonensis CE 208 4.0 polysticha WP 10 1.0 clevelandia WZ 201 7/30 mortoni postmortoni PE-TS 39,990 3.2 MB 45,350 MR-NF 45,832 cortezi CM 45,832 210 potomacensis AQ 2,092 522 creola MB 2,196 0.5 praecincta praecincta PE-TS 39,714 5.8 dobyensis DBT 10 1.0 praecincta virginiensis AQ 2,092 3.2 dumblei CM 8,306 3.0 “prehumerosa” BS 10 15 dutexata CM 122,794 3.0 mortoni “premortoni” BS 10 1:5, еигупоте PE-TS 14,443 5.8 rina CM-UL-MCB 130,581 7.0 MR-NF 890 rivurbana MB 7,453 0.3 femina WE 65 3.0 saffordi CL 1,446 4.0 BA 10 stenzeli KI 42 4.0 gilberti BA 35.923 3.2 subrina CM-UL-MCB 123,016 7.0 hilli KI 10 4.2 subtilis CH 10 5:0) houstonia CM 20:177 3.0 tennesseensis CL 13,720 4.0 humerosa AQ 2,092 292 toulmini CL 10 4.0 infans CM B 3.0 turneri QC 10 3.0 SCB 10 RK 622 kincaidensis KI 346 4.2 vaughani MCB 10 2.0 erally undesirable because of the high chance of ho- moplasy (cf, Hecht, 1976). 4) It seems likely that the species of the humerosa group are not particularly closely related to the other Coastal Plain species, for the following reasons: i) possible relationships between the humerosa group and turritellid species from elsewhere are more sup- ported than for the other Coastal Plain species groups (e.g., Merriam (1941) for the Pacific coast, Rutsch (1943) for the Caribbean and South America); ii) as PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 39 Table 9.—Character states for Paleocene and Eocene turritellid species from the Gulf and Atlantic Coastal Plains discussed in this paper. Conventions and abbreviations as in Table 1. growth line apical sculpture basal lateral sinus apical pleural whorl Species formula Sinus type depth apex angle angle angle profile size alabamiensis C1 B2 АЗ 4 2 M B OR-P 20 J M aldrichi СІ B2 АЗ 4-5 2 M B OR 14 Q M alveata C1 B1 A2 3-4 2 M A-B OR 18 D M apita C1 B1 А2 3 2 M B OR 20 V S-M arenicola C1 B1 A2 3-4 2 M A-B OR-P 23 20 С M carinata C1 B2 A3 4 2 M B n 15 19 J-L M-L chirena C1 Bl А2 4 2 S-M B OR 27 J S-M claytonensis к; ? 19 Q-X M-L clevelandia C1 B1 A2 4-6 2 M B Р 16 Ј M cortezi i ? 16 27 U M-L creola СІ B1 A2 3 1-2 M A-B OR-OP 22 С M dobyensis C2 B1 A3 3 2 M B OR 15 12 С м dumblei C1 B1 A2 4—5 2 M B P-OR T 17 J M dutexata C1 B1 A2 A 1 M A-B OR 21 C-J M eurynome ў 6? 2 M B Р 14 Q-X L femina C1 B1 A2 2-3 1-2 M B P-OR 23 C-J M gilberti C1 B2 АЗ 4-5 1 M A-B OR 20 D-J S-M hilli ve 4 2 M-D B OR-P 20 J-Y M-L humerosa C1 B2 A3 3 2 M A-B OR 17 X L-VL infans C1 B2 A3 9 1-2 M A OR 19 J S kincaidensis ES 4-6 2 м B Р 15 J M levicunea ? 4 2 S-M B B 22 30 H-L M-L lisbonensis ? 3 2 M B P-OR 19 7 C-D M-L martinensis C1 B3 A2? (1) 2 2 M B OR 14 L M-L mortoni mediavia ? 4 2 M B OR 21 23 J-Y L mortoni mortoni СІ Bl А2 4 2 м В P-OR 22 J-Y L-VL multilira 4 S 2 M A-B P 16 9 X-Q L perdita C1 B2 A3 3—4 3? M B P 16 J-D M pleboides C1 B2 A3 3 1 M A OR 20 18 E S-M mortoni postmortoni " 4 1-2 M A-B Р 23 21 Y-Z L-VL praecincta 3 4-6 2 M B ју 19 14 X-W L-VL prehumerosa C1 B2 A3 (2) de 2 M A-B P 17 15 Q-X M-L mortoni “premortoni” C1 B2 АЗ/СІ ВІ A2 (3) 4 2 M B-C P 22 J-Y M-L rina СІ B2 АЗ 4 2 м А-В OR 18 L-Y L rivurbana ? 4-5 2 M A-B OR 16 L-U M stenzeli ? 4 1 M A OR 17 J-U M subrina ? 4 2 M B OR 23 U L tennesseensis C1 B2 A3 С 2-3 M B OR-P 17 U S-M toulmini ү 3 2 M B OR 10 X-C M-L already mentioned, Spiller (1977) states that “ Turri- tella” humerosa has a multispiral protoconch, sugges- tive of a more extended planktonic phase and higher potential for dispersal, and this may have allowed more contact with regions outside the Coastal Plain; iii) the adapically carinate whorl profile of the humerosa group would not appear to be morphogenetically very close to the basally carinate profile of the rina group, with which the humerosa group shared a common apical sculpture formula; it seems more reasonable to assume that if these two lineages are related at all, that they developed their distinct whorl profiles independently from a noncarinate ancestor. Relationships within Species Groups: a Stratophenetic Approach Following from the discussion in the Methods sec- tion above, the stratigraphic ranges of the component, species of the four lineages are plotted on an evolu- tionary tree in Text-figure 15, together with a set of hypothesized branching relationships. These relation- ships are based on stratophenetic reasoning (Bretsky, 1979; Gingerich, 1979; Lazarus and Prothero, 1984): morphologically similar species found in close geo- graphic and stratigraphic proximity are assumed to be closely related. Exact branching order within a cluster 40 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 mortoni group A humerosa group rina group creola group m r ү: МЕ eee epp ah 55 ст 5855 2 SESS е Se fe о g а е g $824 Eo 58568 8 “9 5 o 2 ERE E E 885988 BUS. 6 Шала ыы. JB Wut suu о €sl52 BEES Cece Seiten Cee ЕШЕсі 5584 5 S SSE SESE SPSLSGESESES S929 оров 8888 9 зсобкафео 4852825 885555 525887458 LELE | 5a>b Зр—а T Збос - 1 mortoni humerosa group rina group creola group group res E r 1-3 TIT a tg EST. SESA n 5892 2 g @ = 2 88%- 67 55 ESPE ізде, б 255 VECES шесе Z f a Боа трдн 922555555 5555 3 5 [d Ss go Еж 22030 <= 8 з Ва ELSIE 88855 09585 950655 БЕБЕ ғ--------- rivurbana o на 3a Ы | 6a-b Haan o ЕЧ 5 5 5 | [== — biboraensis T0355 | 9b—»a ШЕ тогіопі rina group creola group group humerosa group C en r Y Ў 1 Lu f pores | 585: SES 2528 a S 2 sg” 2% o ва? 52 2 5420 TU MM 8 55555 бе og 50508 ERRE p 88868556 585285, 55552 Seow 5059 5555 85286596645 о EEE ES 52258 Бє ЕБЕ g g Б 5 389546 босар $35su50tis5 5555 525855555 а 2628458 95555 90589488868 БЕБЕ асоаЕаеЕб5 | | oii + | T 2a—b F5a-b tap 8 а-эс la—b Text-figure 14.— Three alternative cladograms showing possible relationships of four of the principal species groups of turritellid gastropods from Paleocene and Eocene sediments of the U.S. Gulf of such species is determined by the balance among the three variables of morphology, stratigraphy and geography. For example, of three species in the same area and stratigraphic horizon, the two sharing the most similarities are assumed to be more closely related; of three equally dissimilar species, the two occurring clos- est in either time or space are most closely related. Individual supraspecific relationships based on this stratophenetic approach are discussed in the system- atic section below. Supraspecific Taxa in the Present Study A persistent pattern in the literature on North Amer- ican Cenozoic turritellids is the reluctance of investi- gators to assign species to formal supraspecific groups. Palmer (1937), for example, while noting that “The Claibornian species do not belong to Turritella s.s. according to the lines of growth," uses the name Tur- ritella *in the broad sense of the genus." The reasons for this reluctance are summed up by Merriam (1941, p.35): According to the standards of taxonomy herein proposed, the European and austral genera, subgenera and sections could be eval- uated only by study of complete and well-preserved “genotype” material. This has been done for only one or two of these. Until this study is accomplished for all, it is considered inexpedient to propose additional generic or subgeneric names, or, in America at least, to attempt to use most of those already existent. Allison and Adegoke (1969, p.1250) similarly state that “Much additional study of earliest apical ontogeny is yet necessary before Bowles’ [1939] groups can be reconstituted naturally. Until this information is avail- able, formal nomenclature should not be provided for his groupings as they now stand” (emphasis in origi- nal). The only commonly discussed basis for suprageneric — and Atlantic Coastal Plains. These cladograms differ only in the nature of the outgroup, which is hypothetical in each case. A. Out- group with beaded sculpture, paucispiral protoconch, C,B,A, apical sculpture formula, and round whorl profile. B. Outgroup with beaded sculpture, paucispiral protoconch, C,B,A, apical sculpture formula, and adapically carinate whorl profile. C. Outgroup with beaded sculpture, paucispiral protoconch, C,B,A, apical sculpture formula, and basally carinate whorl profile. See text for further discussion. Characters and character states are as follows: 1. beaded sculpture, a. present, b. absent; 2. protoconch, a. paucispiral, b. multispiral; 3. whorl profile, a. basally carinate, b. round, c. adapically carinate; 4. apical sculpture formula, a. C,B,A,, b. С,В,А,; 5. strength of B spiral in adults, a. well developed, b. weak to absent; 6. strength of C spiral, a. moderate, b. very strong to carinate. Cladogram 1 or 2 is preferred based on likely form of Cretaceous ancestor (i.e., outgroup) and probability that humerosa group is only distantly related to other Coastal Plain clades (i.e., that similarities are shared primitive char- acters). See text for further discussion. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 4] | PALEOCENE | EOCENE Midwayan Sabinian Claibornian Jacksonian = Ф A D elc = 3 70 = = ar Jm 2 о 5 = w [^] o ә|9 O o ¿IU 9 (еі = о | 5 13512 а бан ба ва 525223 eS x => = S. = ш 0 | a ES m du = == 2х0 2. |р а а је х| o 36,8. ~ olo| N | б o о || о [95 |= = 230 |00 = |Б о | 2 |-2і-91-ж 135 | џ| © ue O-O |e NO I es »EOSZIO|G аео |69 |5910 21915 о | © + = |9 » ==" 339 Z| © |=: 90/50/5249 5 |4|ш = Фан ® өсі ару o |» = II ~ jso С = єз a Ф о ~ ~ = < 3 3 a о RM ~ Ф 5) w 3 ж 3 ES c = (а Еч femina dutexata Vii» [Y | ! Wes ep У а а i. Wo. | apita eis T =~ 1 1 AY = е Шы : arenicola Ф Ф lisbonensis |- - -еявтниланнинан | li | 2 i creola ! pleboides заны mortoni mediavia | m — рњ clevelandia IN Ww in plummeri | | kincaidensis hilli | levicunea í cortezi ; Я Реут ВА EN i 4 rivurbana gl eg | AX пије le E 1 stenzeli ; | ен kun subrina ı І | — ť ng | } | и | | tennesseensis a 1 rina | EOS | | ner 1 4" [fischeri 1 alabamiensis ! = : carinata Jer f Be и ial 1 | gilberti | infans perdita | O) DINNER нн ---------- AE m — — — — — ышын | aldrichi Жаы eurynome multilira Text-figure 15.—Phylogenetic tree of turritellid species discussed in this paper. Known stratigraphic ranges are represented by solid lines, inferred relationships by dashed lines. Stratigraphic relationships based on Toulmin (1977), Dockery (1980), and Carter (1984). assignment of American Paleogene species has been the growth line classification of Guillaume (1924) (Text- figure 9). As pointed out by Dollfus (1926), Stewart (1927) and Woodring (1928), Guillaume fails to cor- relate his groups with already existing supraspecific names, some of which appear to match his descrip- tions. The conventional pairing (e.g., Woodring, 1928) is as follows: groupe de 7. terebralis Lamarck = Turritella Lamarck, 1799 groupe de 7. imbricataria Lamarck = Haustator Montfort, 1810 groupe de 7. turris Basterot = Archimediella Sacco, 1895 groupe de T. subangulata Brocchi = Torculoidella Sacco, 1895 Guillaume’s “groupe de T. hybrida” apparently does not match up with an existing name, and it is this growth line group into which American Paleogene spe- cies have usually been placed (e.g., Palmer, 1937; Bowles, 1939). The only name ever proposed for the species with hybrida-type lateral sinuses is Turricula, proposed by Douvillé (1929) as a subgenus of Turritella. The name Turricula, however, is a much preoccupied name, and is unavailable (Bowles, 1939, p.270; Marwick, 1957a, p.161). Douvillé also proposes the genus name Tor- quesia to contain Eurasian Cretaceous species with lat- eral growth line traces intermediate between those of Guillaume’s 7. imbricataria and Т. hybrida groups and bearing prominent adapical carinae and beaded sculp- ture on the spiral ribs. The use of Torquesia is extended by Rutsch (1943, pp.163ff) to include a number of species from the lower Tertiary of the Americas as well as Eurasia. Marwick (1957a) suggests placing these and other similar Tertiary species in Zspharina Vialov and Soloun, 1936, as a subgenus of Torquesia. As made clear by Allison and Adegoke (1969), however, Js- pharina is a nomen nudum, because neither type spec- imens nor type locality is designated and adequate de- scriptions and illustrations of the type species are not provided. Govoni (1983) and Govoni and Hansen (in press) have followed Rutsch (1943) in suggesting that Torquesia be used to include lower Tertiary species (including the species here included in the “humerosa group") with adapically carinate whorls and Aybrida- type lateral sinuses. Despite these opinions, I believe that the described differences between the nominate genera Torquesia and Haustator are ambiguous. Marwick (1957a, p.154) states that the type species of the genus Haustator, H. imbricatarius (Lamarck) from the Upper Eocene of England, has a growth line with a "moderately deep, moderately oblique lateral sinus of a general shape that is widespread in the Turritellinae", and suggests that homoplasy was likely in this rather simple growth line type. As already mentioned, Guillaume (1924) em- PALAEONTOGRAPHICA AMERICANA, NUMBER 59 phasizes that the differences between his T. hybrida and T. imbricataria types are small; my own exami- nation of these species supports this opinion (Text- figure 12). The basal sinus of T. imbricataria is some- what deeper than that of T. hybrida, but even this is somewhat variable in the specimens I have examined. According to Marwick (19572, p.154), the type species of Haustator has a three-whorled protoconch and an apical sculpture formula of C, В, Аз. Although the apical ontogeny is apparently unknown for both the type species of Torquesia from the Cretaceous and for any of the less sculptured forms assigned by Marwick to the genus from the lower Tertiary, an as yet unde- scribed species (“prehumerosa” Govoni (1983); Go- voni and Hansen, in press) placed by Govoni in Tor- quesia, has a protoconch of about two whorls and an apical sculpture formula of d C, B, Аз. In erecting Torquesia as a genus separate from Haustator, Dou- villé (1929) places most emphasis on the whorl profile; species assigned to Torquesia have prominent adapical carinae while those in Haustator typically have basal carinae. Marwick (1957a, p.160), however, states that these differences in whorl profile “аге not taxonomi- cally important.” Nevertheless, Marwick declares Tor- quesia to be a “useful genus for many Cretaceous spe- cies of Guillaume’s Aybrida group.” In summary, Torquesia s.s. and Haustator do not appear to differ in any features other than the presence or absence of beaded sculpture and the vertical position of carination on the whorl, both of which are features Marwick (1957, p.160) declares to be “apparently of no more than specific value." If the unbeaded lower Tertiary forms are included in Torquesia, as suggested by Govoni (1983) and Govoni and Hansen (in press), then one of even these two diagnostic characters is lost. Based on the data currently available, Guilllaume's T. hybrida and T. imbricataria lateral sinus types do not appear to be sufficiently discrete or non-overlap- ping criteria on which to base supraspecific designa- tions. The genera Haustator and Torquesia, therefore, can be said to share a common lateral sinus form. Further work may show that some of these species differ in form of the basal sinus. Data currently avail- able, however, indicate that such differences are minor. The genus Torquesia, if it is to be recognized at all, should be restricted to the species for which it was erected: the chiefly Eurasian, adapically carinate Cre- taceous species bearing prominent beaded sculpture. In the lower Tertiary of the western hemisphere, adap- ically carinate species largely lacking beaded sculpture, but sharing the Aybrida-imbricataria growth line type (e.g., “Т.” humerosa Conrad) should be placed in a genus-group separate from Torquesia. Itis possible that many of the Eurasian species discussed under Jsphar- PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 43 Text-figure 16.—Three possible supraspecific classifications of Paleocene and Eocene Coastal Plain turritellids. Diagrams on the left assume either of cladograms A or B in Text-figure 14; diagrams on right assume cladogram C in Text-figure 14. H = humerosa group, R = rina group, M = mortoni group, C = creola group. At, A2. Two supraspecific taxa, one including the humerosa group, the other the mortoni, creola and rina groups. B1, B2. Two supraspecific taxa, one including the humerosa and rina groups, the other including the mortoni and creola groups. C1, C2. Three supraspecific taxa, one including the humerosa group, one the rina group, and the other the mortoni and creola groups. This is the scheme adopted here. ina by Marwick may also belong here. (If it should prove to be the case that “Turritella” humerosa has a small, multispiral protoconch suggestive of a long planktonic interval (as indicated by Spiller, 1977), such wide dispersal of this clade would be more likely.) The choices for supraspecific assignment of the Pa- leocene and Eocene Coastal Plain species are sum- marized in Text-figure 16, superimposed on the pos- sible cladograms discussed in the previous section. The three choices are as follows: 1) Based on whorl profile, the mortoni, rina and perhaps creola species groups could be placed in the genus Haustator Montfort (choices la and 1b), but this would mean that species with two distinct apical on- togenetic formulae (C, B, A; and C, B, A,) would be included in a single genus. As discussed above, most previous work, as well as the present study, indicates that lineages can be recognized on the basis of apical ontogeny. 2) If apical ontogeny is given the most emphasis, the 44 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 rina group might be placed with the humerosa group in a supraspecific taxon separate from the mortoni and creola groups (choices 2a and 2b). Against this alter- native is the possibility that the humerosa group is, as discussed above, not particularly closely related to the other lower Tertiary Coastal Plain species, but rather to similar species from Eurasia or elsewhere. The growth line form and lack of beaded sculpture that unite the Coastal Plain species discussed here could both be shared primitive characters. 3) A final alternative is to recognize three separate supraspecific taxa among the lower Tertiary Coastal Plain species. This is the option I have adopted here. The first taxon, consisting of the “mortoni group” and the “creola group,” is characterized by a basally carinate to rounded whorl profile, C, B, A, apical sculpture formula, hybrida-imbricataria-type lateral sinus, moderately deep basal sinus, and a protoconch of 1-2.5 whorls with small P1. It appears that no name is currently available for such a taxon. I therefore pro- pose the name Palmerella for it (see p. 45). The second taxon, consisting of the “rina group," is characterized by prominent basal carination formed by a well developed D spiral, an apical sculpture for- mula of C, B, A, with a tendency for the B spiral to become obsolete late in ontogeny, especially in later members of the group, Aybrida-imbricataria-type lat- eral sinus, moderately deep basal sinus, and a proto- conch of 1-2.5 whorls with a small P1. These species can be assigned to the genus Haustator Montfort, 1810, as a derived branch that eventually gave rise to the genus Torcula Gray, 1847 in the Late Eocene (Allison and Adegoke, 1969). The third group of species includes “Turritella” hu- merosa Conrad and its apparent relatives. These spe- cies are characterized by moderate to pronounced adapical carination, an apical sculpture formula of C, B, A,, hybrida-imbricataria-type lateral sinus, mod- erately shallow basal sinus, and a protoconch possibly varying from 1-1.5 to 3-4 whorls with a small РІ. Information is presently insufficient to assign these spe- cies to a formal genus-level taxon with confidence, and I reluctantly follow tradition by assigning it to “Тиу- ritella” sensu lato. SYSTEMATIC PALEONTOLOGY In the descriptions that follow, the locality infor- mation given represents all locality records known to me for each species. Abbreviations of localities refer to those listed in Appendix 2. At least one source for each record is indicated in the parentheses following the abbreviation. Abbreviations for these sources are as follows: A— Allison (1965, 1967); AA—Allison and Adegoke (1969); ANS— ANSP collection; B— Bowles (1939) (documen- tary specimens for most of Bowles’ records are in the USNM collection); C—Cooke (1943); CA—Campbell (1992); D-Dockery (1977, 1980); G—Gardner (1935); GO —Govoni (1983), Govoni and Hansen (in press); GSA — Geological Survey of Alabama collection; HA— Harris (1894b); HP— Harris and Palmer (1947); HU— Hughes (1958); JA—private collection of James Allen, Alexandria, Louisiana; L—LeBlanc in Barry and LeBlanc (1942); MCZ—author’s collection, deposited in the Department of Invertebrate Paleontology of the Museum of Comparative Zoology; P—Palmer (1937); PB—Palmer and Brann (1966); PL—Plummer (1933); RP—Richards and Palmer (1953); RS—Renick and Stenzel (1931); S40—Stenzel (1940); S53—Stenzel (1953); ST—Stenzel and Turner (1940, 1942); T— Toulmin (1977); TLM—Toulmin and LaMoreaux (1963); TX—Texas Memorial Museum/Bureau of Economic Geology, Austin; US—USNM collection; V—Vaughan (1896); VC—Van Nieuwenhuise and Colquhoun (1982); VS— Veatch‘and Stephenson (1911); WW-Wassem and Wilbert (1943); W-Wilbert (1953); WD-— Ward (1985). In the locality lists, ? after a locality number means that the species identification of the specimen(s) is un- certain. (?) after a locality number means that the lo- cality is not definitely known. As in Tables 1 and 9, the size classification of Mar- wick (1971) is used in referring to maximum shell height of each species: Very Large = > 100 mm, Large- 50- 100 mm, Medium = 20-50 mm, Small=10-20 mm, Very Small=< 10 mm. Sculptural ontogenies for most species are shown as _ “Marwick Diagrams" in Text-figure 7. Growth line traces are shown in Text-figure 12. Protoconch mea- surements are given in Table 3, and other character states summarized in Table 9. Measurements given in tables for locality numbers represent /argest individuals observed from these lo- calities. Apical and pleural angles have been measured on type specimens, unless otherwise indicated. See Text-figures 3,4,6 and 8 for illustration of mor- phological terms used in descriptions. Abbreviations of Repository Institutions ANSP— Academy of Natural Sciences of Philadel- phia, Philadelphia, PA, USA FGS— Florida Geological Survey type collection; now held at Florida Museum of Natural History, University of Florida, Gainesville, FL, USA FMNH-UC- Field Museum of Natural History, Chicago, IL, USA PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 45 FLMNH-UF--Florida Museum of Natural History, University of Florida, Gainesville, FL, USA GSA — Geological Survey of Alabama, Tuscaloosa, AL, USA LSU — Museum of Geoscience, Louisiana State Uni- versity, Baton Rouge, LA, USA MCZM — Department of Mollusks, Museum of Comparative Zoology, Cambridge, MA, USA MCZIP — Department of Invertebrate Paleontology, Museum of Comparative Zoology, Cambridge, MA, USA MGS — Mississippi Office of Geology, Jackson, MS, USA PRI—Paleontological Research Institution, Ithaca, NY, USA TMM/TBEG — Texas Memorial Museum/Bureau of Economic Geology, University of Texas, Austin, TX, USA UCMP- Museum of Paleontology, University of California, Berkeley, CA, USA UNC- Department of Geology, University of North Carolina, Chapel Hill, NC, USA USGS —U.S.Geological Survey, Reston, VA, USA USNM--U.S.National Museum, Washington, DC, USA Systematics Class Gastropoda Cuvier, 1797 Subclass Prosobranchia Milne-Edwards, 1848 Order Mesogastropoda Thiele, 1925 Superfamily Cerithiacea Fleming, 1822 Family Turritellidae Lovén, 18472 Genus PALMERELLA, new genus Type Species. — Turritella mortoni Conrad, 1830 Diagnosis. —Small to very large turritellids with round to basally carinate adult whorl profile and apical sculpture formula of С, B, A;. Description. —Shell small to very large, of 10-25 whorls. Protoconch of 1-2.5 whorls with small P1. Apical sculpture formula C, B, A;. Profile of adult whorls round to basally carinate. Sculpture consists of multiple simple, moderately developed spiral ribs. Apical angle usually equal to, but occasionally less than, 2 The name “Turritellidae” was introduced independently by Clark (1851) and Woodward (1851), with priority usually attributed to Woodward (e.g., Knight, 1960, p.1317). The first use of a formal suprageneric name for the group, however, is apparently that of Lovén (1847, p.194) as “Turritellea”. According to Article 1161, ICZN, “A family-group name of which the latinized suffix is incor- rect is available with its original authorship and date, but with a corrected suffix” (ICZN, 1985, p.27), and thus Loven is the correct authority for the name Turritellidae. pleural angle. Lateral sinus of the “hybrida-imbrica- taria” type, basal sinus moderately deep and simple. Stratigraphic and Geographic Distribution. — U.S.Gulfand Atlantic Coastal Plain (VA, NC, SC, GA, AL, MS, LA, TX, AR), Mexico, Trinidad? South America?; Lower Paleocene— Upper Eocene. Etymology. —Named after Dr. Katherine V.W. Palmer in recognition of her many contributions to knowledge of Coastal Plain fossil mollusks, especially turritellid gastropods. Palmerella alveata (Conrad in Wailes, 1854) Plate 5, Figures 7-9 Turritella alveata Conrad in Wailes, 1854, pl.17, Ғ.7 (reprint, 1939, pl.4, f.7); Conrad, 1856, p.263 (reprint, 1939, p.9); Harris, 1894b, p.120, 169, 181; Bowles, 1939, p.306, pl.32, f.1; Stenzel and Tur- ner, 1942, card 43; Palmer in Harris and Palmer, 1947, p.288, pl.36, f. 7-12; Brann and Kent, 1960, p.897; Allison, 1965, p.671; Palmer and Brann, 1966, p.980; Dockery, 1977, p.44, pl.3, £.5. Mesalia alveata (Conrad). Conrad, 1865a, p.33; 1866, p.25. Turritella (Haustator) alveata Conrad. Cossmann, 1912, p.118. Description. —Shell medium sized, of 18 to 22 whorls. Maximum observed whorl diameter 12.2 mm. Apical angle 18°; apical and pleural angles approximately equal. Sutures slightly to moderately incised. Protoconch erect, homeostrophic, turbinate, of 0.75-1.0 smooth, round- ed, somewhat inflated whorls; Pl small. Apical sculp- ture formula C, B, A,. Primary spirals B and C ap- pearing simultaneously in lower У; of whorl, abruptly becoming prominent and of equal strength. Primary spiral A appearing within 1.0 whorl on upper % of whorl, remaining slightly weaker than Band С for 1– 2 whorls, after which all primaries of approximately equal strength. Secondaries r, s and t appearing be- tween primaries around whorl 8-9. Teleoconch whorls wider than high. Profile of early teleoconch whorls roughly evenly convex; later whorls rounded and slightly basally convex to straight-sided. Area between C spiral and suture deeply excavated and smoothly concave. Lateral aspect of growth line trace slightly prosocline, with upper and lower inflection points. Lat- eral sinus moderately deep with apex about at whorl middle. Spiral and basal sinuses roughly symmetrical and relatively shallow. Measurements. —See Table 10. Table 10.—Measurements of Palmerella alveata (Conrad). Ab- breviations: MD = maximum diameter (including carina if present) in mm., MH = maximum height, in mm., WN = whorl number, to the nearest 0.5. MD MH WN ANSP 13213 lectotype 12:2 48.1 17 ANSP 13213 paratype КӨ 44.5 15:5 46 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Stratigraphic and Geographic Distribution. — MS, LA: Moody's Branch Formation; AR: White Bluff For- mation; lower Jacksonian Stage, Upper Eocene. Type Locality. — Moody's Branch, Jackson, Hinds County, Mississippi (MS-HN-4). Other Localities. – MS-HN-1b (B), MS-HN-1c (D), MS-HN-1d (B), MS-HN-2 (D), MS-HN-3 (D), MS- HN-4b (B), MS-CL-3 (B), LA-FR-1 (B), LA-GR-1 (B), LA-GR-1a (B), LA-LS-1 (B), LA-CL-1b (D), LA-CL- 1c (D), LA-CL-1e (D), LA-CL-1g (D), LA-CL-1h (D), LA-CL-2 (D), LA-CL-3 (D). Type Material. —lectotype and paratype of T. alveata Conrad ANSP 13213. Remarks/Discussion. — The type specimens are much larger than average for the species; mean height of specimens I have examined is closer to 30-40 mm than to 40-50 mm. In sculpture and whorl profile Palmer- ella alveata is most similar to P. creola (Palmer) and P. arenicola (Conrad), also from the Moodys Branch Formation. All three of these species show generally rounded whorls, widest in the lower half, and narrow excavated bands just above the suture. Palmerella al- veata has the least inflated whorls of the three. P. al- veata often co-occurs with but is less abundant than Haustator perdita (Conrad) at most exposures of the Moodys Branch; at Bunker Hill Bluff on the Ouachita River (LA-CL-2) and Riverside Park in Jackson, Mis- sissippi (MS-HN-6), however, P. alveata is the most common turritellid and among the most common ma- crofossils. Palmerella apita (de Gregorio, 1890) Plate 4, Figures 6,7 Turritella carinata H.C.Lea, 1841, p.96, pl.1, f.10; 1849, p.107; Meyer, 1887, p.54, pl.3, £.1,1a; Harris, 18955, p.10; not I.Lea, 1833; p.129. Turritella apita de Gregorio, 1890, p.123, pl.11, £.8, £.26, copy T. carinata H.C.Lea, 1841, f.27a, 27b, copies Meyer, 1887; Coss- mann, 1893, p.29; Harris, 1895b, p.82; Palmer, 1937, p.195, pl.24, f.1,3,7,10; in Harris and Palmer, 1947, p.285; Bowles, 1939, p.275, pl.31, £.4; Stenzel and Turner, 1942, card 44; Brann and Kent, 1960, p.898; Glibert, 1962, p.103; Allison, 1965, p.672; Palmer and Brann, 1966, p.980-981; Toulmin, 1977, p.301, pl.50, f.2. Description. — Shell small to medium sized. Maxi- mum observed whorl diameter 10.0 mm. Apical angle 20*; apical and pleural angles approximately equal. Su- tures only moderately incised. Protoconch and earliest teleoconch whorls not seen; Palmer (1937, p.196) and Allison (1965, p.672) report an apical sculpture for- mula of d C, B, a;. On earliest whorls examined, pri- mary spiral B quickly becoming most prominent, forming single sharp keel on middle of whorl. C spiral much less prominent and A even less. Whorl profile between C spiral and lower suture relatively straight, sloping evenly inward, and marked by fine, evenly Table 11.— Measurements of Palmerella apita (de Gregorio). Ab- breviations as in Table 10. MD MH WN USNM 494978 8.0 26.0 10.5 MCZIP 29290 109) 24.8 6 MCZIP 29291 8.0 21.8 729 spaced spiral ribs. Lateral aspect of growth line trace orthocline to slightly prosocline with upper and lower points of inflection. Lateral sinus moderately deep with apex at about middle of whorl. Spiral and basal sinuses relatively shallow. Measurements. —See Table 11. Stratigraphic and Geographic Distribution. —AL: Gosport Sand; GA: McBean Formation? (?); LA: Cook Mountain Formation (?); upper Claibornian Stage, Middle Eocene. Type Locality.—Claiborne Bluff, Alabama River, Monroe County, Alabama (AL-MO-1a). Other Localities. — AL-CL-4 (MCZ), GA-JO-1 (VS) ?, LA-WI-1 (V) ? Type Material.—holotype of T. apita De Gregorio PRI 26438; lectotype of T. carinata H.C.Lea ANSP 13173. Remarks/Discussion. —I have not been able to con- firm records for this species from beds of Cook Moun- tain age, and for the present consider it to be restricted to the Gosport Sand, in which it is relatively rare. I have not seen a well preserved apex; if the apical sculp- ture formula given by Palmer (1937) and Allison (1965) is correct, it is another illustration of an apparently “unicarinate” species not being truly so on its earliest whorls (see also Palmerella arenicola (Conrad), below). This is in contrast to “Turritella” dobyensis Dockery, 1980, which appears to be truly unicarinate. Possible phylogenetic relationships of Palmerella apita are dis- cussed below under P. arenicola. Palmerella arenicola (Conrad, 1865b) Plate 5, Figures 3-6 Turritella plebeia Say. Owen, 1860, pl.9, f.6; not Say, 1824, p.125. Mesalia ? arenicola Conrad, 1865b, p.141, pl.10, f.11; 1866, p.11 {locality not Oregon as given]. Turritella carinata Y.Lea. Heilprin, 1884, p.37; Call, 1891, p.8; not 1- Бей, 1955 Turritella arenicola (Conrad). Dall, 1891, footnote in Call, 1891, p.8; 1892, p.232,310; Stewart, 1927, p.354; Bowles, 1939, p.275, pl.31, £.5-7; Stenzel and Turner, 1942, card 45; Palmer in Harris ? The status of the “McBean Formation" is unclear. It is most probably a sandy facies of the upper Claibornian Santee Limestone, and equivalent to the upper part of the Lisbon/Cook Mountain Formation and/or the Gosport Sand in Alabama (see Huddlestun and Hetrick, 1985; Campbell, 1992, 1995; Dockery and Nystrom, 1992; Fallaw and Price, 1992). PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 47 and Palmer, 1947, p.281, pl.34, f.8-11; Brann and Kent, 1960, p.898—9; Allison, 1965,p.673-4; Palmer and Brann, 1966, p.981; Dockery, 1977, p.43-4, pl.3, f.16,17; 1980, p.82, pl.56, f.7,8,10 Toulmin,1977, p.335, pl.62, f.12,13. Turritella arenicola branneri Harris, 1894b, p.169, р1.6, £.7; Schuch- ert, Dall, Stanton, and Bassler, 1905, p.676; Palmer, 1937, p.197, pl.23, f.1,2; in Harris and Palmer, 1947, p.283, pl.34, £.2,3,6,7; Stenzel and Turner, 1942, card 49; Wilbert, 1953, p.123, pl.2, £.5; Brann and Kent, 1960, p.899; Allison, 1965, p.674. Turritella arenicola danvillensis Stenzel and Turner, 1940, p.841, pl.47, £.4,5; 1942, card 58; Palmer in Harris and Palmer, 1947, p.284, pl.34, f.1,4,5, pl.35, f.1; Brann and Kent, 1960, p.899,900; Allison, 1965, p.672-673; Palmer and Brann, 1966, p.981-982. Description. —Shell medium sized, of 18 to 22 whorls. Maximum observed whorl diameter 12.0 mm. Apical angle 25°; pleural angle sometimes slightly less than apical angle. Sutures only slightly incised. Protoconch erect, homeostrophic, turbinate, of about 2.0 smooth, rounded, somewhat flattened whorls; P1 small. Apical Sculpture formula C, В, A. Primary spirals B and C appearing approximately simultaneously as broad prominence at around middle of whorl. B quickly be- coming more prominent, forming single, conspicuous, sharp carina on middle of whorl. Primary A appearing shortly after B and C, but remaining faint. B remaining most prominent for up to 10 whorls, after which other primaries and secondaries emerging abruptly, covering upper surface of intermediate and late whorls with nu- merous, evenly spaced, subequal spirals, changing whorl profile from keeled to rounded and slightly basally con- vex. B and C becoming equally prominent and the most conspicuous sculptural elements. Area between C spiral and suture widening and becoming slightly concave. Lateral growth line trace generally prominent, usually prosocline, with upper and lower inflection points and relatively shallow lateral sinus; apex just above middle of whorl. Spiral and basal sinuses rela- tively shallow. Measurements. —See Table 12. Stratigraphic and Geographic Distribution. —MS, LA: Moody’s Branch Formation, Yazoo Formation (Dan- ville Landing Member); AR: White Bluff Formation; TX, MX: horizon unknown; GA,SC: McBean Formation’ ?; upper Claibornian 2 and Jacksonian Stages, Upper Eocene. Type Locality.—Garland Creek, Clarke County, Mississippi (MS-CL-3). Other Localities. -MS-CL-15 (D), MS-CL-18 (D), MS-YZ-4 (B), MS-YZ-5 (B), MS-HN-1 (D), MS-HN-4 (D), LA-GR-1 (B), LA-GR-1d (B), LA-RA-1 (B), LA- RA-2 (B), LA-CL-6 (D), AR-BR-1(B), AR-DR-1 (B), AR-JF-1 (W,US), AR-SF-1 (B), AR-SF-2 (B), AR-SF-4 (B), AR-CL-1b (H), AR-CL-4 (H), AR-CL-8 (H), TX- ST-1 (B), TX-SA-2b (B), MX-TA-1 (B), MX-TA-2 (B), * See Footnote 3, p.46. Table 12.— Measurements of Palmerella arenicola (Conrad). Ab- breviations as in Table 10. MD MH WN USNM 489002 12.0 41.0 10.5 USNM 495144 9:0 36.0 13.3 MS-YZ-5 10.2 34.2 10 MS-CL-3 10.4 32.3 11 LA-CA-1 10.0 22.9 3 MX-NL-3 (B), GA-BU-1 (VS), GA-JO-1 (VS), SC-O-1 (CA). Type Material.—lectotype of T. arenicola Conrad ANSP 15536; holotype of T. a. branneri Harris USNM 135141; syntypes of T. a. danvillensis TMM/TBEG 20962, 20963. Remarks/Discussion. — Allison (1965, pp.672-3) states that the upper Jacksonian subspecies Turritella arenicola danvillensis Stenzel and Turner has an apical sculpture formula of C, B, A, but that arenicola s.s. shows С, B, Аз. I have not see specimens with the latter pattern; the description given above is for spec- imens from the Moodys Branch Formation in Clarke County, Mississippi, and suggests that both Yazoo and Moodys forms show similar apical ontogenies and be- long to a single taxon. The similar apical sculpture and early strength of the B spiral in Palmerella apita (de Gregorio) and P. ar- enicola suggest a close relationship between these two species. The rounded whorl profile and strength of the two lowest spirals on later whorls in P. arenicola sug- gest a relationship with the lineage of P. femina (Sten- zel), including P. dutexata (Harris), and P. lisbonensis (Bowles) (see discussion of these forms below). The smaller pleural angle in arenicola in particular may indicate relationship with P. lisbonensis. Palmerella apita and P. arenicola may represent descendants of the femina lineage in which the B spiral became stron- ger than the C spiral, this pattern persisting to adult- hood in apita, and ending much earlier in the ontogeny of arenicola. Palmerella chirena (Stenzel and Turner, 1940) Plate 3, Figures 7-9 Turritella chirena Stenzel and Turner, 1940, p.838, р1.47, f.3; 1942, card 54; Palmer and Brann, 1966, p.984. Description. —Shell small, of 10 to 15 whorls. Max- imum observed whorl diameter 7.0 mm. Apical angle 27°; apical and pleural angles approximately equal. Su- tures only slightly incised. Teleoconch whorls wider than high. Protoconch suberect, homeostrophic, tur- binate, of 2.0-2.5 smooth, rounded whorls; Pl small. Apical sculpture formula С, B, A;. Primary spirals B and C appearing approximately simultaneously as faint, 48 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 13.— Measurements of Palmerella chirena (Stenzel and Tur- ner). Abbreviations as in Table 10. MD MH WN TMM/TBEG 20960 holotype Gat 19:5 975 broad prominence just below middle of whorl. Within 1.0 whorl A appearing very faintly, B and C diverging and profile between C and suture becoming excavated and concave. Shape ofearly to intermediate teleoconch whorls generally rounded, of late whorls straight-sided and subquadrate. Lateral aspect of growth line trace orthocline, showing only lower point of inflection and relatively shallow lateral sinus whose apex is above whorl middle. Spiral and basal sinuses of moderate depth. à Measurements. —See Table 13. Stratigraphic and Geographic Distribution. — TX: “Cane River Formation" (see remarks below); Clai- bornian Stage, Middle Eocene. Type Locality. —road from Chireno to Hwy. 21, Nacogdoches County, Texas (TX-NA-8). Other Localities. — TX-NA-7 (MCZ). Type Material. —holotype of 7. chirena Stenzel and Turner TMM/TBEG 20960. Remarks/Discussion. — The stratigraphic position of beds in East Texas identified by Stenzel and Turner as *Cane River" is unclear. The Cane River Formation was named by Spooner (1926) for beds in Louisiana occurring stratigraphically above those of the Wilcox Group and below the Sparta Sand. Ellisor (1929) rec- ognizes the Cane River in the subsurface in East Texas, underlying the Reklaw, and it is so indicated on a USGS correlation chart (Wilmarth, 1930). Eargle (1968) states that the Louisiana Cane River beds correlated with beds of the Weches, Queen City and Reklaw in East Texas. I have not been able to relocate the type locality of P. chirena near the town of Chireno; all outcrops in ditches between the town and Highway 21 lack macrofossils. A good outcrop containing fairly well preserved specimens of P. chirena (locality TX- NA-7), however, occurs on Hwy. 21, approximately 7 km west of Chireno. Exposed here are approximately 5 m of highly glauconitic, clayey sand containing small to medium sized oysters identifiable as Cubitostrea smithvillensis (Harris) and/or C. lisbonensis (Harris), both of which are characteristic of the Weches For- mation. Lithologic and faunal evidence is therefore consistent with assigning this outcrop and, provision- ally at least, the type occurrence of P. chirena to the Weches Formation. Palmerella chirena shares with P. dumblei (Harris) from the overlying Stone City and Cook Mountain beds a similar apical sculpture formula and basally carinate whorl shape, but differs in its greater apical angle and somewhat smaller size. Both P. chirena and P. dumblei may be derived from a form similar to specimens here noted for the first time as occurring in the upper Sabinian Bashi Formation and assigned to P. femina (Stenzel) (see p. 51). The origins of Palmerella chirena, and its apparent relative P. dumblei, are unclear. These Middle Eocene species share with the Late Paleocene P. mortoni (Con- rad) the C, B, A; apical sculpture formula and a basally carinate whorl profile, but are much smaller. They share with the Middle-to-Late Eocene species of the “creola group" the same apical sculpture formula and rela- tively modest size (see Text-figures 14,15). Based on size alone, it does not seem likely that mortoni was actually ancestral to these forms. There are thus two choices for the origin -of chirena and dumblei. Either they are derived from the “creola group," in which case their basally carinate whorl profile is not homol- ogous to that in mortoni and its close relatives, or they are derived from a lineage that split off from the “mor- toni group," probably in the earliest Paleocene, before the time of P. mortoni mediavia (Bowles) and P. mor- toni ssp. (“premortoni” Govoni, 1983; see Govoni and Hansen, in press). Based on the reasoning and confi- dence estimates discussed above, and summarized in Table 2, I favor the former alternative. It seems un- likely that a group arising in the Coastal Plain region in the Early Paleocene would leave no trace until the Middle Eocene. (The alternative view is, however, sup- ported by the previously unreported presence of P. femina (Stenzel) from the Bashi Marl of Alabama (see below), which provides a potential Lower Eocene an- cestor for the creola lineage. It is possible that, because chirena and dumblei are small, relatively uncommon Texas forms, the somewhat more poorly preserved Pa- leocene-lowermost Eocene Texas macrofossil record has thusfar concealed their derivation from a mortoni- like ancestor.) The “creola group” itself has no definitely known ancestor on the Coastal Plain before the earliest Eo- cene, and by the argument just given might on these grounds be assumed not to have originated in the re- gion. This may well prove to be the case. It is possible, however, that one of the Early Paleocene species with a round whorl profile (e.g., “Turritella” ola Plummer, 1933) for which the apical spiral formula is not known might be ancestral to the creola lineage, and these should be investigated in addition to seeking possible ancestral forms are sought outside the Coastal Plain. Palmerella clevelandia (Harris, 1894b) Plate 5, Figures 1,2 Turritella perdita? Dall, 1891, footnote in Call, 1891, p.8, fide Harris, 1894b, p.92. | \ / \ | | ) ) ) PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 49 Turritella mortoni Conrad. Call, 1891, p.8, fide Harris, 1894b, p.92; not Conrad, 1830. Turritella clevelandia Harris, 1894b, p.170, pl.4, f.9; 1899a, p.74 in part [part = 7. gilberti Bowles]; Vaughan, 1896, p.50; Schuchert, Dall, Stanton and Bassler, 1905, p.676; Palmer, 1937, p.202, pl.26, f.6,7; in Harris and Palmer, 1947, p.290, pl.36, f.1-6; Bowles, 1939, p.308, pl.31, 69,12; Stenzel and Turner, 1942, card 56; Wilbert, 1953, p.123, pl.2, f.4; Brann and Kent, 1960, p.907; Glibert, 1962, p.95; Palmer and Brann, 1966, p.984—5; Dockery, 1977, p.44. Turritella (Haustator) Clevelandica [sic] Harris. Cossmann, 1912, p.118. Description. —Shell small, of 10 to 15 whorls. Max- imum observed whorl diameter 5.5 mm. Apical angle 16°; apical and pleural angles approximately equal. Su- tures moderately incised. Apical sculpture formula C, B, A,. Primary spirals B and C appearing approxi- mately simultaneously just below middle of whorl, of equal strength and beginning to diverge. Primary A appearing within 1.0 whorl, and B becoming more prominent for one to two whorls, after which C be- coming slightly more prominent and remains so. Three primaries retaining relative strengths, giving late whorls a tricarinate, subquadrate form. Secondaries present but remaining faint. Lateral growth line trace proso- cline with upper and lower inflection points and rel- atively shallow lateral sinus with apex around whorl middle. Spiral and basal sinuses relatively shallow. Measurements. —See Table 14. Stratigraphic and Geographic Distribution. — AR: White Bluff Formation; MS: Moody's Branch For- mation; LA: Moody's Branch Formation, Yazoo For- mation (Danville Landing Member) TX?, GA?: ho- rizons unknown; Jacksonian Stage, Upper Eocene. Type Locality. — White Bluff, Arkansas River, Jef- ferson County, Arkansas (AR-JF-1). Other Localities. — AR-CL-1b (B), AR-CL-4 (B), AR- CL-4B (B), AR-CL-7 (B), AR-BR-1 (B), AR-SF-1 (B), AR-SF-2 (D), AR-SF-4 (B), MS-YZ-1 (D), MS-YZ-2 (MCZ), LA-GR-1 (JA), LA-CA-1 (HP), LA-CA-2 (HP), GA-JO-1 ? (VS), GA-GL-2 ? (C). Type Material. —lectotype of T. clevelandia Harris USNM 498010, syntypes USNM 135142. Remarks/Discussion. — This species is relatively rare. In whorl form and adult sculpture it is most similar to Haustator gilberti (Bowles), specimens of which were originally identified as clevelandia, but the apical sculp- ture of the two forms is very different. P. clevelandia differs from the other Moodys Branch turritellids of Table 14. — Measurements of Palmerella clevelandia (Harris). Ab- breviations as in Table 10. MD MH WN USNM 498010 a 24.1 1365) similar apical sculpture in its subquadrate whorl profile and three-ribbed adult sculpture. It may be more close- ly related to small basally carinate forms of Claibornian age, such as P. chirena (Stenzel and Turner). P. clevelandia is one of two Moodys Branch turri- tellids to occur in the Danville Landing Member of the Yazoo Formation, the other being P. arenicola (Con- rad). After noting the similarity of P. dumblei (Harris) to the Oligocene species “Turritella” carota MacNeil, MacNeil (in MacNeil and Dockery, 1984, p.53) sug- gests that carota may have been derived from P. cley- elandia. Palmerella creola (Palmer in Harris and Palmer, 1947) Plate 4, Figures 1-5 Turritella creola Palmer in Harris and Palmer, 1947, p.286, pl.35, f.2,4,5; Brann and Kent, 1960, p.909; Allison, 1965, p.671-2; Palmer and Brann, 1966, p.985. Description. —Shell small to medium sized, of 15 to 20 whorls. Maximum observed whorl diameter 10.0 mm. Apical angle 22°, pleural angle may be slightly less than apical angle. Sutures moderately incised. Pro- toconch erect, homeostrophic, turbinate, of approxi- mately 1.0 smooth, rounded whorls; Pl small. Apical sculpture formula C, B, A,. Primary spirals C and B appearing, faintly at first, around or slightly below mid- dle of whorl, quickly becoming prominent. C may be equal in strength or slightly weaker than B. Primary A appearing two to three whorls later, and remaining faint throughout ontogeny. Sculpture of adult whorls consisting of two prominent spirals (B and C) on lower Y of whorl, separated by a slightly weaker secondary (T). Primary A and secondaries above whorl middle approximately equal in strength but weaker than sec- ondary (T). Teleoconch whorls wider than high, evenly rounded and relatively inflated in profile. Whorl profile below C spiral slightly excavated and concave leading to suture. Lateral aspect of growth line trace orthocline with upper and lower inflection points and moderately deep sinus whose apex is above middle of whorl. Spiral and basal sinuses moderately deep. Measurements. —See Table 15. Stratigraphic and Geographic Distribution. — ПА, TX, MS: Moody’s Branch Formation; lower Jacksonian Stage, Upper Eocene. Table 15.—Measurements of Palmerella creola (Palmer). Abbre- viations as in Table 10. MD MH WN PRI 4579 holotype 325 10.0 Tiles PRI 4580 paratype 239 9.8 =) Type Locality.—Montgomery Landing, Red River, Grant Parish, Louisiana (LA-GR-1). Other Localities. —TX-SA-2b (HP), MS-YZ-2 (MCZ). Type Material. —holotype of 7. creola Palmer PRI 4579, paratypes PRI 4577, 4578, 4580. Remarks/Discussion. — Palmer (in Harris and Palm- er, 1947) suggests that P. creola is most closely related to “the 7. dutexata Harris stock of the lower Clai- borne.” In its rounded, convex whorl profile, creola most closely resembles P. femina (Stenzel), which is close to dutexata (see below). Among Moodys Branch species, creola is most similar to P. arenicola (Conrad) and P. alveata (Conrad). P. creola is most common at its type locality, the now inaccessible outcrop at Montgomery Landing, but it was apparently not even very abundant there. Mr. James Allen of Alexandria, Louisiana kindly made available to me his collection of turritellids from Mont- gomery. It was, he says, accumulated in approximately 50 collecting trips, but includes only approximately 40 specimens of P. creola. Palmerella dumblei (Harris, 1895a) Plate 6, Figures 4-6 Turritella dumblei Harris, 1895a, p.81, pl.9, £.7; Kennedy, 1895, p.123,124,127-29; Palmer, 1937, p.202 in part, pl.26, £.11,15, not f.10 [= T. sp., fide Palmer and Brann, 1966, | p.985]; Bowles, 1939, p.303, pl.32, £9,19; Stenzel and Turner, 1942, card 59; Palmer and Brann, 1966, p.985. Description. — Shell small to medium sized, of 15 to 20 whorls. Maximum observed whorl diameter 9.7 mm. Apical angle 17°; apical and pleural angles ap- proximately equal. Sutures only slightly incised. Pro- toconch of approximately 1.75 smooth, rounded whorls; P1 small. Apical sculpture formula C, B, A;. Primary spirals B and C appearing abruptly and at equal strength in middle and lower М of whorl, re- spectively. Primary A slightly weaker, appearing with- in 0.75 whorl. Lateral whorl profile straight-sided on early whorls, straight-sided to slightly concave on later whorls with whorl base expanding to form a rounded basal carina. Primary spirals relatively reduced on late whorls, combining with secondaries to cover whorl with numerous evenly spaced, fine, subequal spirals. Whorl profile of late whorls slightly concave with a rounded basal carina separated from lower suture by narrow, slightly concave area marked by two to three minor spirals. Lateral aspect of growth line trace or- thocline to slightly prosocline with upper and lower inflection points. Lateral sinus relatively shallow with apex at around whorl middle. Spiral and basal sinuses relatively shallow. Measurements. —See Table 16. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 16.—Measurements of Palmerella dumblei (Harris). Ab- breviations as in Table 10. MD MH WN TMM/TBEG 35661 holotype 8:2 243 8.5 TMM/TBEG 35662 paratype 9.65 АРАНЫ 10.0 TMM/TBEG 35663 paratype US 25.8 13.0 TMM/TBEG 35663 paratype 6.0 23.0 1509, ТММ/ТВЕС 35665 рагаїуре 9:29 13.6 9:9 USNM 498003 ы 26.5 1259 USNM 498003 8.0 16.5 S Stratigraphic and Geographic Distribution. — TX: Stone City Beds, Cook Mountain Formation (Whee- lock Member); middle Claibornian Stage, Middle Eo- cene. Type Locality. —'*Moseley's Ferry”, Stone City Bluff, Brazos River, Burleson County, Texas (TX-BU-1). Other Localities. TX-AT-2 (B), TX-AT-3 (B), TX- BR-1 (B), TX-RO-11 (B), TX-SA-2d (US). Type Material.—holotype of T. dumblei Harris TMM/TBEG 35661, paratypes TMM/TBEG 35662, 35663 (2 specimens), 35664, 35665. Remarks/Discussion. —Its apical sculpture places P. dumblei in the genus Palmerella. Its basally convex whorl profile suggests a closer relationship to P. clev- elandia (Harris) than to the more evenly rounded-con- vex forms such as P. femina (Stenzel) and P. creola (Palmer). Palmerella dutexata (Harris, 1895a) Plate 3, Figures 10-12 Turritella dutexata Harris, 1895a, p.82, pl.9, Ғ.8; Palmer, 1937, p.198, in part, pl.26, f.1,2,4,8,9; in Harris and Palmer, 1947, p.287; not Bowles, 1939, p.285, р1.31, £2 [= P. femina (Stenzel)]; Stenzel and Turner, 1942, card 60; Brann and Kent, 1960, p.910, in part, Nos. 2910,2911 only, fide Palmer and Brann, 1966; Palmer and Brann, 1966, p.986. Turritella cf. dutexata Harris. Palmer and Brann, 1966, p.986. Description. —Shell small to medium sized, of 10 to 15 whorls. Maximum observed whorl diameter 8.0 mm. Apical angle 21°; apical and pleural angles ap- proximately equal. Sutures only slightly incised. Pro- toconch erect, homeostrophic, turbinate, of approxi- mately 1.5 smooth, rounded whorls; P1 small. Apical sculpture formula C, B, A,. Primary spirals Band C appearing as broad prominence on lower !^ of whorl, quickly diverging and becoming equally prominent. Primary A appearing within two to three whorls but remaining faint throughout ontogeny. D spiral com- monly weakly developed on later whorls but other spi- rals usually absent. Whorl profile above B concave; all teleoconch whorls bicarinate in aspect. Lateral aspect of growth line trace prosocline with upper and lower inflection points and relatively shallow lateral sinus; PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 51 Table 17.—Measurements of Palmerella dutexata (Harris). Ab- breviations as in Table 10. Table 18.—Measurements of Palmerella femina (Stenzel). Abbre- viations as in Table 10. MD MH WN MD MH WN PRI 2912 9.5 26.1 6.5 TMM/TBEG 20965 holotype 10.0 23.0 7.5 PRI 2913 6.9 12.0 6.5 TMM/TBEG paratype 959 “ae M TMM/TBEG paratype 120 360 5.5 USNM 139021 53 8 5.0 apex above whorl middle. Basal sinus not visible on Specimens examined. Measurements. —See Table 17. Stratigraphic and Geographic Distribution. — TX,LA,MS: Cook Mountain Formation; AL: Upper Lisbon Formation, Gosport Sand ?; upper Claibornian Stage, Middle Eocene. Type Locality. — Elm Creek, Lee County, Texas (TX- LE-11). Other Localities. -TX-RO-12 (В), TX-AN-2 (B), TX-AN-3 (B), MS-CL-1 (B), MS-CL-2 (B), MS-CL-5 (B), MS-CL-16 (B), MS-CL-17 (B), MS-NE-10 (B), MS- NE-11 (B), MS-NE-1 (B), MS-NE-2 ? (P), LA-BI-1 (B), LA-BI-2 (B), LA-BI-3 (B), LA-CB-1 (B), LA-JA-2 (ST), LA-WI-1 (В), LA-WI-2 (B), AL-MO-1a ? (P), AL-CL- 14 (MCZ). Type Material.—holotype of 7. dutexata Harris TMM/TBEG 1974 lost (fide Erickson, 1982). Remarks/Discussion. —See under P. femina, below. Palmerella femina (Stenzel in Renick and Stenzel, 1931) Plate 3, Figures 3-6 Turritella femina Stenzel in Renick and Stenzel, 1931, p.87,89,107 [reference only], pl.6, f.14; Plummer, 1933, p.644,647,815; Palm- er, 1937, p.203, pl.26, f.5, copy Stenzel, 1931; Palmer in Harris and Palmer, 1947, p.286; Stenzel and Turner, 1940, p.830, р1.46, f.11-13; 1942, card 64; Palmer and Brann, 1966, p.987. Turritella dutexata Harris. Bowles, 1939, p.285, pl.31, f.2; not T. dutexata Harris, 1895a, p.82. Turritella femina oligoploka Stenzel in Stenzel and Turner, 1940, p. 832, pl.46, f.7-10; Stenzel and Turner, 1942, card 88; Palmer and Brann, 1966, p.987. Description. —Shell medium sized, of 15 to 20 whorls. Maximum observed diameter 12.0 mm. Apical angle 22°; apical and pleural angles approximately equal. Su- tures moderately incised. Protoconch erect, homeos- trophic, turbinate, of approximately 1.75 smooth, rounded whorls; Pl small. Apical sculpture formula С, B, A). Primary spirals B and C appearing approx- imately simultaneously just above and just below whorl middle, respectively, moving within 1.0 whorl to about middle and lower Уз of whorl, respectively. Primary A appearing within 0.5 whorl of beginning of B and C, equalling C in strength within one to two whorls. Pri- mary spirals commonly bearing beading or minor scal- loping on early teleoconch whorls. Secondaries ap- pearing above and below A on later whorls, where A is weaker than B and C. Upper Y. of whorl thus covered by numerous, approximately equally spaced spirals of moderate strength, lower Y dominated by two pri- maries lacking associated weaker spirals. Profile of late whorls rounded and evenly convex. Lateral growth line trace prosocline with upper and lower inflection points and relatively shallow lateral sinus; apex around whorl middle. Spiral and basal sinuses very shallow. Measurements. —See Table 18. Stratigraphic and Geographic Distribution.—TX: Weches Formation, (Viesca Member); AL, GA: Bashi Formation; upper Sabinian Stage, middle Claibornian Stage, Upper Paleocene— Middle Eocene. Type Locality.—Cobb Branch, Robertson County, Texas (TX-RO-6). Other Localities. — YX-RO-12 (US), TX-LO-12 (ST), TX-LO-13 (RS), AL-CO-1 (MCZ), GA-RA-3 (MCZ). Type Material.—holotype of T. femina Stenzel TMM/TBEG 20965, numerous unnumbered para- types TMM/TBEG; syntypes of T. femina oligoploka TMM/TBEG 20969, 20970, 20971, 20972. Remarks/Discussion. —In their extreme morpholog- ical variants, P. femina and P. dutexata (Harris) are quite distinct. On its largest whorls dutexata has fewer prominent spirals, and the lower half of the whorl is dominated by only two; the upper half bears only a few relatively weak lines, and is markedly concave in profile. Although they also show two prominent spiral ribs on the lower half of the whorl, the adult whorls of femina characteristically bear numerous additional spirals of equal to slightly lesser strength, and the entire’ whorl is convex and slightly inflated in profile. There appear to be intermediates between these two ex- tremes, however, represented in part by the subspecies femina oligoploka Stenzel and Turner, which has fewer spiral ribs on the upper half of later whorls and a less inflated profile than femina s.s. Stenzel and Turner noted that femina s.s. and f. oligoploka “are end mem- bers of a series” of transitional forms. Both taxa have been recognized only in the Weches Formation of Tex- as, but from adjoining counties. This morphological series, however, can be extended upward to forms usu- ally identified as Turritella dutexata Harris from the 39 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Wheelock Member of the Cook Mountain Formation. Many of these specimens show the reduction in spiral ribbing described in femina oligoploka, and a less in- flated whorl profile. Specimens from the upper Cook Mountain and Upper Lisbon Formations tend to show the extreme reduction of sculpture on the upper whorl and the characteristic concave profile. As noted by Stenzel and Turner, dutexata does not occur with either femina or f. oligoploka, strengthening an hypothesis of evolutionary transition from one to the other. Below the Weches, the femina lineage also appears to be recognizable in specimens I have collected from the upper Sabinian Bashi Formation in eastern Ala- bama and western Georgia. Although apices are not preserved, these specimens clearly show the distinctive double carina on the lower half of the whorl, and have rounded concave whorl profiles, and thus are clearly distinguishable from the only other turritellid so far reported from the Bashi, Haustator gilberti (Bowles). Recognition of these specimens as belonging to P. fem- ina helps to fill an important temporal gap in the his- tory of Palmerella in the Coastal Plain Paleogene, and shows that at least two lineages survived the apparent crisis for turritellids that seems to have occurred near the end of the period of deposition of the Tuscahoma Formation, when all of the large, characteristic Sabi- nian turritellid species disappeared. A Sabinian representative of the lineage leading to femina also may make the search for Paleocene Coastal Plain ancestors more fruitful. The only form currently recognized that may fill this role is “Turritella” ola Plummer from the Lower Paleocene Clayton Forma- tion of Texas. This species shows two strong carinae on the lower half of the whorl, and a rounded, convex whorl profile. Unfortunately, it is known from only a single specimen from a locality that seems to have disappeared. The only feature that calls into doubt the relationship of Palmerella femina to other Coastal Plain turritellid species is the form of its basal growth line trace, which shows a very shallow sinus. The basal sinus of P. du- texata (Harris) is unfortunately not visible on any of the specimens I have examined. The basal sinus on the closely related form P. lisbonensis (Bowles), as well as that of P. pleboides (Vaughan), however, are similarly shallow. Whether this is sufficient to separate these forms at a high level from other upper Claiborne and Jackson species (e.g., P. creola (Palmer), P. apita (de Gregorio), etc.), which have deeper basal sinuses, is not clear. Some indication that it is not is provided by the basal sinus of Haustator perdita (Conrad), which varies from nearly as shallow to nearly as deep as the extremes shown by these other species. The evolutionary relationships among Palmerella femina (Stenzel), P. dutexata (Harris) and P. lisbonen- sis (Bowles) are uncertain. The very shallow to almost straight basal sinus of P. femina may call into question its relationship to any other Coastal Plain species. It is, however, very similar in its sculpture to dutexata and lisbonensis, all three species having pronounced B and C spirals and relatively reduced A spirals through- out ontogeny. It therefore seems likely that dutexata is the direct phyletic descendant of femina, with lis- bonensis branching off from this lineage. Palmerella hilli (Gardner, 1935) Plate 11, Figure 8 Turritella levicunea Harris. Plummer, 1933, p.815, pl.10, f.4,4a; not T. mortoni levicunea Harris, 1896. Turritella hilli Gardner, 1935, p.292, pl.25, f.4-6; Bowles, 1939, p.301; Stenzel and Turner, 1942, card 68; Palmer and Brann, 1966, p.989. Description. —Shell medium sized, of perhaps 15 to 20 whorls. Maximum observed whorl diameter 14.0 mm. Apical angle 20°; pleural angle usually slightly greater than apical angle. Protoconch and early apical whorls unknown. Smallest whorls examined showing three spirals of approximately equal strength, spaced equally over the whorl and separated by single smaller spirals. All these spirals becoming reduced in strength on following whorls except lowermost, which increases in strength and forms sharp, flaring basal carina. Ad- ditional, anterior rib expanding to equal this spiral, thickening and rounding the carina on last 4-5 whorls. Area between this spiral and the suture on latest whorls becoming slightly convex. Profile of early whorls more or less straight, of later whorls markedly concave as the carina increases in strength. Spirals finely beaded by the intersection of the growth lines with the fine spiral ribs. Lateral aspect of growth line trace orthoc- line, with upper and lower inflection points and a deep sinus with apex above whorl midpoint. Antispiral and basal sinuses moderately deep. Measurements. —See Table 19. Stratigraphic and Geographic Distribution.—TX: Table 19.—Measurements of Palmerella hilli (Gardner). Abbre- viations as in Table 10. MD MH WN USNM 337054 holotype 13.0 Do 8.5 USNM 12112 159 27.2 55) USNM 12112 13.4 25.8 > USNM 12112 PEZ SIND 9.0 USNM 12112 12.4 34.9 7.0 USNM 12112 THES) ST 8.5 | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 53 Kincaid Formation; lower Midwayan Stage, Lower Pa- leocene. Type Locality. —approx. 1 km above mouth of Dry Creek, Colorado River, Bastrop County, Texas (TX- BA-2b). Other Localities. —TX-BA-1c (B), ТХ-ВА-15 (B). Type Material. —holotype and 2 paratypes of T. hilli Gardner USNM 337054. Remarks/Discussion. — Without giving the size of the whorls, Gardner (1935) states that Early apical whorls sculptured with 2 strong simple spirals on the anterior half of the whorls, another of almost equal strength a little behind the middle and except on the very earliest whorls, a finer spiral directly in front of the posterior suture." Although the apical sculpture is unknown, the Sculpture described by Gardner on the earliest known whorls is consistent with a formula of C, B, A; such as shown by P. mortoni (Conrad). The adult whorl profile and sculpture of P. hilli are also similar to that of P. mortoni, although illi is substantially smaller. Govoni (1983) and Govoni and Hansen (in press) state that an undescribed form (*premortoni," P. mortoni Ssp. herein, see p.64), which they describe from the Brightseat Formation of Maryland (apparently equiv- alent to part of the Kincaid Formation in Texas), is very similar to P. hilli. They note that the Brightseat form "differs from [hilli] in details of relative strength and position ofthe primary spirals in the juvenile whorls and in having a more rounded and slightly less prom- inent basal carina." Palmerella levicunea (Harris, 1896) Plate 11, Figure 4 Turritella mortoni var. levicunea Harris, 1896, p.110, pl.11, f.9; Brann and Kent, 1960, p.928; not T. levicunea Plummer, 1933, p.815, pl.10, f.4,4a [= Palmerella ћи! (Gardner)]; not Т. mortoni Conrad, 1830. Turritella levicunea Harris. Bowles, 1939, p.301; Stenzel and Turner, 1942, card 74, £.1,9; Palmer and Brann, 1966, p.992-3; Toulmin, 1977, p.174, pl.8, f.9. Description. —Shell medium sized to large, of per- haps 10 to 15 whorls. Maximum observed whorl di- ameter 38.0 mm. Apical angle 22°; pleural angle 30°. Sutures generally inconspicuous. Protoconch and ear- liest teleoconch whorls unknown. Sculpture on earliest known whorls consisting of three moderately promi- nent spirals, one in approximately the middle of upper У of whorl, опе in middle of lower Y, and one very close to lower suture. Upper and middle spirals sep- arated from each other by at least two finer spirals; upper spiral from suture and middle from lower by Only one. Upper two spirals decreasing in strength on succeeding whorls as additional intercalary spirals ap- Table 20.—Measurements of Palmerella levicunea (Harris). Ab- breviations as in Table 10. MD MH WN PRI 43 holotype 18.0 36.0 7:5 USNM 15145 27.6 60.1 7.0 MCZIP 29277 20.0 37.6 8.5 pear, sculpture of later whorls consisting of numerous, more or less evenly spaced, very fine spirals. Lower- most spiral expanding to form edge of sharp basal ca- rina, in earlier whorls just above suture but in later whorls separated from suture by narrow, slightly con- cave to straight-sided area. Lateral aspect of growth line trace prosocline with indistinct upper and lower inflection points and moderately deep sinus whose apex is just above whorl middle. Antispiral and basal sinuses relatively shallow. Measurements. —See Table 20. Stratigraphic and Geographic Distribution. — AL: Clayton Formation, Porters Creek Formation (Mat- thews Landing Member), Naheola Formation; Mid- wayan Stage, Lower Paleocene. Type Locality. —Dale Branch, near Oak Hill, Wilcox County, Alabama (AL-WI-12). Other Localities. — AL-WI-9 (B), AL-WI-13 (MCZ), AL-WI-53 (T), AL-WI-54 (T), AL-WI-59 (ST). Type Material. —holotype of T. levicunea Harris PRI 43. Remarks/Discussion. — This species is very rare, the USNM collections containing only 14 specimens, and my own collections only a single specimen from Mat- thews Landing (MCZIP 29277). The species is, how- ever, highly distinctive in its earlier whorls for its smooth shell surface and large apical and pleural an- gles. In later ontogenetic stages as space begins to open up between the carina and the lower suture, it begins to resemble Palmerella mortoni (Conrad) more closely. In the absence of more knowledge of the apical sculp- ture, it is assigned only tentatively to Palmerella; the basal carina of this species could ally it to either Pal-: merella or Haustator. Palmerella lisbonensis (Bowles, 1939) Plate 3, Figures 1,2 Turritella nasuta Gabb. Palmer, 1937, p.200, in part, pl.25, f.3,9; Brann and Kent, 1960, p.928, in part, Nos. 2901, 2905, fide Palmer and Brann, 1966, p.993; not Gabb, 1860, p.385. Turritella lisbonensis Bowles, 1939, p.287, pl.31, £.1,3; Palmer and Brann, 1966, p.993; Toulmin, 1977, p.303, pl.51, £.7. Turritella dutexata lisbonensis Bowles. Stenzel and Turner, 1942, card 75. Table 21.—Measurements of Palmerella lisbonensis (Bowles). Ab- breviations as in Table 10. MD MH WN USNM 498009 holotype 8.5 59.5 155 Description. —Shell medium sized to large, of prob- ably 20 to 25 whorls. Maximum observed whorl di- ameter 14.2 mm. Apical angle 19°; pleural angle 7°. Protoconch and earliest teleoconch whorls unknown. Earliest whorls examined (dia. = 2.5 mm) bear four spiral ribs, two very prominent of approximately equal strength on lower half, two very weak of approximately equal strength on upper /. Additional weaker spirals appearing above, between and below the two lower primaries, which decrease in strength until whorl is covered by numerous, subequal spirals, approximately equally spaced. Whorl profile changing from basally bicarinate to slightly convex, with maximum width below whorl middle. Area between lowest strong spiral and lower suture remaining slightly concave. Sutures moderately incised. Lateral aspect of growth line trace prosocline, with weak upper and lower inflection points. Antispiral and basal sinuses shallow. Measurements. —See Table 21. Stratigraphic and Geographic Distribution. — AL: Lisbon Formation; GA, SC: McBean Formation?, Lis- bon Formation; TX, MS: Cook Mountain Formation; middle— upper Claibornian Stage, Middle Eocene. Type Locality.—approx. 1.5 km south of Lisbon Landing, Alabama River, Monroe County, Alabama (AL-MO-4d). Other Localities. — AL-CL-14(MCZ), AL-CL-16 (B), AL-CN-1 (B), AL-MO-1b (MCZ), MS-CL-1 (B), MS- CL-2 (B), MS-CL-7 (B), MS-CL-16 (B), MS-CL-17 (B), MS-CL-18 (B), MS-CL-19 (B), MS-NE-1 (B), MS-NE- 10 (B), SC-AI-1 (B), SC-LE-1 (B), SC-LE-2 (B), SC- OR-5 (B), SC-OR-6 (B), SC-OR-7 (B), SC-OR-10 (B), SC-OR-11 (B), GA-BU-3 (B), GA-BU-7 (B). Type Material. —holotype of T. lisbonensis Bowles USNM 498009, paratype USNM 497952. Remarks/Discussion. — As noted by Erickson (1982), Bowles states that /isbonensis is “confined to the Clai- borne of the eastern Gulf region," but then gives sev- eral Texas localities for its occurrence, apparently re- ferring to specimens referred to Palmerella dutexata (Harris). This confusion highlights the similarity of lisbonensis to P. femina (Stenzel) and P. dutexata (Har- ris) which, as discussed above (p.57), are probably 5 See Footnote 3, p.46. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 themselves ends of an intergrading series. Palmerella lisbonensis shares with these forms the prominence of a pair of carinae on the lower portion of early as well as late whorls, but it is distinct in attaining a greater size and much more elongate overall shape. In early whorls it shows a relatively high apical angle, similar to femina, but soon changes to a more columnar, cy- lindrical form. On the basis of these morphological and stratigraph- ic relationships, it seems reasonable to suggest that Р. dutexata (Harris) of the Cook Mountain, Upper Lisbon and Gosport Formations is a direct phyletic descen- dant of P. femina (Stenzel) from the Weches Forma- tion, and that P. lisbonensis is a product of a clado- genetic event from this lineage, probably during de- position of the Cook Mountain/Upper Lisbon for- mations. As mentioned above, /isbonensis may have been close to the ancestry of P. apita (de Gregorio) from the Gosport and P. arenicola (Conrad) from the Moodys Branch. Palmerella mortoni mortoni (Conrad, 1830) Plate 1, Figures 4-6, Plate 2, Figures 6-8 Turritella mortoni Conrad, 1830, p.213-215, 221, pl.10, f.2; 1834, p.4 in part; 1835a, p.40 in part, not pl.15, Ғ11 [= Haustator carinata (1.Lea)]; 1846, p.219; 1865a, p.32; 1866, p.11; Tuomey, 1858, p.267, not p.270-272; not Aldrich, 18862, p.9,13,46,55,58- 60 [= Palmerella mortoni postmortoni Harris]; not Smith and Johnson, 1887, p.59 in part [= Palmerella alabamiensis Whitfield]; de Gregorio, 1890, p.122 in part, pl.11, f.7, copy Conrad, 1830, not pl.11, f.3-6,9 [= Haustator carinata (1.Lea)]; Cossmann, 1893, p.29; Harris, 1894a, p.302-304, f.3; 1896, p.110; not 1899a, p.74, pl.10, #3,4 [= P. m. postmortoni]; not 1899b, p.299,308, р1.52, £9 [=Turritella sp., fide Palmer and Brann, 1966, p.994], pl.55, £4 [= P. m. postmortoni]; Clark, 1895, p.4; 1896, p.69, pl.13, f.1a- е; Clark and Martin, 1901, p.147, pl.26, f.1—4; not Veatch, 1906, p.14, f.5,5a [= P. m. postmortoni]; Clark and Miller, 1912, p.92- 100, 102,119,120; Guillaume, 1924, p.289,290; 1926, p.426; not Cooke, 1926, p.259,261,264, pl.94, f.2; not Semmes, 1929, f.59- 2 [= P. m. postmortoni]; Palmer, 1937, p.195, р1.23, f.6,11, pl.24, f.14 [not 13 as in text]; Bowles, 1939, p.293, pl.33, f.14; Stenzel and Turner, 1942, card 83; Shimer and Shrock, 1944, p.493, pl.201, f.14-17, copy Clark and Martin, 1901; Brann and Kent, 1960, p.927 in part, nos. 2880,2883,2892,3904 only, fide Palmer and Brann, 1966, p.995; Vokes, 1961, p.49, pl.10, f.1; Palmer and Brann, 1966, p.994—5. Turritella mortoni mediavia Bowles, 1939 in part, p.294, pl.33, Ғ3; not f.5. Description. —Shell large to very large, of 20 to 25 whorls. Maximum observed whorl diameter 40 mm. Apical angle 22°; apical and pleural angles approxi- mately equal. Sutures moderately deeply incised. Pro- toconch erect, homeostrophic, turbinate, of approxi- mately 1.7-2.0 smooth, rounded whorls; P1 small. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 55 Apical sculpture formula С, B, A. Primary spirals C and B appearing approximately simultaneously as broad, slightly angular prominence just below middle of whorl, and quickly diverging and becoming more acute, B becoming stronger than C. Primary A ap- pearing within 0.5-0.75 whorl, remaining weaker than C for one to two whorls after which it is of approxi- mately equal strength. Secondaries and finer spirals beginning at about whorls 8-10. Primary Band one or more secondaries remaining prominent on upper slope Of carina, but more adapical spirals becoming obsolete. Whorl profile changing shortly after appearance of nu- merous secondary spirals, from medially to basally car- inate, C becoming relatively more prominent and low- er 1⁄4 of whorl expanding outward. Area between C spiral and suture expanding and changing from rela- tively straight-sided to variably convex in profile. Whorl profile above carina becoming increasingly concave. Lateral aspect of growth line trace prosocline, with pronounced upper and lower inflection points and rel- atively deep sinus whose apex just above whorl middle. Lower inflection point usually high on lower part of whorl, causing spiral sinus to be relatively narrow and basal sinus relatively wide although both moderately deep. Measurements. —See Table 22. Stratigraphic and Geographic Distribution. — MD, VA: Aquia Formation (Piscataway and Paspotanza Members); SC: Black Mingo Group, Williamsburg Formation; Upper Paleocene. Type Locality. —probably Piscataway Creek, Prince Georges County, Maryland (MD-PG-5). Other Localities. -MD-AA-5 (В), MD-CH-7 (B), MD-PG-3(B), MD-PG-10 (B), VA-ST-4 (B), VA-ST-5 (B), VA-ST-6 (B), VA-ST-7 (B), VA-HA-2 (MCZ), SC- SU-2 (B), SC-BE-3 (VC), SC-CL-2 (B). Type Material. —holotype? and 2 paratypes of 7. mortoni Conrad ANSP 15508. Remarks/Discussion. —‘‘Turritella mortoni Con- rad" and its close relatives are of special importance in both the evolutionary history of their clade in the Coastal Plain Paleogene, and the history of its study Table 22. — Measurements of Palmerella mortoni mortoni (Con- rad). Abbreviations as in Table 10. MD MH WN ANSP 15508 holotype ? 2009 59.9 9.5 ANSP 15508 paratype 22:7 50.6 8.5 USNM 497990 26.5 98.0 14.0 MCZIP 29307 31.0 110.0 115 MCZIP 29099 37.9 104.5 9.5 over more than a century and a half. Turritella mortoni was one of the first species to be described from the “Eocene” of North America (Conrad, 1830), and since that time it has been a commonly used index fossils (e.g., Shimer and Shrock, 1944; Toulmin, 1977), and is one of the most characteristic and widely recognized macrofossil taxa of the lower Tertiary. The systematic status of“ T. mortoni”, however, has remained poorly understood. Many of the numerous references to it in the literature, as Bowles (1939) has noted, refer to other species, and the name “7. mortoni" has commonly been used “more in the sense of the group of basally carinate Paleocene and Lower Eocene turritellas than as a distinct specific entity" (1939, p. 293). Four very similar named forms comprise what can be called “Turritella mortoni sensu lato.” These аге T. mortoni s.s. Conrad, 1830 from the Sabinian-aged Aquia Formation of Maryland and Virginia and Sa- binian-aged Williamsburg Formation (Black Mingo Group) of South Carolina; 7. postmortoni Harris, 1894a from the Sabinian-aged Nanafalia and Tuscahoma Formations (Wilcox Group) of the Gulf Coastal Plain; T. mediavia Bowles, 1939 from the Midwayan-aged Clayton Formation of the Gulf Coastal Plain; and 7. *premortoni" (Govoni, 1983; Govoni and Hansen, in press) from the Midwayan-aged Brightseat Formation of Maryland. (As discussed above, 7. hilli Gardner from the Midwayan-aged Kincaid Formation of Texas may be very close to 7. “ premortoni" and may rep- resent a fifth member of this group.) (See Text-figure 15 for stratigraphic relationships.) All four of these taxa share the following features: 1) a moderate and round- ed to pronounced and sharp-edged carina, in which the C spiral is most prominent, 2) moderate apical angles, approximately equal to pleural angles, 3) me- dium to large size. Morphologic relationships among these forms have usually been approached only in very general and qual- itative terms. Clark and Martin (1901, p.147) note that‘ T. mortoni” from Maryland and Virginia “shows very great variation in the form and decoration of the whorls, and if it were not for the great number of in- termediate types one might readily establish several independent species." Andrews (1971, 1974) has at- tempted a preliminary morphometric analysis of this variation, but uses only 37 specimens and does not make reference to taxa from elsewhere on the Coastal Plain. He is thus unable to draw any systematic con- clusions. I have attempted to resolve the morphological re- lationships among at least some of these forms by mul- tivariate morphometric analysis. Details of the meth- ods and specimens used are presented in Appendix 1. 56 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 23.—Results of factor analysis of specimens of the Recent species Turritella terebra (Linnaeus). Variance explained by first five factor axes. cumulative proportion Factor of variance 0.4018 0.5645 0.6396 0.7077 0.1333 л > 6 — For mortoni and its close allies, I have subjected large samples of specimens from Virginia and Maryland (mortoni s.s.) and Alabama (usually assigned to post- mortoni Harris) to factor analysis. These results can be compared to a similar analysis of a Recent species, the type species of the genus Turritella Lamarck, Tur- ritella terebra (Linné). Text-figure 17 shows the results for T. terebra. A total of 78 variables on each of 86 specimens were analyzed. The first three factors together explain al- most 64% ofthe total variance (Table 23). In the factor Table 24.— Results of factor analysis of specimens of the Recent species Turritella terebra (Linnaeus). Rotated factor loadings of each variable on each of the first three factor axes. Variables as indicated in Text-figure 3. Variable factor 1 factor 2 factor 3 Variable factor 1 | factor 2 factor 3 УН! – 0.438 0.143 – 0.090 WWII 0.419 0.125 0.462 SWI —0.440 0.142 —0.108 IN11 0.275 0.032 0.448 WWI —0.439 0.142 —0.108 WH12 0.668 0.083 0.252 INI —0.439 0.141 220-013 SWI2 0.692 0.058 0.250 WH2 —0.649 0.057 —0.060 Ww12 0.704 0.103 0.212 SW2 —0.646 0.059 —0.042 IN12 0.499 0.084 0.205 WW2 —0.650 0.059 —0.049 WH13 0.782 0.246 0.009 IN2 —0.639 0.049 —0.051 SWI13 0.786 0.210 0.016 WH3 0.318 —0.099 0.128 WW13 0.786 0.241 0.011 SW3 —0.810 —0.095 0.138 IN13 Qu 0.249 0.023 WW3 —0.806 03101 0.120 WH14 0.829 0.327 —0.143 IN3 2:0 789 —0.096 0.126 5%14 0.828 0.297 = 0439 WH4 —0.830 0:270 0.196 wwil4 0.829 0.326 —0.129 SWA —0.818 —0.261 0.220 IN14 0.822 0.340 EO} WWA —0.806 — 0.268 0.194 WHI5 0.795 0.393 = 04198 IN4 —0.746 —0.189 0.242 SWIS 0.796 0.363 —0.185 WHS —0.607 —0.514 0.363 WWIS 0.795 0.388 —0.185 SWS —0.540 0520 0.367 INIS 0.780 0.404 = 021163 WWS —0.530 06021 0.338 WHI6 0.716 0.463 = 0292 INS 220995 —0.400 0.352 SW16 02/22, 0.432 —0.262 WH6 —0.298 —0.613 0.564 WW16 0722 0.453 —0.263 SW6 =0. F90 220:617 0.577 IN16 0.617 0.429 2201229 wwe (РТУ —0.611 0.519 WH17 0.479 0.612 —0.354 IN6 —0201 —0.452 0.510 SW17 0.482 0.584 2201370 WH7 2-001273 —0.650 0.590 WWI7 0.483 0.600 0309 SW7 0:156 —0.634 0.603 IN17 0.427 0.660 —0.345 WW7 —0.079 —0.621 0.495 WH18 0.249 0.714 —0.430 IN7 = 0.152 – 0.314 0.496 SW18 0.253 0.685 —0.451 WH8 —0.160 —0.296 0.863 WWI8 0.249 0:717 —0.429 SW8 =0070 —0.269 0.873 IN18 0.217 0.794 220/525 WWS —0.009 —0.260 0.761 WHI9 0.073 ` 0.869 —0.007 IN8 —0.164 SOLOS 0.607 SW19 0.077 0.852 —0.016 WH9 =0:012 220221 0.876 WW19 0.074 0.868 —0.003 SW9 0.052 —0.197 0.884 IN19 0.066 0.894 0.042 WWO 0.133 – 0.145 0.737 WH20 0.015 0.807 0.138 IN9 0079 —0.043 0.578 SW20 0.015 0.807 0.137 WHIO 0.032 —0.065 0.911 WW20 0.016 0.808 0.140 SW10 0.074 270/059 0.916 WH WWI0 0.158 —0.014 0.767 SW IN10 —0.058 0.044 0:57] WW WHIl —0.103 0.116 0.252 IN SWI1I 0.346 0.094 0.619 | | І PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON $7 l—factor 2 plot (Text-fig. 17A), most (84%) of the Specimens analyzed cluster around the factor 1 axis. The 14 specimens that do not do so fall along a neg- atively sloping line in the first quadrant, scoring high on factor 2. Ten of these 14 specimens also fall away from the rest in the factor 2— factor 3 plot (Text-fig. 17B), scoring negatively on factor 3. The loadings in the varimax-rotated factor matrix (Table 24) suggest that these 14 specimens differ primarily in the later whorls (17—20). These 14 specimens are not randomly drawn from all samples analyzed; 11 of the 14 come from two Specimen lots from the Philippines (ТЕКЕВКІ, TEREBR18; Appendix 1), suggesting the possibility that the variability noted here may represent differ- entiation at the population level. This possibility is supported by the distribution of 7. terebra in the west- ern Pacific. It is found today in considerable numbers in Taiwan (Kuroda, 1941), the south-central coast of the Chinese mainland (Chin, 1941; Chung, 1981), Hong Kong (Wu and Richards, 1981), the Philippines (Faus- tino, 1928), New Guinea (Hinton, 1972), Java (Rob- erts et al., 1982) and northern Australia (Garrard, 1972; Wilson, 1993). This range is larger than that of most Recent turritellid species, raising the possibility that T. terebra may include a number of diverging popu- lations. Text-figure 18 shows the results for mortoni/post- mortoni. A total of 90 variables on each of 354 spec- imens were analyzed. The first three factors together explain 57.296 of the total variance (Table 25). The varimax-rotated factor matrix (Table 26) indicates that the first factor is dominated by variables on whorls 4,5, and 6, whereas the second factor is dominated by variables on later whorls (7—9) and the third factor by variables on earlier whorls (1—3). Alabama specimens Score higher on all three axes than do Maryland and Virginia specimens, indicating differences throughout ontogeny. It is clear from the plots in Text-figure 18 that specimens from Maryland and Virginia are, as a group, distinct from specimens from Alabama, but not more distinct than any of the populations of Turritella terebra analyzed (Text-figure 17). It would have been very desirable to include other forms and samples in this analysis, especially speci- mens assigned to mortoni from the Sabinian-aged Black Mingo Group of South Carolina (e.g., Van Nieuwen- huise and Colquhoun, 1982), as well as specimens of T. mediavia Bowles, 1939 and T. “premortoni” Go- voni, 1983, but too few well-preserved specimens of any of these were available for quantitative analysis. These limitations of material produce substantial lim- itations of interpretation in attempts at a comprehen- Sive understanding of evolution within “mortoni sensu lato": 1) Without morphometric data on Sabinian forms from South Carolina, it is difficult to determine wheth- er the differences shown in Text-figure 18 between specimens from Virginia-Maryland and Alabama are clinal or more characteristic of discrete subspecific dif- ferentiation. 2) Without morphometric data on Mid- wayan forms from Maryland (“premortoni” Govoni, 1983; Govoni and Hansen, in press) and the Gulf coast (mediavia Bowles, 1939), it is difficult to decide how many discrete taxa should be recognized during this interval and, more importantly, how Midwayan forms evolved into Sabinian forms. Qualitative observations are, however, possible. Bowles (1939, p. 295) states that mediavia differs from mortoni s.s. and postmortoni of Harris “in the persis- tence of the fairly strong secondary carina that makes the profile ofthe whorls more rounded," but specimens with similar sculpture can be found among samples of mortoni from Virginia and Maryland (e.g., Plate 1, figure 4, Plate 2, figure 6). Bowles (1939, p. 295) notes that some specimens from the Black Mingo Group of South Carolina are “identical in every way with the holotype of Turritella mortoni mediavia,” yet they also agree closely with specimens of mortoni s.s. from Vir- ginia. Among the Midwayan forms, virtually all spec- imens referred to mediavia Bowles are considerably larger and more robust (and thus more similar to both mortoni s.s. and postmortoni) than all known speci- mens of “premortoni” Govoni. (In size and adult sculpture, 7. “premortoni” in fact most closely resem- bles 7. hilli Gardner.) Based on the multivariate analysis presented above, and comparison with the Recent species Turritella ter- ebra, it is reasonable to conclude that the patterns of variation showed by the Sabinian-aged specimens from Maryland-Virginia and Alabama are consistent with intra-specific variation, and that Gulf and Atlantic populations can be considered as representing two dis- tinct subspecies within mortoni. Carolinian specimens of the same age appear intermediate but this cannot be documented quantitatively. There are at least four possible evolutionary rela- tionships among these taxa. In the first (Text-figure 19A), only one of the Midwayan-age forms (mediavia or “premortoni’’) is ancestral to both postmortoni and mortoni s.s. In favor of *premortoni" being this single ancestral form is the close similarity of its protoconch and early teleoconch whorls to those of mortoni s.s. (Govoni, 1983, p. 103; see Plate 1, figs. 2,6). In favor of mediavia as representative of this single ancestral form is the greater (albeit qualitative) similarity in adult size and sculpture of the Sabinian forms to mediavia than to “premortoni”. Against this interpretation is the lack of apparent mediavia descendants where they might 58 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 A iani О A ed 3 ir a a N 27 5 о © D ia = o CH Е Cu n - О ГІР! H- | | COLO) Ө : g | 5—4 2.5 2 er er ae mo 1 en 2 - - - o - -0. п Оп г o - об әт. gp O AE o 24 Factor 1 B p o o о ге | | | | [ale == -2 -1 1 2 Macy 4 5 о 5 5 o ІР Imi -1 o+ o o AD tE O SARI B E a -2.5 + cg O ak Factor 2 Text-figure 17.—Results of factor analysis of specimens of Turritella terebra. A. first and second factor axes. B. Second and third factor axes. See Appendix 1 for specimens used and details of analysis. be expected if mediavia was directly ancestral to post- mortoni (i.e., in post-Clayton fossiliferous sediments ofthe Gulf coast, such as the Matthews Landing Mem- ber of the Porters Creek Formation, and the Naheola Formation). The second scenario (Text-figure 19B) suggests that Atlantic and Gulf Coastal Plain lineages evolved more or less separately during the Paleocene; “premortoni” is ancestral to mortoni s.s., and mediavia to postmor- toni. Such separation is supported by the differences within both Midwayan and Sabinian forms. Taken as a whole, mortoni s.s. from the Aquia of Maryland and PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 59 + xO УХ уу N | | cst OIE ~ x x е -4 -3 хах, х «i 5 Ж XK 2- хх о Au x % х х “x Sa sica Factor 1 X Virginia L] Alabama B ds a Ber ЕНЕ о 5 uds o ¡El 4 + о 8 3 + x Ы nn в 2 a О ims | | | дет e S dibus | 8 e Macs 7” 2 ma go te : 2 4 6 Factor 2 Text-figure 18.—Results of factor analysis of specimens of Palmerella mortoni sensu lato. A. first and second factor axes. B. Second and third factor axes. x represents specimens from Virginia (mortoni 5.5.); O represents specimens from Alabama (usually referred to postmortoni Harris). See Appendix 1 for specimens used and details of analysis. Virginia shows differences in the ontogenetic devel- expansion stops before largest size is reached, whereas opment of the basal carina compared to postmortoni in mortoni s.s. it continues throughout growth; the from the Alabama Wilcox. The expansion of the carina greater degree of carination in postmortoni is a result is allometric in both forms, but in postmortoni this ofa higher allometric scaling exponent (Allmon, 1994). 60 Table 25.—Results of factor analysis of specimens of Palmerella PALAEONTOGRAPHICA AMERICANA, NUMBER 59 mortoni (Conrad). Variance explained by first five factor axes. Factor cumulative proportion of variance AUNI 0.3505 0.4828 0.5720 0.6376 0.6953 Govoni's “premortoni” is distinct from mediavia, dif- fering in size and degree of sculpture, but is a good candidate for ancestry of Aquia mortoni s.s. not only because of its geographic position and apical form, but also its moderate, rounded carina which is similar to that of the lowest mortoni in the Aquia. Bowles’ me- diavia is similar to both postmortoni and some late Aquia mortoni in its pronounced carina and large size. Under this scenario the four forms should probably be considered separate species, since temporally coexist- Table 26.—Results of factor analysis of specimens of Palmerella mortoni (Conrad). Rotated factor loadings of each variable on each of the first three factor axes. Variables as indicated in Text-figure 3. Variable factor 1 factor 2 factor 3 Variable factor 1 factor 2 factor 3 УНІ —0.047 0.064 0.747 MW7 0.602 0.574 0.054 CHI-1 —0.048 0.068 0.740 CW7 0.643 > 092 0.070 CH2-1 —0.044 0.055 0.742 CAN7 0.567 0.546 —0.003 SWI —0.046 0.060 0.744 WH8 = 0150 0.346 0.111 MWI —0.046 0.059 0.749 CH1-8 0.251 0.699 0.002 CWI —0.046 0.061 0.751 CH2-8 0.400 0.482 0.107 CANI —0.042 0.057 0.744 SW8 0.433 0.612 —0.014 WH2 0.132 0.017 0.840 MWS8 0.459 0.633 0.020 СН1-2 0,125 0.022 0.851 CW8 0.539 0.631 0.054 CH2-2 0.144 0.002 0.769 CAN8 0.331 0.632 —0.047 sw2 0.127 0.018 0.829 WH9 —0.248 07972, 0.071 MW2 0.132 0.016 0.846 СН1-9 -0.341 0.542 -0.118 CW2 0.135 0.015 0.840 CH2-9 0.053 0.523 —0.044 CAN2 0.140 0.010 0.832 SW9 —0.147 0.504 —0.160 WH3 0.463 —0.058 0.622 MW9 = 0/158 0.538 —0.142 СН1-3 0.497 —0:095 0.613 CW9 20072 0.543 =0.125 CH2-3 0.493 ОШО 0.561 CAN9 0207 0.447 —0.155 SW3 0.489 —0.094 0.609 WHI10 —0.503 0.229 0.057 MW3 0.501 —0.096 0.604 СН1-10 —0.803 0.060 —0.053 CW3 0.502 20.097 0.604 CH2-10 —0.647 0.177 —0.021 CAN3 0.516 = АИ 0.568 SW10 —0.739 0.044 —0.062 WH4 0.743 —0.046 0.334 MWI0 —0.758 0.056 —0.059 CH1-4 02132 —0.039 0.329 Cw10 2:0273:7 0.068 20055 CH2-4 0.708 —0.062 0.318 CANIO —0.765 0.007 —0.076 SWA 0.722 —0.033 0.343 WH11 OO 0.107 0.067 MWA 0737 --0:056 0.344 СН1-11 —0.804 —0.161 —0.030 CWA 0.734 —0.034 0.344 CH2-11 20:739 0015 20:911 САМА 0.741 — 0.043 0.314 Swill 0:765 —0.191 —0.044 WHS5 0.841 (02157 0.146 MW11 —0.783 —0.167 —0.036 СН1-5 0.820 0.163 0.137 CWII О —0.164 —0.037 CH2-5 0.793 0.118 0.151 CANII 290/7270 “(202 —0.048 SW5 0.802 0.179 0.127 WHI12 2:019 30 0.203 0.106 MWS 0.830 0.176 0.119 СН1-12 -0.578 -0.500 —0.013 CWS. 0.827 0.178 0.134 CH2-12 —0.561 —0.480 = 0012 CANS 0.836 0.161 0.097 Sw12 = (0.572 – 0.508 – 0.014 WH6 0.767 0.392 0.064 MW12 05979 —0.504 —0.014 CH1-6 0.714 0.393 0.088 CW12 =0.573 —0.514 —0.014 CH2-6 0.609 0.363 0.074 CAN12 2015/75 —0.487 DIS SW6 0.714 0.378 0.049 WH13 065 0720 —0.047 MW6 0.736 0.397 0.054 СН1-13 ONIS 057271 —0.047 CW6 0.754 0.388 0.086 CH2-13 OSO ЕДІ) —0.047 CAN6 0.746 0.358 0.014 SW13 —0.134 exits) —0.047 WH7 —0.090 0.327 0.116 MW13 0.135 22017115 —0.047 СН1-7 0.534 0.599 0.036 CWI13 089 s0712 —0.047 CH2-7 0.492 0.438 0.154 CAN13 20:192 —0.724 —0.047 SW7 0.563 0.553 0.030 | | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 61 ing forms are morphologically distinct and the two lineages are presumed to have evolved independently Over geological time. Finally, Midwayan forms may represents a single, variable and geographically widespread species that evolved, in toto, into another variable and widespread Species in the Sabinian (Text-figure 19C). In support of this hypothesis is the highly variable morphology within mortoni from the Aquia. Andrews (1971) sug- gests that a less carinate form evolved into a more carinate form within the Aquia section. Text-figure 18 indicates that postmortoni and mortoni s.s. are both variable, but along different multivariate axes. Wheth- er *premortoni" and mediavia could have been соп- Specific is not clear, given their size and sculpture dif- ferences, but it is interesting to note that mediavia Occupies a geographic area in-between those of the morphologically similar and approximately synchro- nous hilli in Texas and “premortoni” in Maryland. А possible variant on this scenario is similar to one sug- gested that has been discussed for human evolution (see Van Valen, 1966). Different populations ofa single Species could have crossed a morphological threshold to a new morphology independently. The descendant populations would still belong to a single and more or less genetically continuous species, but its populations had not all evolved simultaneously or with constant levels of genetic interchange. Based on all information currently available, I ten- tatively favor the last of these hypotheses (Text-figure 190). Data are sufficient to show that mortoni s.s. from Maryland-Virginia is distinct from postmortoni from Alabama, but not substantially more so than different living populations of Turritella terebra; specimens from geographically intermediate areas may (or may not) link the two morphologically. Data are not sufficient to indicate whether “premortoni” and mediavia (and hilli ?) can be united into a single, variable species or which Midwayan form most closely resembles (and gave rise to) which Sabinian form. It is possible that larger samples of the forms that were not analyzed quantitatively here can be assembled in the future. Pending such analyses, I choose to include all four forms within a single, variable species mortoni, and to recognize them as geographic/chronologic subspecies. Palmerella mortoni mediavia (Bowles, 1939) Plate 2, Figures 1,9 ? Turritella mortoni Conrad var. Harris, 18945, p.48, pl.3, f.5; Brann and Kent, 1960, p.928. Turritella mortoni Conrad ? Harris, 1894b, p.47,48; 1896, p.16,17; McCallie, 1908, p.335 [not p.235 as in Bowles, 1939]; Veatch and Stephenson, 1911, p.219,220; Ste- phenson and Veatch, 1915, p.69; Stephenson and Crider, 1916, p.48, 49, 51-53; Brantly, 1920, p.144,146 [only]; Cooke, 1936, p.50; not Conrad, 1830; Toulmin, 1977, pl. 8, Е8. ? Turritella saffordi Gabb. Harris, 1896, p.110, in part, pl.11, Ғ8, [Т. mortoni var. on plate caption]; not Gabb, 1860, p.392. Turritella mortoni mediavia Bowles, 1939, p.294 in part, pl.33, £.5; not pl.33, £3 [= ? Palmerella mortoni s.s. Conrad]; Stenzel and Turner, 1942, card 79; Palmer and Brann, 1966, p.995-996. Description. — Original description: *Spire abruptly tapering for the genus. Apical whorls sharply angulated anteriorly, concave in profile; adult whorls more rounded, almost convex in profile, sharply constricted at the sutures, which are linear but distinctly impressed between the apical whorls, gradually becoming deeper and wider between the later whorls. Sculpture of apical whorls consisting of a strong basal carina with a less prominent revolving cord near the posterior suture; both the carina and the posterior cord persistent on the adult whorls, the posterior giving the whorls a more rounded appearance than is general in the group. Sec- ondary sculpture consisting of numerous very fine re- volving lines, beaded on some of the early whorls by the intersection of the growth lines. Incremental striae distinct and sharply flexed medially, curving out again and then slightly reflexed at the anterior suture. Ap- erture unknown." (1939, p.294) Protoconch and earliest teleoconch whorls un- known. Measurements. —See Table 27. Stratigraphic and Geographic Distribution. — AL, GA, MS, TN, AR: Clayton Formation; lower Midwayan Stage, Lower Paleocene. Type Locality. — Prairie Creek, Wilcox County, AI- abama (AL-WI-9). Other Localities. — AL-HE-2 (TLM), AL-HE-3 (VS), GA-CL-3 (B), GA-CL-6 (T), GA-SL-3 (VS), GA-MA-1 (B), TN-HA-12 (US), AR-PU-4 (SC), AR-WH-1 (HA), AR-WH-2 (SC), AR-LO-1 (HA). Type Material. —holotype of T. mortoni mediavia Bowles USNM 495146, paratype USNM 495147. Remarks/Discusssion. —See above, under P. mor- toni mortoni. Palmerella mortoni postmortoni (Harris, 18942) Plate 2, Figures 2-5 Turritella mortoni Conrad. Aldrich, 18862, in part, p.13,55,58; not p.9,46,59,60; Harris, 1899a, р.74,р1.10, f.4; 1899b, p.308, in part, pl.55, £.4, not pl.52, Ғ9 [= Turritella sp., fide Palmer and Brann, 1966]; Veatch, 1906, pl.14, f.5,5a; Guillaume,1924, p.289 in part [not Claiborne as indicated]; 1926, p.425; Cooke, 1926, p.259,261,264, pl.94,f.2; Semmes, 1929, Ғ59-2; not Conrad, 1830. Turritella mortoni postmortoni Harris, 1894a, p.302-304, in part, f.1, not £2 [= Palmerella mortoni s.s.]; 1899a, p.75, pl.10, Ғ3; Clark and Martin, 1901, р.147-148 in part, not pl.26, Ғ5 [=P. mortoni s.s.]; not Grabau and Shimer, 1909, f.1064d [= P. mortoni s.s.]; Bowles, 1939, p.298, pl.33, f.4; Palmer, 1937, p.188, pl.23, f.8-10; Stenzel and Turner, 1942, card 95; LeBlanc in Barry and LeBlanc, 1942, pl.104, in part, pl.13, f.11; Shimer and Shrock, о ч Ы wod pow EI 3 | Жос њи Т d N E сч zi БЕНЕН ae he NT mm ` г Feuer / шина i \ 2 ч ty 2 Wd} I дош Y TEE 9 i ! Џ ! ) 5 i | 1 ' | 2 E- — – – - – – – – - ин poU б І d ЈОШ | udi i Е | ! ! М 552 2 i E Oo oO OOO GEO O | ШЕШШ, | | 5 : wod pau \ wod ! | "Deu 1 © I | б | ! A / ! E A Балы | Жы " | < шоа рәш | аи i ' A, | юха \ 1 с- ETEND ы — 7. | 119 SEE | ~ = | __- ity к= U « са 62 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 63 Table 27.— Measurements of Palmerella mortoni mediavia (Bowles). Abbreviations as in Table 10. Table 28.— Measurements of Palmerella mortoni postmortoni (Harris). Abbreviations as in Table 10. MD MH WN MD MH WN USNM 495146 holotype 19.0 41.0 7.5 USNM 497992 hypotype 23.0 65.0 10.5 MCZIP 29308 26.6 743 10.5 1944, p.498 in part, not pl.201, f.17 [=P. mortoni s.s.]; Brann and Kent, 1960, p.928, in part, not No. 3903 [= P. mortonis.s.]; Palmer and Brann, 1966, p.996; Toulmin, 1977, p.232, pl.29, f.9. Turritella humerosa Conrad. Trowbridge, 1932, pl.31, Ё6,7, copies Veatch, 1906, after Harris, 1899a [PRI 270, 271, not USNM 11754 as in Trowbridge]; not Conrad, 1835b. Turritella postmortoni Harris. Toulmin, 1969, pl. 1, f.14. Turritella sp. Toulmin, 1977, pl.29, f.12,13. Description. —Shell large, of 15 to 20 whorls. Max- imum observed whorl diameter 27.0 mm. Apical angle varies from 19°; apical and pleural angles approxi- mately equal. Protoconch and earliest teleoconch whorls unknown. Sutures not deeply incised. Spiral sculpture on earliest whorls observed (dia. = 0.75 mm) consisting of three moderately prominent spirals, one in middle of whorl and one close to each suture, middle spiral being strongest. In succeeding whorls, upper and lower Spirals moving toward center, space between upper and middle spirals becoming greater than that between low- er and middle. Within four to five whorls, secondary spirals appearing between upper and middle primaries, between upper primary and suture, and immediately associated with lower suture. Further intercalaries con- tinuing to appear on succeeding whorls. Teleoconch whorls wider than high. Profile of earliest known whorls generally straight-sided, angulated in middle by prom- inent middle primary spiral. On succeeding whorls, lower primary spiral surpassing middle spiral in strength; whorl profile becoming basally carinate; area between lower primary spiral and spiral at suture be- coming slightly concave in profile. This area remaining approximately same size on succeeding whorls, while area beneath lowest spiral expands, the point of contact of following whorl shifting relatively inward with ca- rina coming to overhang suture with a surface origi- nally roughly horizontal, becoming angled downward and slightly convex outward in profile with carina moving relatively upward on the whorl. On late whorls profile markedly concave above the carina and slightly convex below. Carina varying in profundity from acute angulation in lower half of whorl to flaring, adapically turned flange several mm wide, often undulating in its lateral trace. Outer edge of carina almost always made of single spiral rib, but single weaker spiral just below edge usually visible. Lateral aspect of growth line trace markedly prosocline, with well developed upper and lower inflection points and a moderately deep sinus, apex of which is just above whorl middle. Lower in- flection point usually relatively high on lower Y. of whorl, causing spiral sinus to be relatively narrow and basal sinus to be relatively shallow. Measurements. —See Table 28. Stratigraphic and Geographic Distribution. — AL,GA: Nanafalia Formation (Ostrea thirsae beds, Grampian Hills Member), Tuscahoma Formation (Bells Landing Member, Greggs Landing Member); TX,LA: Pendle- ton Formation, Marthaville Formation; Sabinian Stage, Upper Paleocene. Type Locality.—Bells Landing, Alabama River, Monroe County, Alabama (AL-MO-3) Other Localities. — AL-WI-1 (B), AL-WI-18b (B), AL- WI-18c (C), AL-WI-55 (B), AL-WI-57 (T), AL-WI-58 (T), AL-MA-1 (B), AL-PI-1 (B), AL-PI-6 (T), AL-CH-6 (B), AL-CH-11 (T), AL-CH-12 (T), AL-DA-1 (МС2), AL-DA-4 (T), AL-MO-5 (B), AL-HE-4 (T), TX-SA-5 (B), TX-SA-5C (L), TX-SA-19 (L), LA-NA-5 (L), LA- NA-12 (L), LA-NA-15 (L), LA-NA-16 (L), LA-NA- 17 (L), LA-SA-20 (WW), LA-SA-21 (WW), LA-SA- 22 (WW), GA-HE-1 (MCZ), GA-CL-2 (MCZ), GA- CL-7 (ANS). Type Material. —lectotype of Т. mortoni postmor- toni Harris lost (fide Palmer and Brann, 1966, p.996), hypotype (Bowles, 1939) USNM 497992. Remarks/Discussion. — Harris’ postmortoni din qualitatively from mortoni s.s. principally in being smaller, and in frequently showing a sharper and more pronounced basal carina, but overlaps considerably in — Text-figure 19.— Four possible evolutionary scenarios for the forms most closely related to Palmerella mortoni (Conrad). mor = Palmerella mortoni s.s., pom = Palmerella mortoni postmortoni, prm = Palmerella mortoni ssp. (“premortoni”), med = Palmerella mortoni mediavia, hil = Palmerella hilli Gardner. A1, A2. Only one phyletic link between Sabinian and Midwayan forms, either between the Gulf coast forms mediavia and postmortoni, or the Atlantic coast forms mortoni s.s. and “premortoni.” B. Atlantic and Gulf lineages are separate from each Other throughout Midwayan and Sabinian times. C. Gulf and Atlantic forms continually interbreeding throughout Paleocene and earliest Eocene times, forming one broad and highly variable phyletic lineage. This is the hypothesis favored here. 64 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 general shell form. The results of the factor analysis (Text-figure 18) indicate that there are whorl shape differences between the Alabama and Virginia/Mary- land specimens at all ontogenetic stages. An interesting feature showed by several individuals of postmortoni is a doubling of the carina by the ex- pansion of a secondary spiral just above the C spiral (see Plate 2, Figures 2,3,5). Toulmin (1977, pl.29, f.12,13) suggested that these forms represent a distinct species, but all intermediate conditions exist between a simple, single carina and a truly double carina of two equal-strength ribs. Most interesting is that some in- dividuals of “Т.” praecincta Conrad also show a dou- ble carina (Plate 14, Figure 3). Palmerella mortoni, ssp. Haustator premortoni Govoni, 1983, p. 104, pl.5, f.1-5. Description. —Govoni (1983) provides the following description: “Shell medium-size, tapering, turriculate. Suture narrowly impressed. Protoconch erect, hom- eostrophic, turbinate, of about 3 smooth, rounded, rap- idly expanding volutions; first volution minute, ini- tially somewhat depressed; protoconch merges into te- leoconch without percepible break. Whorls of teleo- conch wider than high; total number of volutions unknown. Primary spiral threads B and C may appear on earliest juvenile whorl simultaneously (B, C,) or, more frequently, C may appear just slightly ahead of B (B; C); within a single volution, primary spiral thread a appears just slightly posterior of the midpoint be- tween B and the posterior suture, while at the same time B shifts slightly anteriorly of midwhorl and strengthens rapidly relative to c, forming a sharp, raised cord, and yielding the sculptural pattern which dom- inates the remaining juvenile whorls (a, B, c, d); pri- mary spiral thread d seems to appear very early but remains more or less obscured beneath succeeding whorls. First fine secondary thread appears no later than fourth to fifth teleoconch whorl between primary spiral a and the posterior suture; additional secondary threads added progressively starting at sixth to seventh whorl, first between primaries a and B, then between B and c; a single secondary thread does not appear between c and anterior suture until eigth to ninth (early adolescent) volution, with 3 or 4 additional threads or weak cords not appearing in interspace until later ad- olescent to mature whorls. Starting in earlier adoles- cent whorls, many spirals, particularly stronger ones, may become finely but distinctly beaded where inter- sected by growth lines; beading persists throughout mature whorls but may become subdued, or late in ontogeny beads may merge and be incorporated into strong, elongate, wavey wrinkles as growth lines strength- en and become raised and crinkled. Whorl profile con- vex and submedially carinate in juvenile whorls; with addition and strengthening of secondary spirals in con- junction with relative weakening of primaries a and b, whorl sides of adolescent whorls initially flatten then quickly become shallowly concave medially above low, sharp, carinate basal angulation that overhangs suc- ceeding whorl; basal angulation formed by relative strengthening of primary c and sometimes the second- ary spiral below; in mature whorls basal carination becomes rounded and increases in prominence, whorl sides become somewhat more constricted medially, and posterior third of whorl may become broadly con- vex or develop a very low secondary carination at po- sition of primary spiral a. Mature whorls covered by numerous fine spiral threads that tend to be of subequal strength on posterior third of whorl, of similar but more obviously alternating strength medially, and strongest on and below the basal carina; in latest whorls spirals appear wavey and disrupted by intersection of strongly wrinkled growth lines; flattened whorl base covered by numerous relatively coarse subequal spiral threads. Growth lines sinuous, indistinct at first, be- coming increasingly prominent in mature whorls; on upper surface of mature whorls above basal carina, growth lines form a deep, nearly symmetrical spiral lateral sinus with gently curved and acutely inclined limbs, with vertex immediately above midwhorl and corresponding to point of maximum constriction of whorl side; lines recurve to form prospiral sinus of similar width and depth centered upon basal carina, then continue in broad antispiral arc across whorl base to columella. Aperture incompletely known, possibly subquadrate; parietal region calloused.” Measurements. —See Table 29. Stratigraphic and Geographic Distribution. — MD: Brightseat Formation; Lower Paleocene. Type Locality.—Southwest Branch, W of Capital Beltway, Prince Georges County, Maryland (MD-PG- 12). Other Localities: MD-PG-13 (GO), MD-PG-14 (GO). Type Material: holotype and 75 paratypes of Haus- tator "premortoni" Govoni USNM Remarks/Discusssion. —See above, under P. mor- toni mortoni and Govoni and Hansen (in press). Table 29.— Measurements of Palmerella mortoni ssp. (*premor- toni" Govoni). Abbreviations as in Table 10. MD MH WN USNM holotype 14.9 39:9 6.5 USNM paratype 4.8 1581 8 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 65 Palmerella pleboides (Vaughan, 1895) Plate 6, Figures 1-3 Turritella pleboides Vaughan, 1895, p.213, nomen nudum, Bowles, 1939, p.311, pl.33, £7; Stenzel and Turner, 1942, card 92, f.4- 6, copies Vaughan, 1895, f.7; Palmer and Brann, 1966, p.1001. Mesalia pleboides Vaughan, 1896, p.36,44, pl.3, f.4-6; Schuchert, Dall, Stanton and Bassler, 1905, p.401; Palmer, 1937, p.208, pl.27, f.5,11,12; Brann and Kent, 1960, p.549,550. Description. —Shell small, of 10 to 15 whorls. Max- imum observed whorl diameter 7.0 mm. Apical angle 20°; pleural angle may be slightly less than apical angle. Sutures relatively deeply incised. Protoconch suberect, homeostrophic, turbinate, of 1.5-1.75 smooth, round- ed whorls; P1 small. Primary spirals B and C appearing approximately simultaneously, C perhaps just slightly before B, as a pair of faint angulations on the lower У of the whorl, rapidly becoming distinct ribs. Primary A appearing within 0.5 whorl after B and C, achieving equal strength within 1.0 whorl, giving early whorls a subquadrate aspect. Secondary ribs appearing at about whorl 6-7. Sculpture on late whorls consisting of even- ly spaced primaries and secondaries of approximately equal strength, separated by weaker intercalaries. Growth lines beginning to show marked positive relief at about whorl 3, producing axial costae particularly on subsutural shelf developed on early whorls between A spiral and suture. Beading caused by intersection of growth lines and spiral ribs persisting after whorl pro- file becomes rounded at about whorl 9-10. Teleoconch whorls wider than high, evenly rounded and somwhat inflated in profile. Area between C spiral and suture concave in profile. Lateral aspect of growth line trace orthocline to prosocline, with only lower inflection point. Sinus of moderate depth with apex just above whorl middle. Antispiral and basal sinues shallow. Measurements. —See Table 30. Stratigraphic and Geographic Distribution. — LA: Cook Mountain Formation; upper Claibornian Stage, Middle Eocene. Type Locality. — Hammetts Branch, 2 mi NE of Mt. Lebanon, Bienville Parish, Louisiana (LA-BI-7). Other Localities. —LA-BI-1 (B), LA-BI-2 (B), LA- BI-4 (B), LA-BI-5 (B), LA-BI-6 (B), LA-BI-8 (B), LA- CB-2 (B). Type Material. —lectotype of T. pleboides Vaughan USNM 495175, paralectotypes USNM 147040. Remarks/Discussion. —The apical sculpture places Table 30.—Measurements of Palmerella pleboides (Vaughan). Ab- breviations as in Table 10. MD MH WN USNM 495175 lectotype 7.0 22.0 9.5 this form in the mortoni group and so in the genus Palmerella. Its convex whorl profile and shallow basal growth line sinus ally it with P. femina (Stenzel), al- though it shows finer sculpture than this Weches spe- cies. Palmerella potomacensis (Clark and Martin, 1901) Plate 15, Figures 1,2 Turritella potomacensis Clark and Martin, 1901, p.149, pl.27, f.2,3; Clark and Miller, 1912, p.104,106,107, 119,120; Bowles, 1939, p.310; Stenzel and Turner, 1942, card 96; Palmer and Brann, 1966, p.1002. Description. —Shell medium-sized, of 12-15 whorls. Maximum observed whorl diameter 11.5 mm. Apical angle approximately 12-15°, approximately equal to pleural angle. Sutures relatively deeply incised. Pro- toconch and earliest teleoconch whorls are unknown. Earliest known whorls (diameter = approx. 1.0 mm) showing two spirals of approximately equal strength in about middle of both upper and lower halves of whorl. Secondary spirals soon appearing. Basal spiral quickly becoming stronger and being joined by a sec- ond subequal spiral beneath it giving the whorls a ba- sally carinate-cingulate appearance. Above basal ca- rina, adult whorl surfaces covered with three to four principal spirals of approximately equal strength and an equal number of finer secondaries. Lateral aspect of growth line trace prosocline, with lower and upper inflection points very close to sutures. Sinus of mod- erate depth just below middle of whorl. Basal aspect of growth line trace unknown. Aperture unknown. Measurements. —See Table 31. Stratigraphic and Geographic Distribution. — V A: Nanjemoy Formation, Woodstock Member; Lower Eocene. Type Locality. —1 mi. below Popes Creek, Charles County, Maryland (MD-CH-2). Other Localities. —- V A-KG-3 (B), VA-HA-2 (WD, MCZ). Type Material. — 2 syntypes of T. potomacensis Clark and Martin (plastic molds) USNM 207075, 207076. Remarks/Discussion. —' The earliest known spiral sculpture, as well as the profile of late whorls, of po- tomacensis are similar to those of P. mortoni mortoni from the underlying Aquia Formation. Although a well Table 31.— Measurements of Palmerella potomacensis (Clark and Martin). Abbreviations as in Table 10. MD MH WN USNM 207075 syntype 8.0 395.0 12 USNM 207076 syntype 11.0 38.0 3.5 66 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 preserved apex is needed for confirmation, these fea- tures are sufficient to unite potomacensis with the mor- toni group for the time being. Palmerella sp. Plate 11, Figure 5 Turritella sp. Ward, 1985, pl.4, f.12, 13. Description. —Shell small to medium sized, of prob- ably 10 to 15 whorls. Maximum observed whorl di- ameter 7.6 mm. Apical angle 20°; pleural angle may be slightly less than apical angle. Sutures indistinct on early whorls, only slightly incised on later whorls. Te- leoconch whorls wider than high; early whorls straight- sided in profile, later whorls evenly rounded and con- vex. Protoconch and earliest apical whorls unknown. Earliest known whorls (dia. = 1.2 mm) bearing four spiral ribs. Two on lower Y. of whorl most prominent, located at around whorl midpoint and in middle of lower half, respectively. Next strongest on about mid- dle of upper Y. of whorl, followed by one much weaker midway to upper suture. Lower two spirals remaining most prominent for several whorls, as intercalaries ap- pear and come to equal strength with upper spirals. Profile of earliest whorls dominated by lower two ribs, upper Y. of whorl sloping straight down to them from the suture. Within approximately six whorls of earliest observed, whorl profile becoming evenly rounded as spirals assume more equal strengths, maximum di- ameter occurring in lower '4 of whorl. Sculpture on latest whorls consisting of approximately evenly spaced, equal strength, moderately pronounced spirals alter- nating with finer spirals. Sutures inconspicuous. Lat- eral aspect of growth line orthocline with weak upper and lower inflection points and moderately deep sinus with apex just above whorl middle. Antispiral and bas- al sinuses relatively shallow. Measurements. —See Table 32. Stratigraphic and Geographic Distribution.—VA: Nanjemoy Formation, Woodstock Member; Lower Eocene. Localities. -VA-HA-3 (WD,MCZ), VA-HA-4 (WD,MCZ). Remarks/Discussion. — This form appears to repre- sent an undescribed species, but insufficient material Table 32.— Measurements of Palmerella sp. Abbreviations as in Table 10. is available to name it formally at this time. The ear- liest known spiral sculpture is similar to that of Pal- merella mortoni and its close relatives, although a com- plete apex is required for positive determination. The whorl profile, adolescent and adult sculpture, and basal aspect of the growth line are most similar to those of T. femina from Gulf coast deposits of slightly later age. Palmerella stenzeli, new species Plate 11, Figures 9,10 Diagnosis. — Medium-sized turritellid with conspic- uous, rounded basal carina, fine, even spiral sculpture and a depressed zone across the middle of the whorl. Description. — Shell medium sized, of probably 15 to 20 whorls. Maximum observed whorl diameter 18.6 mm. Apical angle 17°; pleural angle slightly greater than apical angle. Sutures moderately deeply incised. Protoconch and earliest teleoconch whorls unknown. Smallest whorl examined (dia. = 0.14 mm, paratype MCZIP 29271) bearing three spirals of moderate strength, one on middle of upper Y of whorl, one just below whorl middle and one on about middle of lower 13 of whorl. Lower two spirals slightly stronger than upper one, and lowest spiral forming edge of basal carina, below which whorl profile slightly concave. Sec- ondaries of lesser strength appearing irregularly spaced between these spirals over next three to five whorls; original spirals decreasing in strength so that sculpture of later whorls consisting of numerous more or less equally spaced, equally fine spirals. Spirals not beaded. Whorl profile becoming increasingly basally carinate, basal carina becoming broader and rounded and com- prising approximately lower М of whorl. Middle of whorl becoming markedly concave between height of uppermost original spiral and carina. Lateral aspect of growth line trace orthocline to slightly prosocline, with only lower inflection point. Sinus moderately deep, with apex just above whorl middle. Antispiral nd basal sinuses moderately deep. Measurements. —See Table 33. Stratigraphic and Geographic Distribution. — TX: Kincaid Formation (Tehuacana Limestone); lower Midwayan Stage, Lower Paleocene. Type Locality. —old Flat Rock Quarry, approx. 5 mi. S of Ola, Kaufman County, Texas (TX-KA-5). Table 33. —Measurements of Palmerella stenzeli n. sp. Abbrevi- ations as in Table 10. MD MH WN MD MH WN USNM 366511 9:9 1222 10.5 MCZIP 29270 holotype 9.85 33.4 thes, USNM 366512 59 14.4 4 MCZIP 29272a paratype 12:0 34.0 10 MCZIP 29280 б.7 20.8 752 MCZIP 29273а paratype 1255 47.2 13 MCZIP 29280 7.6 20.3 6 MCZIP 29273b paratype 13.0 44.2 11 ) ~ шш У nn PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 67 Other Localities. —TX-LI-3 (MCZ). Type Material.—holotype of Palmerella stenzeli n. sp. MCZIP 29270, paratypes MCZIP 29271, 29272, 29273, PRI 33198. Etymology. —Named for the late H.B.Stenzel, pio- neer of Texas Cenozic invertebrate paleontology. Remarks/Discussion. — The sculpture of the earliest known whorls of this species most closely resembles that of Palmerella levicunea (Harris), for which the protoconch and earliest teleoconch whorls are also un- known. P. stenzeli is distinct from the other basally carinate Midwayan species of Texas (P. hilli (Gardner), “Turritella” kincaidensis Plummer, 1933 “T.” plum- meri Stenzel and Turner, 1940) in its finer, unbeaded Spiral sculpture and especially the medial concave band across the middle of later whorls. The type locality is close to that of “T.” kincaidensis, but the two forms have not been found together. P. stenzeli lacks the adapical concave constriction characteristic of “7.” kincaidensis. The prominent basal carina allies P. sten- zeli with other similarly shaped lower Paleogene spe- cies of Palmerella, but, as for many species discussed here, definite supraspecific placement must await dis- covery of a well preserved apex. This species is common to abundant in the dense limestones exposed at its two known localities. These limestones appear to belong to either the Tehuacana Member of the Kincaid Formation or to one or more of the limestone “lentils” of the underlying Pisgah Member. Genus HAUSTATOR Montfort, 18106 Type Species. — H. gallious Montfort, 1810 (— Tur- ritella imbricataria Lamarck, 1804), Upper Eocene, England. Diagnosis. — Shell medium-sized to vary large, of 15- 25 whorls. Protoconch of 0.75 to 2.5 smooth, rounded whorls with small РІ. Apical sculpture formula C,B>A;. Profile of adult whorls round to basally carinate. Sculp- ture consisting of multiple simple, moderately devel- Oped spiral ribs. Apical angle less than or equal to pleural angle. Lateral sinus of the “hybrida-imbrica- taria" type, basal sinus moderate deep and simple. Stratigraphic and Geographic Distribution. — U.S. Gulf and Atlantic Coastal Plains, Western Europe; Lower Paleocene— Upper Eocene. $ The name Haustator Montfort, 1810 is a junior subjective syn- onym of Aculea Perry, 1810 (Petit and Le Renard, 1990). In view of the widespread use of Haustator, however, Petit and Le Renard (1991) have proposed that it be conserved and that Aculea be sup- pressed. This was approved by ICZN Opinion 1677 (1992). Haustator alabamiensis (Whitfield, 1865) Plate 7, Figure 1-3 Turritella alabamiensis Whitfield, 1865, p.267; de Gregorio, 1890, p.128; Aldrich, 1894, p.239,246, pl.13, f.2; Harris, 1894b, p.30,48; 1896, p.109, pl.10, £.5; Gardner, 1935, p.284, in part; Bowles, 1939, p.299, pl.32, £.21; Stenzel and Turner, 1942, card 41; Palmer and Brann, 1966, p.979; Toulmin, 1977, p.173, pl.8, f.7; not Coss- mann, 1893, p.29 under tiga. Turritella mortoni Conrad. Smith and Johnson, 1887, p.59, in part; not Conrad, 1830. Turritella alabamiensis var. prealaba Conrad. Kennedy, 1895, p.146, nomen nudum. Turritella (Haustator) alabamiensis Whitfield. Cossmann, 1912, p.118. Description. —Shell medium sized, of 15 to 20 whorls. Maximum observed whorl diameter 12 mm. Apical angle 205 pleural angles slightly greater than apical angle. Sutures deep to shallow. Protoconch of 2.0-2.5 smooth rounded whorls; P1 unknown, probably small. Teleoconch whorls slightly wider than high, straight- sided to basally convex in profile. Apical sculpture formula C, В, A,. Primary sprial C appearing as an- gulation on lower Уз of whorl, followed within ap- proximately 0.5 whorl by primaries B and A, which quickly achieve equal strength with C. Primaries de- creasing in relative strength, joined by numerous sec- ondaries and tertiaries, so that whorls soon covered with evenly spaced, fine spirals of roughly equal strength. Primaries and secondaries may be faintly beaded on intermediate whorls. In most individuals, lower % of whorl, bearing C spiral, becoming increas- ingly convex, eventually forming rounded but prom- inent angulation giving whorl a basally carinate form. Some specimens showing more concave whorl profiles with less pronounced development of basal carina. Lat- eral aspect of growth line trace orthocline or slightly prosocline, usually showing only lower inflection point. Lateral sinus moderately deep, with apex just above middle of whorl. Spiral and basal sinuses moderately deep. Measurements. —See Table 34. Stratigraphic and Geographic Distribution. — AL: Clayton Formation, Porters Creek Formation (Lower Member), Porters Creek Formation (Matthews Land- ing Member), Naheola Formation; GA: Clayton For- mation, undifferentiated Midway Group; TX: Kincaid Table 34. — Measurements of Haustator alabamiensis (Whitfield). Abbreviations as in Table 10. MD MH WN FMNH-UC 24522 syntype 10.0 35.0 11.5 Formation, Wills Point Formation?; AR: Porters Creek Formation; Midwayan Stage, Lower Paleocene. Type Locality.—probably Matthews Landing, Ala- bama River, Wilcox County, Alabama (AL-WI-3). Other Localities. — AL-WI-9 (В), AL-WI-16 (B), AL- WI-21 (B), AL-WI-24 (T), AL-WI-25 (T), AL-WI-36 (B), AL-WI-37 (B), AL-WI-38 (T), AL-WI-39 (T), AL- WI-40 (T), AL-WI-41 (T), AL-WI-42 (T), AL-WI-43 (T), AL-WI-44 (T), AL-WI-45 (T), AL-WI-46 (T), AL- WI-47 (T), AL-WI-48 (T), AL-WI-49 (T), AL-BU-1 (T), AL-BU-2 (T), AL-BU-3 (T), AL-HE-1 (T), AL- MA-2 (T), GA-RA-1 (T), GA-SL-2 (VS), TX-BA-30 (B), TX-BA-31 (B), TX-BA-32 (US), TX-CA-1 (B), TX-CA-3 (US), TX-FL-7 (B), TX-HU-1 (B), TX-KA-4 (B), TX-KA-1 (B), AR-PU-1 (PB). Type Material. —11 syntypes of T. alabamiensis Whitfield FMNH-UC 24522. Remarks/Discussion. — Haustator alabamiensis and “Turritella” aldrichi Bowles (see p.81) are often quite similar in their adult whorl shape and may be initially difficult to separate. These two species are very distinct in their early apical sculpture and whorl shape, how- ever, and in the extremes of their adult whorl form as well. This similarity was discussed by Bowles (1939), who defended the separation of the two taxa “because of their apparent relationships to the latter well-estab- lished and distinct groups of T. [herein Palmerella] mortoni and T. humerosa.” Haustator carinata (I.Lea, 1833) Plate 9, Figures 1-11 Turritella carinata I.Lea, 1833, p.129, pl.4, f.120; H.C.Lea, 1849, p.107; Dana, 1863, f.802; 1895, f.1486, copy 1863; Conrad, 1865a, p.32; 1866, p.11; de Gregorio, 1890, in part, pl. 11, £.3-5, f.6, copy I.Lea, 1833, f.9, copy T. mortoni Conrad, 1835a; not Heil- prin, 1891, p.400; Cossmann, 1893, p.29; Harris, 1895b, p.10; Palmer, 1937, p.189, pl.24, f.5,6,8,9,12, p.82, f.1; Bowles, 1939, p.304, pl.33, f.13; Stenzel and Turner, 1942, card 52; Brann and Kent, 1960, p.904, 905; Glibert, 1962, p.95. Palmer and Brann, 1966, p.983; not T. carinata H.C.Lea, 1841, p.96; not H.C. Lea, 1849, p.107 [= “Т”. apita de Gregorio]; Toulmin, 1977, p.301, pl.50, f.3; Dockery, 1980, p.81, pl.29, f.1. Turritella mortoni Conrad. Conrad, 18352, p.40, in part, pl.15, f.11 [Harris reprint 1893, p.40, pl.15, £.11]; not 7. mortoni Conrad, 1830. - Turritella mortoni var. A Conrad, 1835a, p.40; not T. mortoni Con- rad, 1830. Turritella gracilis H.C.Lea, 1841, p.97, pl.1, f.12; 1849, p.107; de Gregorio, 1890, p.127, pl.11, #32, copy LLea, 1833; Harris, 1895b, p.21. Turritella monilifera H.C.Lea, 1841, p.97, pl.1, Ғ11; 1849, p.107; Harris, 1895b, p.29; not T. monilifera Adams and Reeve, 1848 (1850), p.48; not 7. monilifera Deshayes, 1833, p.275. Turritella mut. tiga de Gregorio, 1890, p.126, pl.11, f.22; Cossmann, 1893, p.29. Turritella ghigna de Gregorio, 1890, p.125, pl.11, £19; Palmer, 1937, p.191, pl.24, f.2,4,11,13,15, pl.83, £.1; Stenzel and Turner, 1942, PALAEONTOGRAPHICA AMERICANA, NUMBER 59 card 65; Brann and Kent, 1960, p.914; Glibert, 1962, p.97; Palmer and Brann, 1966, p.988; Toulmin, 1977, p.302, pl.50, f.4. Turritella litripa de Gregorio, 1890, p.125, pl.11, £.20. Turritella carinifera claibornensis de Gregorio, 1890, p.126, pl.11, £.33, copy Т. monilifera H.C.Lea, 1841. Turritella hybrida Deshayes. de Gregorio, 1890, p.126, pl.11, £.23; not T. hybrida Deshayes, 1833, p.278. Turritella eterina de Gregorio, 1890, p.126, pl.11, f.34-36; Coss- mann, 1893, p.29. Turritella claibornensis de Gregorio. Cossmann, 1893, p.29. Turritella (Haustator) claibornensis de Gregorio. Cossmann, 1912, p.118. Turritella (Haustator) claibornensis tiga de Gregorio. Cossmann, 1912, p.118. Turritella (Haustator) claibornensis eterina de Gregorio. Cossmann, 1912, p.118. Turritella mortoni turneri Palmer, 1937, p.194, pl.23, f.3,7; not T. mortoni Conrad, 1830; not T. turneri Plummer, 1933. Turritella carinata palmerae Bowles, 1939, pl.33, f.12; Stenzel and Turner, 1942, сага 61;- Palmer and Brann, 1966, р.983-4; Toul- min, 1977, p.302, pl.50, f.1. Description. —Shell medium sized to large, of 15 to 20 whorls. Maximum observed whorl diameter 18.6 mm. Apical angle 15°; pleural angle varying from ap- proximately equal to apical angle to substantially great- er. Sutures varying from faint to deeply incised. Pro- toconch suberect, homeostrophic, turbinate, of 0.75- 1.25 smooth, somewhat flattened whorls; P1 small. Apical sculpture formula C, B, Аз. Primary C spiral appearing as angulation on lower 3—2 of whorl, fol- lowed after about 1.0 whorl by B spiral at about middle of whorl and closely associated with C. B and C quickly reaching equal strength and diverging. A spiral follow- ing B within about 0.5-0.75 whorls, quickly reaching equal strength with B and C. Very faint secondaries appearing between primaries beginning at whorls 8-9, becoming obsolete or persisting as very fine spirals, never as strong as primaries. Faint beading often pres- ent on primaries on intermediate whorls. Adult whorls slightly to very basally carinate, carina being formed by D spiral at basal angulation of whorl just above suture. Point of contact ofa succeeding whorl may shift inward on base of preceeding whorl from margin of angulation, forming overhanging carina, or may re- main close to margin, pleural angle remaining higher. Measurements. —See Table 35. Stratigraphic and Geographic Distribution. —AL: Gosport Sand, Upper Lisbon Formation; MS: Cook Mountain Formation; GA: McBean Formation’, Lis- bon Formation; SC: McBean Formation’; LA: Cook Mountain Formation; upper Claibornian State, Middle Eocene. Type Locality.—Claiborne Bluff, Alabama River, Monroe County, Alabama (AL-MO- 1a). Other Localities. — AL-MO-4,b,c (P,B), AL-MO-11 7 See Footnote 3, p.46. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 69 Table 35.—Measurements of Haustator carinata (I.Lea). Abbre- viations as in Table 10. MD MH WN ANSP 5661 lectotype 11.6 22.0 SI ANSP 5662 syntype 9.4 28.1 VS) ANSP 13174 312 DE 12.0 ANSP 13175 WI 19:2 8.5 РКІ 2893 16.3 47.1 10.5 РКІ 2889 II 21.8 4.5 PRI 2890 9.3 3241 12:5 PRI 2891 20.0 30.2 2.0 MCZIP 29292 11.9 36.5 8.0 MCZIP 29293 16.0 48.5 8.0 MCZIP 29294 12:2 44.0 10.5 MCZIP 29296 18.0 44.3 6.0 MCZIP 29299 ТӨРІ 44.6 8.5 (Т), AL-CL-4 (MCZ), AL-CH-9 (Т), AL-WA-6(MCZ), MS-NE-1 (D,US), MS-NE-2 (P), MS-NE-3 (PB), MS- NE-7 (D), MS-NE-10 (US), MS-CL-1 (P,B), MS-CL- 16 (B), MS-CL-17 (B), GA-BU-3 (VS), GA-TW-7 (VS), GA-HO-1 (VS), GA-HO-2 (VS), GA-SU-1 (VS), GA- RA-4 (VS), GA-GL-1 (VS), GA-WA-4 (VS), GA-WA-5 (VS), GA-WI-1 (VS), GA-JO-1 (VS), GA-BU-8 (VS), GA-BU-9 (VS), GA-BU-10 (ANS), GA-JF-1 (VS), GA- BI-1 (VS), SC-AI-1 (B), SC-OR-6 (B), SC-OR-7 (B), LA-WI-1 (B). Type Material. —lectotype of T. carinata 1.1 еа ANSP 5661, syntypes ANSP 5662—5667; holotype and para- type ? of T. gracilis H.C.Lea ANSP 13174; holotype of T. claibornensis tiga de Gregorio PRI 26440; ho- lotype of T. ghigna de Gregorio lost (fide Palmer and Brann, 1966, p.988); lectotype and paratype? of T. monilifera H.C.Lea ANSP 13175; holotype of T. m. litripa de Gregorio PRI 26441; holotype of T. eterina de Gregorio lost (fide Palmer and Brann, 1966, p.988); holotype of T. carinata palmerae Bowles USNM 497997. Remarks/Discussion. —Stenzel and Turner (1942), Bowles (1939) and Palmer (1937) all note the vari- ability within “Turritella carinata," and specifically that the forms designated as Turritella ghigna de Gre- gorio and Turritella carinata I.Lea intergrade in adult whorl form and sculpture. Palmer states that “When the intimate relation of the two lines of growth [i.e., ghigna and carinata lineages] is seen, one wonders if they are not dimorphic phases of the same species" (1937, p.189). Although she recognizes the two as sep- arate species, for the sake of “taxonomic convenience," Palmer continues to refer to them as “phases” of a common stock. Typical ghigna and typical carinata two forms differ (and intergrade) in both whorl shape and expression of spiral sculpture on the adult whorls; late whorls of typical ghigna are basally convex, but rounded, and bear prominent primary and secondary spirals, where- as those of typical carinata have sharp basal carinae and are generally smooth. Early whorls of the two forms are generally similar, although some forms of carinata show particularly attenuate upper spires. The intergra- dation in sculptural and whorl shape is in a qualitative sense gradual and complete, even among individuals co-occurring in the same sample, and fully justifies recognition of only a single, albeit highly variable, spe- cies. The variation can be seen clearly in Plate 9. Results of multivariate morphometric analysis of “T. carinata” sensu lato are presented in Text-figure 20. A total of 117 variables was measured on each of 143 specimens. The first three factor axes account for 57.3% of the total variance (Table 36). Loadings on the first three factors (Table 37) indicate that variables on the fourth and fifth whorls load most heavily on the first factor, whereas variables from the eleventh and twelth whorls load most heavily on the second factor and variables on seventh whorl load most heavi- ly on the third factor. It is clear from the distribution of specimens that there are no discrete groupings rep- resented, but rather a continuum of variation, mostly along the first factor axis. The specimens plotting in the upper left hand quadrant of Text-figure 20A are those with later whorls (/.е., whorls 11 and higher) perserved, but these also have the highly carinate mor- phology of carinata s.s. (e.g., Plate 9, figures 6,7). Spec- imens assignable to carinata palmerae (Plate 9, figure 10) and ghigna (Plate 9, figure 3,4) fall along the right or positive side of the graph, along the first factor axis. Qualitatively, H. carinata from the Gosport Sand differs from H. rina (Palmer) from the underlying Lis- bon Formation in the persistence of its B spiral onto adult whorls, as well as the frequently more reduced expression of primaries A and C, which are quite prom- inent on late whorls in H. rina. If only these two forms were known, it might be reasonable to propose an an- cestor-descendant relationship between them. Forms from the Upper Lisbon of Alabama, described by Bowles (1939) as “Turritella carinata palmerae," and the equivalent Cook Mountain of Mississippi, figured by Dockery (1980), however, suggest that H. carinata originated well before the deposition of the Gosport, and that it showed a simlar range of variability early in its history as it did in the Gosport. “7. с. palmerae” is similar to the ghigna end of the Gosport carinata spectrum, whereas the Mississippi Cook Mountain specimens are close to the typical carinata form. Both “Т.” c. palmerae and the Mississippi forms, however, are substantially smaller than the largest sizes of the Gosport specimens, suggesting a trend of increasing size within the lineage. 70 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 a 4 + N 3 d з 7 © ш 2 mu Ba D ED m T El | | | о | E EE TEES H ошоо Ue. ы H oto ego ug uu AA 2 15 co „ar $ 0.5 ы. 1.5 2 | Ds wo Factor 1 B = О s с A | | | EI 1 22 -1 1 2 3 4 5 Г] с o o 5 00556 8 ш -3 4 4 + О o -5 pela Factor 2 Text-figure 20.— Results of factor analysis of specimens of Haustator carinata (I. Lea). A. first and second factor axes. B. Second and third factor axes. See Appendix 1 for specimens used and details of analysis. Table 36.—Results of factor analysis of specimens of Haustator carinata (I. Lea). Variance explained by first five factor axes. Haustator cortezi (Bowles, 1939) Plate 10, Figure 9 Factor Edo wi вену Turritella cortezi Bowles, 1939, p.280, pl.31, Ғ.11,15; Stenzel and Turner, 1942, card 57; Palmer in Harris and Palmer, 1947, p.296- 1 0.3082 297; Palmer and Brann, 1966, p.985; Allison and Adegoke, 1969, 2 0.4767 p.1253. 3 0.5725 | = Description. —Shell small to medium sized, of 10 to 15 whorls. Maximum observed whorl diameter 13.5 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 71 Table 37.—Results of factor analysis of specimens of Haustator carinata (I. Lea). Rotated factor loadings of each variable on each of the first three factor axes. Variables as indicated in Text-figure 3. Variable factor 1 factor 2 factor 3 Variable factor 1 factor 2 factor 3 WHI 0.110 0.192 —0.541 W2-7 0.330 0.250) 0.764 CH2-1 0.110 0.181 —0.540 W3-7 0.344 —0.249 0.754 CH4-1 0.000 0.000 0.000 CW7 0.328 5025 0.765 | SWI 0.109 zone = 0,980 CDC7 0.285 —0.249 0.784 | WI-1 0.110 =0.152 —0.541 WH8 70:385 —0.294 0.634 | W2-1 0.110 0,152 —0.541 CH2-8 =0.352 —0.293 0.611 \ W3-1 0.110 04152 —0.541 CH4-8 —0.431 —0.154 0.420 | Cwl 0.110 20.182. —0.541 SWS8 —0.446 —0.227 0.558 | CDCI 0.110 —0.152 —0.541 W1-8 0,398 0.309 0.611 1 WH2 0.580 ОВ —0.511 W2-8 —0.385 —0.310 0.608 CH2-2 0.578 =0.112 0,520 W3-8 0370) —0.306 0.609 CH4-2 0.339 —0.045 200159 CWS8 —0.382 —0.304 0.606 SW2 0:572 —0.100 —0.478 CDC8 —0.411 = 0,303 0.612 | W1-2 0.581 —0,109 —0.501 WH9 20758 0.075 0.222 | W2-2 0.581 USOS 220/501. CH2-9 —0.700 —0.079 0.208 1 W3-2 0.581 =0.109 – 0.494 СНА-9 = (0622 —0.026 0.158 ; CW2 0.582 —0.108 —0.495 SW9 —0.689 — 0.043 0.222 | CDC2 0.581 —0.109 —0.496 W1-9 —0.754 —0.069 0.207 | WH3 0.768 307102 --();369 W2-9 —0.748 —0.062 0.208 J CH2-3 0.720 —0.109 0350 W3-9 —0.745 —0.071 0.211 CH4-3 0,393 —0.047 —0.110 CW9 0751 0.097 0.205 SW3 0.586 —0.060 5:20:29 9 CDC9 —0.748 2201035 0:213 W1-3 0.776 ОЛО —0.338 WH10 —0.630 0.318 —0.114 W2-3 0:777, 2204112 0850 CH2-10 =0896 0.293 —0.120 1 W3-3 0270, ipm 70.350 CH4-10 —0.534 0.320 —0.065 | CW3 0.777 70:112 —0.355 SWIO —0.582 0.344 —0.074 | CDC3 0.773 —0.114 —0.349 W1-10 —0.641 0.355 —0.128 | WH4 0.819 —0.034 —0.169 W2-10 =0.037 0,332 –0.134 | CH2-4 0.794 —0.035 —0.178 W3-10 0:687 0.332 = 0.193 CH4-4 0.318 0.137 0.180 CW10 —0.638 0.344 0,136 Sw4 0.645 0.040 —0.047 CDC10 0:083 0.344 —0.129 | W1-4 0.816 --(),032 —0.156 WHIl —0.179 0.748 —0.050 | W2-4 0.824 —0.035 —0.170 CH2-11 0:182 0.736 —0.058 W3-4 0.828 —0.034 —0.169 CH4-11 —0.143 0.741 0.005 Cw4 0.830 —0.038 ОЯ sw11 0.174 0.788 —0.036 | CDC4 0.816 —0.031 —0.156 Wi-il —0,185 0.755 —0.066 | WHS5 0.860 —0.063 0.068 W2-11 —0.185 0.767 0071 CH2-5 0.842 —0.069 0.047 W3-11 —0.184 0.763 —0.069 CH4-5 0.382 0.007 0.158 СМ11 – 0.182 0.769 – 0.063 SW5 0.682 2201017 0.127 CDC11 0.178 0.757 —0.056 W1-5 0.850 =0:073 0.095 WH12 —0.062 0.830 —0.057 W2-5 0.862 0.077 0.072 СН2-12 —0,059 0.840 —0.062 W3-5 0.867 —0.078 0.058 CH4-12 EO 0.620 --0,017 | CWS 0.865 —0.082 0.063 SW12 —0.063 0.823 —0.046 | CDC5 0.842 2301079. 0.085 WI-12 —0.057 0.856 0.097 ; WH6 0.765 —0.180 0.319 W2-12 —0.055 0.856 70:052 CH2-6 0.758 —0.182 0.288 W3-12 —0.058 0.848 —0.056 CH4-6 0.369 —0.074 0.354 CWI12 —0.058 0.847 —0.057 SW6 0.486 0182 0.337 CDC12 —0.060 0.839 —0.056 | W1-6 0.744 =0.187 0.343 WH13 0.007 0.674 0.023 | W2-6 0.771 —0.190 0.306 CH2-13 0.007 0.674 0.023 | W3-6 0.782 70.191 0.297 CH4-13 0.007 0.674 0.023 4 CW6 0.770 —0.194 0.302 SWI13 0.007 0.674 0.023 | CDC6 0.731 04190 0.327 W1-13 0.007 0.674 0.023 WH7 0.340 0283 0.787 W2-13 0.007 0.674 0.023 | CH2-7 0.349 —0.234 0.749 W3-13 0.007 0.674 0.023 | СН4-7 0.078 —0.085 0.543 CW13 0.007 0.674 0.023 | SW7 0.151 —0.180 0.703 CDC13 0.007 0.674 0.023 W1-7 0.302 —0.249 0.773 72 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 38.—Measurements of Haustator cortezi (Bowles). Abbre- viations as in Table 10. Table 39.—Measurements of Haustator fischeri (Palmer). Abbre- viations as in Table 10. MD MH WN MD MH WN USNM 495172 holotype 11.0 30.0 8.0 FGS I 7399 holotype T5 39.3 б USNM 495173 paratype 9.0 22.0 6.0 FGS I 7400 paratype 6.2 19.4 15 FGS I 7401 paratype 1970) 31.0 6 FGS I 7402 paratype 14.8 58.9 12 FGS I 7403 paratype 14.0 34.0 © FGS I 7400 topotype 16.8 21.9 3 mm. Apical angle approximately 15°; pleural angle in- FLMNH 19122 topotypes 1 3 m 5 creasing slightly until late whorls where it may decrease slightly; varying from approximately equal to apical angle to slightly greater. Sutures faint, not incised. Pro- toconch and earliest teleoconch whorls unknown. Ear- liest known whorls having A and C spirals equally prominent, with B spiral absent. A spiral expanding in later whorls to be much more prominent than C, mak- ing whorls adapically carinate, but may be again some- what reduced on last whorl. Measurements. —See Table 38. Stratigraphic and Geographic Distribution. —MX, TX: Cook Mountain Formation; upper Claibornian Stage, Upper Eocene. Type Locality.—near Mier, Tamaulipas, Mexico (MX-TA-1). Other Localities. —VX-LA-1 (B), TX-LA-2 (B), TX- LA-3 (B), TX-LA-4 (B), TX-LA-5 (B), TX-ZA-2 (B), TX-ZA-3 (B), TX-ZA-4 (B), TX-ZA-5 (B). Type Material. —holotype of T. cortezi Bowles USNM 495172, paratype USNM 495173. Remarks/Discussion. — Based on the description giv- en by Stenzel and Turner (1942), Allison and Adegoke (1969) suggest that the spiral ontogeny in this species is C, B, a, ^ d C, В, Aj > dC, Б, A, >d C, A,, but add that the A spiral may appear after B and C. With- out well preserved apices, however, the apical sculpture formula remains uncertain. If cortezi actually shows a pattern similar to other species in the rina group, it is probably most closely related to Haustator subrina, which also shows extreme development of primaries A and C; cortezi differs from subrina chiefly in its small- er size, greater apical angle and slightly reduced A rel- ative to C on latest whorls. Haustator fischeri (Palmer in Richards and Palmer, 1953) Plate 10, Figure 10 Turritella fischeri Palmer in Richards and Palmer, 1953, p.14, pl.1, f.1-3; Palmer and Brann, 1966, p.988; Allison and Adegoke, 1969, р.1253; Toulmin, 1977, р.335-6, pl.62, Ғ14. Description. —Shell medium sized, of 15 to 20 whorls. Maximum observed whorl diameter 12.3 mm. Apical angle approximately 20*, approximately equal to pleu- ral angle. Sutures shallowly incised on early whorls, becoming slightly more deeply incised on later whorls. Protoconch and earliest teleoconch whorls unknown. Earliest known teleoconch whorls (1.2 mm dia.) show- ing A, B and C spirals, more or less equally developed. By about whorl 8-10, B spiral becoming weaker, con- tinuing onto later whorls with whorl profile becoming cingulate and C spiral becoming stronger than A. D spiral visible only on later whorls, leading to slightly basally carinate shape of largest whorls. Secondary spi- rals consisting of one or two above A and one between A and B, visible by about whorl 8-10. C and D rarely equal in strength, forming cingulate band around base of whorl. Weathered specimens often appearing adap- ically carinate with rounded carina. Lateral aspect of growth line trace usually orthocline, with apex of lateral sinus just above middle of whorl. Basal growth line trace very simple. Measurements. —See Table 39. Stratigraphic and Geographic Distribution. —FL: Moody's Branch Formation (Inglis Member); lower Jacksonian Stage, Upper Eocene. Type Locality.—pit N of town of Gulf Hammock, Levy County, Florida (FL-LE-1). Other Localities. — FL-LE-2 (RP). Type Material. —holotype of T. fischeri Palmer FGS I 7399, paratypes FGS I 7400-7403. Remarks/Discussion. —Discussing the affinities of Haustator fischeri, Allison and Adegoke (1969, p.1254) suggest that the simultaneous reduction of the B spiral and strengthening of the D spiral “document the re- lation of this distinct species to Turritella rina subsp. rina Palmer." Haustator gilberti (Bowles, 1939) Plate 7, Figures 4,5 Turritella carinata Y.Lea. Aldrich, 1894, p.235; Brantly, 1920, p.156; not T. carinata I.Lea,1833, p.129. Turritella lineata I.Lea. Smith, Johnson and Langdon, 1894, p.156; not T. lineata I.Lea, 1833, p.130. Turritella clevelandia Harris. Harris, 1897b, p.32; not 1894b, p.170. Turritella clevelandia Harris var. Harris, 1899a, p.74, pl.10, £2; Brann and Kent, 1960, p.907. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 73 Table 40.—Measurements of Haustator gilberti (Bowles). Abbre- viations as in Table 10. Table 41.—Measurements of Haustator infans (Stenzel and Tur- ner). Abbreviations as in Table 10. MD MH WN MD MH WN USNM 494993 holotype 10.0 395.5 DES TMM/TBEG 20955 syntype 25 8.5 9 TMM/TBEG 20956 syntype 2.8 9.3 10 TMM/TBEG 20957 syntype 3.0 8.5 9 TMM/TBEG 20958 syntype 34 10.5 11 Turritella gilberti Bowles, 1939, p.302, pl.32, f.16; Stenzel and Tur- ner, 1942, card 66; Palmer in Harris and Palmer, 1947, p.291; Palmer and Brann, 1966, p.989; Toulmin, 1969, pl. 2, Е.10; 1977, p.231-232, pl.29, f.7; Dockery, 1980, p.79-80, pl.2, f.2. Description. —Shell small to medium sized, of 10 to 15 whorls. Maximum observed whorl diameter 9.6 mm. Apical angle 20°; apical and pleural angles ap- proximately equal. Sutures moderately incised. Pro- toconch suberect, homeostrophic, turbinate, of 1.75- 2.0 smooth, rounded, somewhat flattened whorls; P1 small. Apical sculpture formula C, B; Аз. Primary spi- ral C appearing as angulation just below middle of whorl, followed within about 0.5 whorls by primary B which appears just above C. Within next whorl B and С diverging until С lies in lower У of whorl and B just below whorl middle. Primary A appearing within 0.5 whorl after B and quickly attaining strength equal to Band С. Primary spirals and slight excavation between C spiral and suture giving succeeding teleoconch whorls tricarinate, subquadrate appearance. Secondary spirals appearing between primaries beginning around whorl 7 or 8, but remaining faint onto adult whorls. Lateral aspect of growth line trace orthocline or slightly pro- socline with normally only lower inflection point. Lat- eral sinus moderately deep with apex above whorl mid- dle. Spiral and basal sinuses moderately deep. Measurements. —See Table 40. Stratigraphic and Geographic Distribution. — AL, MS, GA: Bashi Formation; LA: Marthaville Formation; upper Sabinian Stage, Upper Paleocene-Lower Eocene. Type Locality. — Woods Bluff, Tombigbee River, Clarke County, Alabama (AL-CL-2). Other Localities. — AL-CL-1b (B), AL-CL-1c (B), AL- CL-1d (B), AL-CO-1 (B), AL-CO-2 (B), AL-DA-3 (US), AL-WA-2 (T), AL-BU-4 (T), AL-CL-7 (T), LA-NA- 14 (L), MS-LA-3 (B), MS-LA-4 (B), GA-RA-3 (MCZ). Type Material. —holotype of T. gilberti Bowles USNM 494993. Remarks/Discussion. — There are at least two alter- natives for the ancestry of Haustator gilberti; either it arose from H. alabamiensis (Whitfield), the only known Midwayan species with а C, В, A; apical sculpture formula, or from an as yet unknown form. Although a transition from alabamiensis to gilberti would have involved a size decrease, and this seems to have been generally uncommon in turritellid evolution, it seems the preferable choice considering the high fossil content of Wilcox and Midway Group units that seem to con- tain no other plausible ancestor. Haustator infans (Stenzel and Turner, 1940) Plate 8, Figure 10 Turritella dumblei Harris n.var., Stenzel in Renick and Stenzel, 1931 p.102. Turritella infans Stenzel and Turner, 1940, p.838, pl.47, £.12-15; 1942, card 71; Brann and Kent, 1960, p.921; Palmer and Brann 1966, p.992. > э Description. —Shell small to very small, known ѕрес- imens of fewer than 10 whorls. Maximum observed whorl diameter 3.7 mm. Apical angle 19°; apical and pleural angles approximately equal. Sutures moderate- ly incised. Protoconch suberect, homeostrophic, tur- binate, of 1.5-2.0 smooth, slightly flattened whorls; P1 small. Teleoconch whorls wider than high, straight- sided to slightly rounded in profile. Apical sculpture formula C, В; А; ог C, B; A,. Primary spiral С ap- pearing as angulation on lower Y of whorl, followed 1.0-1.5 whorls later by A and B, which appear more or less simultaneously or B slightly earlier. C remaining strongest for 1-2 whorls then all three primaries at- taining equal strength and remaining so, giving later whorls a tricarinate appearance. Secondary spirals ap- pearing on approximately whorl 6-7. Area between C spiral and suture somewhat incised giving whorls a subquadrate form but base of whorl not concave so sutures not very profound in appearance. Measurements. —See Table 41. Stratigraphic and Geographic Distribution. — TX: Cook Mountain Formation (Wheelock Member), Stone City Beds; Claibornian Stage, Upper Eocene. Type Locality. —'*Moseley's Ferry", Stone City Bluff, Brazos River, Burleson County, Texas (TX-BU- 1). Other Localities. — TX-BU-2 (PB) 7 Type Material. —syntypes of T. infans Stenzel and Turner TMM/TBEG 20955, 20956, 20957, 20958. Remarks/Discussion. —Stenzel and Turner note that faint beading is present on the spirals but I have not seen this on the syntypes. I have examined only the four syntypes, and only through the light microscope. The species is apparently rare in the fauna of the Stone City Beds, and appears genuinely to be a small species, 74 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 rather than just the young whorls of a larger form. Although examination of SEM is usually required for recognition of apical sculpture, the C spiral is promi- nent enough on the well preserved type specimens to recognize that it is the first to appear. How closely associated the appearances of A and B are must await SEM study of further material. The early whorls of H. infans are very similar to those of H. perdita (Conrad) from the Upper Eocene Moody’s Branch Formation of the central Gulf, show- ing both the same apical sculpture formula and flat- tened whorl shape. They differ chiefly in the apparently later appearance of primary spirals B and A, and the dissociation of B from C when it does appear. Haus- tator gilberti (Bowles) is almost certainly on the an- cestral line of infans; the two form share small size, subquadrate whorl shape and tricarinate late whorl sculpture. Haustator martinensis (Dall, 1892) Plate 15, Figure 3 Turritella indenta var. martinensis Dall, 1892, p.308; Schuchert, Dall, Stanton and Bassler, 1905, p.677. Turritella indenta Conrad. McCallie, 1908, p.357; not T.indenta Conrad, 1841. Turritella martinensis Dall. Cooke, 1915, p.111; Bowles, 1939, p.282, pl.31, £.10; Stenzel and Turner, 1942, card 77; Palmer and Brann, 1966, p.993; Nicol, Shaak, and Hoganson, 1976, p.137ff, f.2. Torcula martinensis martinensis (Dall). Allison and Adegoke, 1969, p.1263, pl.148, f.5. Torcula martinensis henkeri Adegoke in Allison and Adegoke, 1969, р.1263-5, pl.147, f.2, pl.148, f.8,12, text-f.2,A,B,D,F,G. Description. —Shell medium sized, of 15-20 whorls. Maximum observed whorl diameter 13.0 mm. Apical angle approximately 17°; pleural angle approximately equal. Sutures moderately incised. Protoconch and ear- liest teleoconch whorls unknown. Earliest known whorls showing A and C spirals more or less prominent. C spiral expanding with basal portion of whorl in later whorls, until whorls very slightly basally carinate. B spiral never more than faint thread on later whorls in middle of wide sulcus at mid-whorl. At least on other faint thread in sulcus on later whorls. Measurements. —See Table 42. Stratigraphic and Geographic Distribution. — FL: Crystal River Formation; GA?, MS?; Dept. Bolivar, Colombia; Middle to Upper Eocene. Type Locality. —Martin Station, Marion County, Table 42.— Measurements of Haustator martinensis (Dall). Ab- breviations as in Table 10. MD MH WN USNM 498390 lectotype 12.9 35.5 8.5 Florida (FL-MA-1) (not Hernando County as often stated; Nicol et al., 1976). Other Localities. — FL-. Type Material.—lectotype of T. martinensis Dall USNM 498390, syntypes USNM 112589, hypotype (Allison and Adegoke, 1969) UCMP 33743; holotype of Torcula martinensis henkeri Adegoke UCMP 32466, paratypes UCMP 32928, 32922, 32927, 32946, 32959, 32986, 33057. Remarks/Discussion. — The stratigraphic and geo- graphic distribution of this species have been unclear since its description. Allison and Adegoke (1969) con- sider records of the species from the Ocala Limestone and Barnwell Formation of Georgia (Bowles, 1939, p.283), the Forest Hills Sand of Mississippi (MacNeil, 1944, pp.1317,1326,1327), and Oligocene strata from Citrus County, Florida (Vernon, 1951, pp.166, 174- 175) to be questionable, and perhaps assignable to oth- er species such as H. fisheri (Palmer) or H. subtilis (Kellum). Chiefly on the basis of associated macrofos- sils, Allison and Adegoke believe H. martinensis from the type area to be of Oligocene age, possibly restricted to the Early Oligocene (assumed by MacNeil, 1944, 1946) who used the species as a guide fossil for this interval. Nicol et al. (1976), however, have pointed out that on the basis of microfaunas the limestones from the vicinity of Martin are of Late Eocene age and con- clude that, whatever the age of putative occurrences elsewhere, H. martinensis in the type area is of Late Eocene age. They also note that this redetermination is consistent with the age of the subspecies (Т. m. henkeri") described by Adegoke (in Allison and Ade- goke, 1969) from Middle to Upper Eocene deposits of Colombia. Allison and Adegoke (1969) place H. martinensis in the genus Torcula Gray as its earliest known species, chiefly on the basis of its inferred apical sculpture for- mula, which they suggest is d C, b; A, based on the earliest observed teleoconch whorls. These authors are not specific about the ancestry of H. martinensis, but suggest (1969, p.1265) that some member of their “Turritella rina group" (= Haustator herein) is ances- tral to Torcula. Haustator perdita (Conrad, 1865b) Plate 8, Figure 1—9 Turritella perdita Conrad, 18655, p.141, pl.10, f.10; 1866, p.11; Bowles, 1939, p.307, pl.32, f.11; Stenzel and Turner, 1942, card 90; Palmer in Harris and Palmer, 1947, p.292, pl.37, f.1-3,6,8- 11; Brann and Kent, 1960, p.930-1; Palmer and Brann, 1966, p.1000; Dockery, 1977, p.45, pl.3, £.8,9; Toulmin, 1977, p.336, pl.62, f.11. Turritella jacksonensis Cooke, 1926, p.136, f.8. Turritella lowei Cooke, 1926, p.136, f.9. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 75 Turritella perdita jacksonensis Cooke. Stenzel and Turner, 1942, card 72; Palmer in Harris and Palmer, 1947, p.294, pl.37, Ё4,5,7,12, copy Cooke, 1926; Brann and Kent, 1960, p.931; Palmer and Brann, 1966, p.1001; Dockery, 1977, p.45, pl.3, £.7; 1980, p.82, ПО ОНИ SS. Turritella perdita lowei Cooke. Stenzel and Turner, 1942, card 76. Description. —Shell small to medium sized, of 15- 20 whorls. Maximum observed whorl diameter 13.3 mm. Apical angle 16°; apical and pleural angles usually approximately equal, but pleural may be slightly great- er. Sutures moderately incised. Protoconch suberect, homeostrophic, turbinate but slightly flattened, con- sisting of approximately 1.75-2.0 smooth, rounded whorls; Pl small. Teleoconch whorls wider than high, Straight-sided to concave in profile. Apical sculpture formula C, B, Аз. Primary spiral C appearing as an- gulation just below middle of whorl and quickly be- coming prominent. Primary B appearing 0.5-1.0 whorls later at or just above middle of whorl, at variable dis- tance above C, and quickly achieving approximately equal strength. B and C immediately diverging until C lies in lower У of whorl and B at around middle. Pri- mary A appearing in top Y of whorl 0.5-1.0 whorls after B, and in most forms quickly becoming approx- imately equal in strength to B and C. Presence and strength of secondaries after appearance of A variable. Minor spirals may be limited to single weak second- aries between primaries or primaries decreasing in Strength until entire whorl covered by numerous, even- ly spaced, equally fine spirals. Primaries often showing minor beading on intermediate whorls. Beyond inter- mediate whorls, B spiral decreasing in strength, whorl width contracting between primaries A and C, giving later whorls a slightly concave profile. Primary C usu- ally strongest on latest whorls of larger specimens. Area between C spiral and suture more or less excavated, giving whorls a subquadrate aspect. Lateral aspect of growth line trace orthocline to slightly prosocline, usu- ally with upper and lower inflection points and mod- erately deep sinus whose apex may be above, below or on whorl middle. Spiral and basal sinuses of shallow to moderate depth. Measurements. —See Table 43. Stratigraphic and Geographic Distribution. — MS, LA: Moodys Branch Formation; lower Jacksonian Stage, Upper Eocene. Type Locality. — T. perdita Conrad, Garland Creek, Clarke County, Mississippi (MS-CL-3); T. p. jackso- nensis Cooke, T. p. lowei Cooke, Moody's Branch, Jackson, Hinds County, Mississippi (MS-HN-4). Other Localities. — MS-CL-3 (B), MS-CL-15 (MCZ), MS-HN-1b (B), MS-HN-1e (B), MS-HN-3 (D), MS- HN-6 (D), MS-YZ-2 (D), LA-RA-3 (B), LA-GR-1 (D). Table 43. — Measurements of Haustator perdita (Conrad). Abbre- viations as in Table 10. MD MH WN ANSP 13232 lectotype 12:3 41.3 13 ANSP 13232 paralectotype 10:5 35.1 10 ANSP 13232 paralectotype 10.6 37% 10 USNM 353944 745 20.0 8 USNM 353945 Із 23.0 8 MS-CL-15 1323 45.0 14 MS-HN-3 8.6 230 9 MS-YZ-2 8.3 20.0 9 MS-CL-3 11.3 25.0 13 LA-GR-1 ІЛЕ 350 13 Туре Material.—lectotype and 2 paralectotypes of Т. perdita Conrad ANSP 13232; holotype of T. jack- sonensis Cooke USNM 353944; holotype of T. lowei Cooke USNM 353945. Remarks/Discussion. — Results of multivariate mor- phometric analysis on perdita sensu lato are presented in Text-figure 21. A total of 67 variables was measured on 68 specimens. The first three factor axes account for 60.1% of the total variance in the data set (Table 44). Loadings on the first three factors (Table 45) in- dicate that variables on the eighth, ninth and tenth whorls load most heavily on the first factor, whereas variables from the fifth and sixth whorls load most heavily on the second factor and variables on first and second whorls load most heavily on the third factor. It is clear from the distribution of specimens that there are no discrete groupings represented, but rather a con- tinuum of variation, especially along axes representing variation on later whorls. In light of this pattern, there appears to be little justification for recognizing more than a single distinct taxon within perdita. There is, however, some degree of differentiation within perdita in the Moody’s Branch Formation by locality. As in- dicated by the Measurements listed above, H. perdita from Clarke County is consistently larger than those from the Jackson area. Specimens from two localities in Clarke County in eastern Mississippi (Chickasawhay River and Garland Creek, MS-CL-15, MS-CL-3) show similar mean values but different amounts of variation, whereas specimens from 130 km to the east at Riv- erside Park in Jackson, in central Mississippi (MS-HN- 6) shows a different mean and much less variation. This variation may be due to environmental vari- ation within the Moodys Branch sea. As discussed by Dockery (1977), and Hansen and Elder (1981), there is heterogeneity among depositional environments represented in the Moodys Branch Formation across its outcrop. Some of this variation is attributable to facies differences across roughly synchronous shelf en- 76 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 T. perdita Dr A AA а 1.5 + E A A AA A 1 L| à ЈЕ А А - ж = A 1 А А ü 0.5 + a 5 А = © = af E | | Г п | Ge i H | n nd -3 -2.5 -2 -1.5 -1 -0.5 0.5 1 ellus 2 215 -0.5 + 5 x 5 x "| à A Ж Riverside Park x Xx z. А 5 Garland Creek xt х cx A x A A Chickasawhay River a Factor 1 Text-figure 21.—Results of factor analysis of specimens of Haustator perdita (Conrad), first and second factor axes. * represents specimens from Riverside Park in Jackson, Hinds County, MS (locality MS-HN-6). ІШ represents specimens from Garland Creek, Clarke County, MS (locality MS-CL-3). А represents specimens from the Chickasawhay River, Clarke County, MS (locality MS-CL-15). See Appendix 1 for specimens used and details of analysis. vironments, but some may also represent changes in environments through time. Н. perdita appears to re- flect this heterogeneity particularly well, showing a great variety of forms across its relatively restricted strati- graphic and geographic range. At the Town Creek lo- cality in Jackson, Mississippi (MS-HN-3) H. perdita is the most abundant turritellid (Dockery, 1977, pers. comm.), whereas less than 6 km away at Riverside Park (MS-HN-6) Palmerella alveata (Conrad) is most abundant (pers. obs.). Haustator rina (Palmer, 1937) Plate 10, Figures 1,4,5 Turritella mortoni Conrad. Tuomey, 1848, p.159; not Conrad, 1830. Turritella carinata I. Lea. Aldrich, 1886a in part, p. 46; not I. Lea, 1833. Table 44.—Results of factor analysis of specimens of Haustator perdita (Conrad). Variance explained by first five factor axes. cumulative proportion Factor of variance 0.3051 0.4639 0.6010 0.6748 0.7417 чл & ошо о к Turritella carinata var. praecarinata Harris. Cooke, 1936, p.63, no- men nudum; not T. praecarinata Douvillé, 1904. Turritella rina Palmer, 1937, p.192, pl.22, £.3,4,9; in Harris and Palmer, 1947, p.296; Bowles, 1939, p.277, pl.31, f.17; Stenzel and Turner, 1942, card 98; Brann and Kent, 1960, p.933,934; Glibert, 1962, p.99; Palmer and Brann, 1966, p.1003; Toulmin, 1977, p.305, р1.51, £.9; Dockery, 1980, p.79, р1.29, f.2,3,5,7. Turritella rina carolina Palmer, 1937, p.194, pl.22, f.6; in Harris and Palmer, 1947, p.296; Bowles, 1939, p.280; Stenzel and Turner, 1942, card 53; Brann and Kent, 1960, p.934,935; Glibert, 1962, p.99; Palmer and Brann, 1966, p.1003; Allison and Adegoke, 1969, p.1257. Turritella wechesensis Bowles, 1939, p.281, pl.31, f.8,14; Stenzel and Turner,1942, card 108. Turritella rina wechesensis Bowles. Palmer in Harris and Palmer, 1947, p.296; Palmer and Brann, 1966, p.1004. Turritella rina rina Palmer. Allison and Adegoke, 1969, p.1256. Turritella rina Palmer var. Dockery, 1980, p.79, pl.2, f.1. Description. —Shell medium sized to large, of 15 to 20 whorls. Maximum observed whorl diameter 21.2 mm. Apical angle 18°; pleural angle usually greater than apical angle. Sutures deeply incised. Protoconch sub- erect, homeostrophic, turbinate, of approximately 1.75- 2.0 smooth, rounded, somewhat flattened and inflated whorls; Pl small. Apical sculpture formula C, B, A;. Primary spiral C appearing as faint angulation just below middle of whorl, followed within 0.5 whorl by B spiral just above C. B and C quickly diverging and PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 77 Table 45.—Results of factor analysis of specimens of Haustator perdita (Conrad). Rotated factor loadings of each variable on each of the first three factor axes. Variables as indicated in Text-figure 3. Variable factor 1 factor 2 factor 3 Variable factor 1 factor 2 factor 3 WHI —0.005 —0.089 0.794 W2-6 0.788 0.004 —0.146 SWI —0.005 —0.089 0.794 W3-6 0.784 01052 =0.151 WI-1 —0.005 —0.089 0.794 CH6 01775 —0.091 —0.150 W2-1 —0.005 —0.089 0.794 WH7 0.369 01050 —0.689 W3-1 —0.005 —0.089 0.794 SW7 0.087 —0.288 -051 CH1 —0.005 —0.089 0.794 W1-7 0.411 0.236 =0.632 WH2 0.335 —0.061 0.684 W2-7 0.417 0.184 — 0.658 sw2 0.334 2:00:09 0.615 W3-7 0.412 0.099 =0.672 W1-2 0.335 —0.059 0.689 CH7 0.392 0.077 —0.670 W2-2 0.341 —0.050 0.676 WHS8 0.038 0.374 =0,101 W3-2 0.338 —0.056 0.682 SW8 —0.245 0.518 —0.236 CH2 0.339 20:059 0.680 W1-8 EID 0.749 —0.215 WH3 0.648 20:120 0.459 W2-8 OMS 0.742 0227 SW3 0.606 007] 0.363 W3-8 20:129 0:727 =0.229 W1-3 0.674 —0.053 0.448 CH8 —0.130 0.701 0,225 W2-3 0.667 —0.083 0.441 WH9 —0.091 0.871 0439 W3-3 0.657 —0.101 0.447 SW9 —0.076 0.810 —0.148 CH3 0.659 —0.098 0.444 W1-9 —0.099 0.886 —0.128 WH4 —0.049 —0.160 —0.082 W2-9 2:0:095 0.881 —0.134 SW4 0.617 —0.381 0.078 W3-9 —0.106 0.876 041189) W1-4 0.742 0228 0.195 CH9 —0.085 0.852 0.125 W2-4 0.732 —0.260 0.176 WH10 0 0.851 0.043 W3-4 0.730 —0.285 0.185 SW10 —0.025 0.803 0.028 CH4 0.724 —0.300 0.175 W1-10 —0.145 0.846 0.048 WHS 0.814 —0.238 —0.006 W2-10 04187, 0.849 0.043 SW5 0.699 0820 —0.105 W3-10 03435 0.849 0.044 W1-5 0.838 —0.090 0.000 CH10 70.4107 0.836 0.048 W2-5 0.835 —0.124 —0.009 WH11 —0.324 0:525 0.035 W3-5 0.832 —0.160 —0.010 SWII —0.286 0.507 0.029 CHS 0.828 —0.158 0.005 WI-11 =0:335 0.521 0.040 WH6 0.760 —0.160 081 W2-11 —0.336 0.521 0.040 SW6 0.592 —0.145 —0.164 W3-11 0:385 0.523 0.040 W1-6 0.786 0.064 E9197 B quickly becoming as strong or stronger than C. Pri- mary A appearing on upper !^ of whorl within 0.25 whorl after B, but not reaching strength equal to C for three to four whorls by which point B becoming weak- er. B rapidly becoming obsolete and disappearing by whorl 8—9. If present, secondaries very faint. Spirals A and C persisting at moderate strength throughout on- togeny, area between becoming concave. Whorl profile below C spiral and above suture usually relatively Straight-sided leading to prominent D spiral. By about whorl 15, point of adapical contact between succeeding whorls beginning to move inward; D spiral becoming Sharp and prominent basal carina. Lateral aspect of growth line trace slightly prosocline, usually with upper and lower inflection points and moderately deep lateral Sinus with apex just above whorl middle. Spiral and basal sinuses moderately deep. Measurements.—See Table 46. Stratigraphic and Geographic Distribution. —H. rina s.s.: AL: Lower and Upper Lisbon Formation; MS, LA: Cook Mountain Formation; SC, GA: McBean Formation’; H. rina carolina: SC: McBean Formation; H. rina wechesensis: TX: Weches Formation; Clai- bornian Stage, Middle Eocene. Type Localities. — T. rina s.s.: Claiborne Bluff, Ala- bama River, Monroe County, Alabama (AL-MO- 15b). T. rina carolina: Orangeburg County, South Carolina (SC-OR- 1). T. rina wechesensis: well at Percilla, Hous- ton County, Texas (TX-HO-13). ; Other Localities. — T. rina в.в: AL-MO-4 (B), AL- CL-13 (B), AL-CN-1 (B), MS-CL-1 (B), MS-CL-2 (B), MS-CL-16 (B), MS-CL-17 (B), MS-NE-1 (B), MS-NE-9 Table 46.— Measurements of Haustator rina (Palmer). Abbrevi- ations as in Table 10. MD MH WN PRI 2874 holotype 18.0 47.0 9.5 PRI 2868 paratype 10.0 25.0 755 MCZIP 29348 50.1 IL 12.0 8 See Footnote 3, p.46. 78 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 (B), MS-NE-10 (B), MS-NE-11 (B), LA-WI-1 (B), GA- BU-3 (B), SC-AI-1 (B), SC-LE-1 (В), SC-WI-1 (В); 7. r. wechesensis: TX-RO-12 (B), TX-BA-3 (PB). Type Material.—holotype of T. rina Palmer PRI 2874, paratypes PRI 2868, 2869; holotype of T. rina carolina Palmer PRI 2872; holotype of 7. rina we- chesensis USNM 497958 paratype USNM 497959. Remarks/Discussion. — As already discussed, Haus- tator rina and H. carinata are very similar, both show- ing basal carination by expansion of the D spiral. Al- though it shows its first occurrence later than H. rina, H. carinata probably diverged independently from a common ancestor with rina, rather than from rina it- self; it seems unlikely that a lineage would sharply reduce its B spiral, as rina and its close relatives do, and then give rise to a descendant with a better de- veloped B, such as carinata. Haustator rivurbana (Cooke, 1926) Plate 10, Figure 3 Turritella rivurbana Cooke, 1926, p. 136, f.10; Bowles, 1939, p.283; Stenzel and Turner, 1942, card 99; Palmer in Harris and Palmer, 1947, p.295, pl.38, f.6,7,9, f.8, copy Cooke, 1926; Brann and Kent, 1960, p.935; Palmer and Brann, 1966, p.1004; Dockery, 1977, p.45-6, pl.3, f.6. Turritella rivurbana rivurbana Cooke. Allison and Adegoke, 1969, p.1257-8. Turritella rivurbana chiapasensis Allison in Allison and Adegoke, 1969, p.1258-9, pl.147, f.3,5,6,8-12,14, pl.148, f.2,11. Turritella rivurbana mexicana Allison in Allison and Adegoke, 1969, р.1259-62, р1.147, £.4,13, pl.148, f.1,3,4,6,7,9,10,13,14. Description. — Shell small to medium sized, of about 15 whorls. Maximum observed diameter approxi- mately 8.6 mm. Apical angle approximately 20°; pleu- ral angle approximately equal. Sutures moderately strongly incised. Protoconch and earliest teleoconch whorls unknown. Earliest known whorls showing A and C spirals with C spiral stronger. B spiral often present as thin thread, slightly closer to C than A. Sulcus between A and C spirals becoming broader with age. A and C staying very sharp and narrow, C always slightly stronger. Weak D spiral on later whorls form- ing a slight basal carination. Measurements. —See Table 47. Stratigraphic and Geographic Distribution. — MS: Moodys Branch Formation; lower Jacksonian Stage, Upper Eocene. Table 47.—Measurements of Haustator rivurbana (Cooke). Ab- breviations as in Table 10. MD MH WN USNM 353946 holotype 8.0 1755 4.5 unnumbered MGS specimen 8.6 30.3 16.5 Type Locality. — Town Creek, Jackson, Hinds Coun- ty, Mississippi (MS-HN-3). Other Localities. — MS-HN-6 (D), MS-YZ-1 (В), MS- YZ-6 (B). Type Material.—holotype of T. rivurbana Cooke USNM 353946; holotype of T. rivurbana chiapacensis Allison UCMP 12361, paratypes UCMP 30099, 30100, 31300, 32108, 32161, 32403, 32404, 32601, 32604, 32634; holotype of 7. rivurbana mexicana Allison UCMP 30091, paratypes UCMP 32638, 32639, 32645, 32646, 32648, 32650. 32652, 32657, 32691, 32699, 32757, 327799. Remarks/Discussion. — H. rivurbana is rare in the Moodys Branch Formation (pers. obs.; D.T.Dockery, pers. comm.). Allison and Adegoke (1969) have sug- gested that the Moodys is not the first occurrence of the species and have described a subspecies (Н. r. mex- icana) of “latest middle Eocene age” from near Si- mojovel, Chiapas, Mexico. These authors also, how- ever, reiterate the suggestion of Palmer (in Harris and Palmer, 1947, p.296) that rivurbana is so similar to H. rina s.s., differing principally in size, that it may be conspecific with it, in which case the Mexican form would be a form of H. rina of approximately the same age. H. rivurbana is more similar to H. martinensis (Dall) than is H. fischeri (Palmer), and may be an in- termediate form between Claibornian rina group spe- cies and the Upper Eocene to Recent genus Torcula Gray (cf., Allison and Adegoke, 1969). Haustator subrina (Palmer, 1937) Plate 10, Figure 2 Turritella eurynome Whitfield. Aldrich, 1894, p.233, not p.237; not Whitfield, 1865. Turritella rina subrina Palmer, 1937, p.194, pl.22, f.1,2,5,7,8,10,11; Bowles, 1939, p.279, рі.31, f.16; Stenzel and Turner, 1942, card 103; Harris and Palmer, 1947, p.296; Brann and Kent, 1960, p. 934,935; Glibert, 1962, p.99; Palmer and Brann, 1966, p.1003, as sabrina [sic]; Allison and Adegoke, 1969, p.1257; Toulmin, 1977, p.305, pl.51, f.6. Description. — Shell medium sized to large, probably of 15 to 20 whorls. Maximum observed whorl diameter 20.0 mm. Apical angle 23°; pleural angle usually slight- ly less than apical angle. Protoconch and earliest apical whorls unknown. Early ontogeny of spiral sculpture resembling that of Haustator rina (Palmer), but pri- mary spirals A and C becoming more prominent more quickly; B often weakening and becoming obsolete ear- lier. Although C stronger than A on earlier whorls, A becoming stronger on later whorls, often expanding to form adapically flaring carina on upper Уз of whorl. Secondary spiral R becoming moderately prominent just below suture on late whorls, as does D, although not to degree shown in H. rina. Fine secondary visible n PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 79 Table 48.— Measurements of Haustator subrina (Palmer). Abbre- viations as in Table 10. Table 49.— Measurements of Haustator subtilis (Kellum). Abbre- viations as in Table 10. MD MH WN MD MH WN PRI 2866 syntype 15.0 17.0 2.0 USNM 353240 holotype 7.0 24.0 10.0 PRI 2867 syntype 10.0 10.0 2.0 PRI 2870 syntype 22.0) 68.0 6.0 PRI 2871 syntype 12.0 33.0 8.0 PRI 2875 t 14.0 41.0 8.0 T қ : USNM ipti) б " 14.0 42.0 15.0 spirals with C strongest and A and B of approximately MCZIP 29304 20.9 58.8 8.5 equal strength. B spiral weakening to a fine thread and below A spiral on late whorls. Growth line trace as in H. rina, except that lateral sinus may be somewhat deeper. Measurements. —See Table 48. Stratigraphic and Geographic Distribution. — AL: Upper and Lower Lisbon Formation; TX, MS: Cook Mountain Formation; SC: McBean Formation’; mid- dle—upper Claibornian stage, Middle Eocene. Type Locality.—1 mile S of Lisbon Landing, Ala- bama River, Monroe County, Alabama (AL-MO-4d). Other Localities. — AL-MO-1b (B), AL-CL-13 (B), TX-HO-2 (B), TX-LE-12 (B), TX-SA-3 (PB), MS-NE-2 (PB), MS-CL-1 (PB), SC-OR-1 (PB), SC-OR-3 (PB), SC-OR-7 (PB). Type Material. —syntypes of T. rina subrina Palmer PRI 2866, 2867, 2870, 2871, 2873, 2875. Remarks/Discussion. — Although Palmer designated subrina as a subspecies of Haustator rina, she described it as morphologically distinct. This highly sculptured form in fact co-occurs with H. rina in the Lisbon and Cook Mountain formations without morphological in- termediates. It is clearly closely related to rina, but its extreme development of primaries A and C, without the basal carination and development of D, distinguish it. The lack of intermediates between distinct co-oc- Curring morphotypes argues for recognition of two sep- arate species-level taxa. Haustator subtilis (Kellum, 1926) Plate 15, Figure 4 Turritella subtilis Kellum, 1926, p.27, pl.5, £5; not Stephenson, 1927, p.21 [= T. kellumi Stephenson, 1939, fide Palmer and Brann, 1966]; Bowles, 1939, p.283; Stenzeland Turner, 1942, card 104, f.5; Palmer and Brann, 1966, p.1005. Description. —Shell small to medium sized, of ap- proximately 15 whorls. Apical angle approximately 15°; pleural angle approximately equal. Protoconch and earliest teleoconch whorls unknown. Sutures slightly incised. Earliest known whorls showing A,B and C ? See Footnote 3, p.46. whorl becoming cingulate by whorl 6-8. Adult whorls not markedly carinate. Second, equally fine thread joining the B spiral on later whorls; two approximately equally spaced between the A and C spirals. Measurements. —See Table 49. Stratigraphic and Geographic Distribution. —NC: Castle Hayne Formation; Middle Eocene. Type Locality. —city rock quarry near Smith Creek, E side of Wilmington, New Hanover County, North Carolina. Other Localities. —known only from the type local- ity. Type Material.—holotype of T. subtilis Kellum USNM 353240. Remarks/Discussion. — This form is known from a single specimen, an external mold in glauconitic marl. Bowles (1939, p.283) suggests that 1t 1s closely related to Т. martinensis Dall and to T. rivurbana Cooke, both of which are herein placed 1n Haustator, differing from these species in being “more slender and more grad- ually tapering. . ." and having whorls “marked by more prominent secondary lirae.” Allison and Adegoke (1969) agree, and tentatively include it in their rina group. The holotype of subtilis differs from martinensis in having A and C spirals narrower, sharper and less pronounced, and the sulcus is accordingly shallower. The B spiral and faint secondaries in the sulcus are stronger than in martinensis. Haustator tennesseensis (Gabb, 1860) Plate 7, Figures 6-9 Turritella tennesseensis Gabb, 1860, p.392, р1.68, f.13; Harris, 1896, p.108, pl.11, £5; Aldrich, 1921, p.25; Bowles, 1939, p.284, pl.32, f.12; Stenzel and Turner, 1942, card 105, #5; Brann and Kent, 1960, p.938, Palmer and Brann, 1966, p.1005. Turritella sp., Lowe, 1933, p.9. Turritella mortoni Conrad. Lowe, 1933, p.10 in part, stations 6496 and 6497 only; not Conrad, 1830. Description. —Shell small to medium sized, of 15 to 20 whorls. Maximum observed whorl diameter 9.5 mm. Apical angle 17°; apical and pleural angles ap- proximately equal. Protoconch suberect, homeos- trophic, turbinate but flattened, of about 2.5 smooth, rounded whorls; P1 small. Protoconch and earliest te- leoconch whorls unusually tall, almost columnar, with 80 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 50.— Measurements of Haustator tennesseensis (Gabb). Ab- breviations as in Table 10. Table 51.—Measurements of Haustator vaughani (Bowles). Ab- breviations as in Table 10. MD MH WN MD MH WN USNM 497989 hypotype 9.5 25.5 7.0 USNM 497955 holotype 252) 10.1 Rs: very small apical angle, increasing on around second or third teleoconch whorl. Apical sculpture formula C, В, А». Primary spirals B and C appearing approxi- mately simultaneously on lower У; of whorl, С as rel- atively sharp angulation just above suture and B as faint prominence just below whorl middle. B quickly becoming stronger, forming subcentral keel for several whorls. Primary A spiral appearing as faint ridge in middle of upper У of whorl. B soon surpassed by C as whorl profile changes to trapezoidal, widest on the bot- tom, then to hourglass-shaped as whorl middle con- stricts. Profile of latest whorls markedly concave, still slightly wider at the bottom. Spiral ribs greatly reduced in strength. Lateral aspect of growth line trace slightly prosocline, lacking well developed inflection points. Lateral sinus only of moderate depth. Basal aspect of growth line trace not observed. Measurements. —See Table 50. Stratigraphic and Geographic Distribution. —TN, MS: Clayton Formation; lower Midwayan Stage, Lower Pa- leocene. Type Locality. —“ Нагдетап County, Tenn., marls of the Ripley Group" (Gabb, 1860). Other Localities. —TN-HA-7a (B), TN-HA-7b (В), MS-TI-8 (В), MS-TI-9 (B), MS-TI-10 (B). Type Material. —holotype of 7. tennesseensis Gabb unknown (fide Bowles, 1939; Palmer and Brann, 1966, p.1005), hypotype (Bowles, 1939) USNM 497989. Remarks/Discussion. — This species is allied with Palmerella dutexata (Harris) and P. lisbonensis (Bowles) in the “bicostate group" by Bowles (1939). Examina- tion of early apical sculpture by SEM, however, sug- gests that its affinities lie with the “7. rina group" and that it should be placed in Haustator. Haustator vaughani (Bowles, 1939) Plate 15, Figure 5 Turritella vaughani Bowles, 1939, p.282, pl.32, £.14; Stenzel and Turner, 1942, card 107, f.14; Palmer and Brann, 1966, p.1006. Description. —Shell very small, of 15-18? whorls. Largest observed whorl diameter 2.2 mm. Protoconch and earliest teleoconch whorls unknown. Apical angle approximately 10*; pleural angle approximately equal. Shell elongate, narrow and straight-sided. Earliest known whorls bearing only subequal, rounded A and C spirals, with no trace of B spiral. A becoming slightly stronger on later whorls; otherwise whorls gently curved, cingulate in profile. Very faint trace of one to two threads above A, at least one below C on later whorls. Sutures only slightly incised. Growth lines not visible. Measurements. —See Table 51. Stratigraphic and Geographic Distribution. — SC: McBean Formation!°; upper Claibornian Stage, Mid- dle Eocene. Type Locality. —Poosers Hill, 5.1 mi N of Orange- burg, Orangeburg, South Carolina (SC-OR-6). Other Localities. —known only from the type local- ity. Type Material. —holotype of Т. vaughani Bowles USNM 497955. Remarks/Discussion. — Commenting on the phylo- genetic position of this species, Allison and Adegoke write: “Тһе relationships of Turritella vaughani are in doubt. This species is known from only two specimens, the holotype and a topotype. Allison examined them and is of the opinion that they represent only portions of the juvenile whorls. The pleural angle of this species is extrmely narrow; the earliest whorls bear A and C . with no sign of other ribs, although the earliest apical whorls and protoconch are missing. This species may have had an early tricostate or near tricostate stage, in which case it would belong to the Turritella rina group, but in the absence of the early apical whorls it is im- possible to evaluate its relationships. It seems doubtful, however, that it belongs to Torcula” (1969, p.1262). Genus 2 (“Turritella humerosa group") Remarks/Discussion. — Of all the species considered in detail in this paper, the least is known about those comprising the humerosa group. I have been able to find well preserved apices only for Turritella aldrichi Bowles, and sample size of several other species (7. biboraensis Gardner, T. claytonensis Bowles, T. gard- nerae LeBlanc) are very small or preservation is poor (T. toulmini, new species). These species are united only by their distinctive whorl profile, a character be- lieved to be prone to homoplasy, and so might not comprise a monophyletic group. Phylogenetic conclu- sions about these species are therefore especially dif- ficult. The origins of the four larger Late Paleocene 10 See Footnote 3, p.46. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 81 species is the biggest mystery. As discussed in more detail in the descriptions below, T. humerosa differs from multilira, praecincta and eurynome in having a broad, rounded carina made up of several spiral ribs, whereas the carinae of the others is sharper and com- posed of only the expanded R secondary spiral rib. T. humerosa occurs only on the Atlantic coast, 7. mul- шта Whitfield and 7. eurynome Aldrich only on the Gulf coast, and T. praecincta Conrad in both areas. In Maryland the Aquia Formation, containing prae- cincta and humerosa, is unconformably underlain by the Brightseat Formation containing the very similar T. “prehumerosa” Govoni (1983; Govoni and Hansen, in press). On the Gulf coast, the time represented by the unconformity between the Brightseat and Aquia appears to be represented by the highly fossiliferous Matthews Landing Marl Member of the Porters Creek Formation. This unit contains no humerosa-like forms other than T. aldrichi. It is possible that humerosa (and praecincta?) evolved from “prehumerosa” on the At- lantic coast, whereas eurynome and multilira (and praecincta?) arose from aldrichi on the Gulf coast. Al- ternatively, all three Gulf coast forms could have been derived from Atlantic coast ancestors, and aldrichi could have been a sterile branch, related or unrelated to the others. Given the data presently available, all that can be done at this point is to put forth these alternatives in the hope of future testing. “Turritella” aldrichi Bowles, 1939 Plate 12, Figures 1—4 Turritella humerosa ? Conrad. Aldrich, 1886a, p.59; not Conrad, 1835b, p.340. Turritella multilira Whitfield. Aldrich, 1886a, p.59; not Whitfield, 1865, p.266. Turritella humerosa var. multilira Whitfield. Aldrich, 1894, p.246, pl.13, £.3 as multileia [sic]; Harris, 1896, p.111, pl.11, £.10, copy Aldrich, 1894. Turritella humerosa Conrad var. Aldrich, 1894, p.246, pl. 13, f.5. Turritella humerosa Conrad. Harris, 1896, p.111, in part, pl.11, f.11, copy Aldrich, 1894; Veatch and Stephenson, 1911, p.217, 219,223,224; Stephenson and Veatch, 1915, p.68-69; Brantly, 1920, p.145-146; not Conrad, 18355. Turritella aldrichi Bowles, 1939, p.315, pl.34,f.11,17; LeBlanc in Barry and LeBlanc, 1942, p.96, pl.13, f.3,4; Stenzel and Turner, 1942, card 42; Palmer and Brann, 1966, р.979–80; Toulmin, 1969, pl. 1. £13: 1977; »0732 518, LLO, Description. —Shell medium sized, of 25 to 30 whorls. Maximum observed whorl diameter 11.1 mm. Apical angle 14°; apical and pleural angles approximately equal. Sutures inconspicuous on early whorls, more deeply incised on later whorls. Protoconch erect, homeos- trophic, turbinate, of 1.0—1.5 smooth, rounded, some- what flattened whorls; P1 small. Apical sculpture for- mula C, B, A,. Primary C spiral appearing abruptly as distinct, raised band just below whorl middle, mov- ing downward to middle of lower ^ of whorl. Primary B appearing within one whorl as less distinct promi- nence in middle of upper Y. of whorl, moving down- ward to just above middle of whorl. Primary A ap- pearing as similarly faint prominence just below upper suture two to three whorls later, moving downward to middle of upper % of whorl. Whorl profile becoming basally carinate with appearance of B spiral; area above C spiral relatively straight-sided, area below to suture sharply concave. Profile changing at about whorl 12- 15 with appearance of secondary spiral (R) above A, R and A both increasing in size to form adapical carina on later whorls. Secondary spirals appearing over rest of whorl and strengthening to equal primaries B and C. Lower 73 of whorl swelling to become roundly con- vex; area just below adapical carina remaining some- what constricted. Narrow band above lower suture be- coming excavated and concave. Primary and second- ary spirals on late whorls usually faintly to moderately beaded by intersection with growth lines. Lateral as- pect of growth line trace slightly prosocline, with upper and lower inflection points and moderately deep sinus, apex just above whorl middle. Spiral and basal sinuses moderately deep. Measurements. —See Table 51. Stratigraphic and Geographic Distribution. — AL: Clayton Formation, Porters Creek Formation (Lower Member), Porters Creek Formation (Matthews Land- ing Member), Naheola Formation; GA: undifferen- tiated Midway Group; LA: Lime Hill (“Logansport”) Formation; Midwayan Stage, Lower Paleocene. Type Locality. — Matthews Landing, Alabama Riv- er, Wilcox County, Alabama (AL-WI-3). Other Localities. — AL-WI-2 (T), AL-WI-3 (B), AL- WI-9 (B), AL-WI-21 (B), AL-WI-36 (B), AL-WI-37 (B), AL-WI-42 (T), AL-WI-43 (T), AL-WI-44 (T), AL- WI-45 (T), AL-WI-46 (T), AL-WI-47 (T), AL-WI-48 (T), AL-WI-50 (T), AL-WI-51 (T), AL-WI-52 (T), AL- WI-53 (T), AL-BU-1 (T), AL-BU-2 (T), AL-BU-3 (T), AL-MA-2 (T), LA-NA-6 (LB), GA-WE-1 (B), GA-SL-1 (B). Type Material.—holotype of 7. aldrichi Bowles USNM 495148, paratype USNM 495149. Remarks/Discussion. — This is the only member of Table 52.—Measurements of “Turritella” aldrichi Bowles. Ab- breviations as in Table 10. MD MH WN USNM 495148 holotype 9.0 39.0 13:5 USNM 495149 paratype 10.0 41.0 9.5 MCZIP 29305 7.0 38.9 24.5 the humerosa group for which I have observed well preserved apices. “T.” aldrichi shares a common apical sculpture formula with members of the rina group (Haustator herein), but is distinguishable from the most similar member of that group, Haustator alabamiensis (Whitfield), by its very different spiral sculpture and whorl shape on early teleoconch whorls, and by distinct adapically carinate adult whorls in most individuals. As mentioned above under H. alabamiensis, however (p.67), some specimens of both species may be difficult to distinguish. Whether aldrichi is closely related to any later mem- bers of the humerosa group is not clear. It differs from most of these species, as well as from most of the other adapically carinate Midwayan species (“Т.” clayto- nensis Bowles, “Т.” “prehumerosa” Govoni, “T.” toulmini n.sp., “T.” biboraensis Gardner) in being very small, delicate, narrow and having more than 20 whorls. It appears to persist virtually unchanged throughout much of the Midwayan Stage (perhaps as long as 7.0 my), and is perhaps best viewed as a more derived side branch than a generalized ancestral form. At Matthews Landing on the Alabama River, ald- richi and alabamiensis are two of the most common mollusks, but in museum lots the two species occur in very different relative abundances. The exposures at Matthews Landing consist of continuously weathering horizontal bedding planes, and the differing compo- sition of samples from here may suggest some mi- croenvironmental heterogeneity of the original com- munity. “Turritella” biboraensis Gardner, 1935 Plate 14, Figure 5 Turritella biboraensis Gardner, 1935, p.290, pl.25, f.3; Stenzel and Turner, 1942, card 47. Turritella sp. cf. T. humerosa Conrad. Gardner, 1935, p.289, pl.25, De Turritella humerosa biboraensis Gardner. Bowles, 1939, p.313; Palmer and Brann, 1966, p.991. Description. — Available material very poorly pre- served. Shell medium sized, total whorl number un- known. Maximum observed whorl diamter 15.9 mm. Apical. angle approximately 1097, approximately equal to pleural angle? Sutures moderately deeply incised. Protoconch and earliest teleoconch whorls unknown; earliest known whorls show pronounced, square adap- ical carina immediately below suture. Whorls straight- sided, marked with numerous, fine secondary spirals of more or less equal strength. May be slightly stronger spiral cord (C spiral?) at base of earliest whorls in ho- lotype. Growth lines not visible on holotype. Measurements. —See Table 53. Stratigraphic and Geographic Distribution. — TX: PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 53. — Measurements of “Turritella” biboraensis (Gardner). Abbreviations as in Table 10. MD MH WN USNM 370989 holotype 15:9 42.4 5 Kincaid Formation; lower Midwayan Stage, Lower Pa- leocene. Type Locality. — Bibora Tank, Indio Ranch, 18 mi SE of Eagle Pass, Maverick County, Texas (TX-MV- 1). Other Localities. -TX-LI-2a (B), TX-MV-4b (B), TX-MV-6 (B). Type Material. —holotype of T. biboraensis Gardner USNM 370989. Remarks/Discussion. —I have examined only the ho- lotype of this species. It shares with “T.” humerosa Conrad a round, broad adapical carina and relatively straight whorl profile. It is distinguished chiefly on the basis of the substantial difference in the age of the two. Study of further material, particular well preserved api- ces, may show that such a taxonomic separation is not justified. “Turritella” claytonensis Bowles, 1939 Plate 12, Figure 8 Turritella humerosa Conrad. Harris, 1896, p.110 in part, pl.11, f.12 only; not Conrad, 1835b. Turritella claytonensis Bowles, 1939, p.314, р1.34, f.14,15; Stenzel and Turner,1942, card 55; Palmer and Brann, 1966, p.984. Description. —Shell large, of perhaps 10-15 whorls? Maximum observed whorl diameter 26.0 mm. Apical angle approximately 15?, approximately equal to pleu- ral angle. Protoconch and earliest teleoconch whorls unknown. Earliest known whorls show more or less uniform, fine spiral lines over entire whorl. Adapical carina develops on later whorls, probably from a sec- ondary above the A spiral. Carina relatively narrow, subrounded, not more than 1596 of whorl height; all other spirals faint. Sutures not incised, but perched immediately above carina. Basal portion of whorl im- mediately above suture slightly inflated. Lateral aspect ofgrowth line trace approximately orthocline, with two inflection points, one above and one below whorl mid- point; lateral sinus of moderate depth. Basal aspect of growth line trace not observed. Measurements. —See Table 54. Table 54.— Measurements of “Turritella” claytonensis Bowles. Abbreviations as in Table 10. MD MH WN USNM 131637 holotype 26.0 68.0 Hie PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 83 Stratigraphic and Geographic Distribution. — AL: Clayton Formation; lower Midwayan Stage, Lower Pa- leocene. Type Locality. —Prairie Creek, Wilcox County, АЈ- abama (AL-WI-9). Other Localities. — AL-BA-1 (B). Type Material. —holotype of T. claytonensis Bowles USNM 131637. Remarks/Discussion. —The USNM collections ap- pear to contain only two specimens of this form, one (the holotype) from Prairie Creek in western Alabama, and the other from the type section of the Clayton Formation in eastern Alabama. I have found turritel- lids to be common at the Clayton type locality, but to consist entirely of the apparently undescribed species here named “Turritella” toulmini; I have not found any other specimens of “T.” claytonensis. The short Spire and large apical angle make “Т.” claytonensis a very distinct form not easily confused with others; its rarity may therefore be real. “Turritella” eurynome Whitfield, 1865 Plate 13, Figures 6-8 Turritella eurynome Whitfield, 1865, p.266; Aldrich, 1887, p.81; 1894, p.237, not p.233 [= Haustator subrina (Palmer)] de- Gregorio, 1890, p.127; Harris, 1899a, p.75, pl.10, £7; Bowles, 1939, p.292, pl.31, £.13, pl.32, £.18; LeBlanc in Barry and LeBlanc, 1942, p.97, pl.13, £.5; Stenzel and Turner, 1942, card 62; Palmer and Brann, 1966, p.987; Toulmin, 1977, p.231, pl.29, f.6. Turritella bellifera Aldrich, 1885, p.150, pl.3, f.13; 1886a in part, p.34,55, pl.1, f.13, not 58; 1894 in part, p.237, not p.130,233,238; Smith and Johnson, 1887, p.29; not Clark, 1891, p.63; Cossmann, 1893, p.29; not Brantly, 1920, p.161; Plummer, 1933, p.815, pl.10, f.7; Bowles, 1939, p.317, pl.34, f.16; Stenzel and Turner, 1942, card 46; Palmer and Brann, 1966, p.982; Toulmin, 1977, p.230- 1. pL29 £5: Turritella (Proto) cathedralis bellifera Aldrich. de Gregorio, 1890, p.127, pl.11,f.17, copy Aldrich, 1885. Turritella humerosa Conrad. Harris, 1899a, p.75, pl.10, Ғ.5; 1899b, p.308, pl. 55, f.5; Brann and Kent, 1960, p.920; not Conrad, 1835b. Turritella (Haustator) bellifera Aldrich. Cossmann, 1912, p.118. Turritella sp., Plummer, 1933, p.815, pl.10, f.9,9a. Turritella humerosa Conrad “var.” Brann and Kent, 1960, p.920. Description. — Shell large, of perhaps 15 to 20 whorls. Maximum observed whorl diameter 20.0 mm. Apical angle 15°, pleural angle 7°. Sutures inconspicuous. Pro- toconch and earliest teleoconch whorls unknown. Ear- liest whorls observed with three moderately promi- nent, approximately equally spaced spirals, the lower two soon becoming stronger as a fourth appears above the upper spiral, closer to the suture than the spiral. This uppermost spiral becoming most prominent on later whorls of most individuals, forming an adapical carina. Four spirals sharp and distinct on later whorls. Secondaries faint if present. Growth lines often con- Table 55.—Measurements of “Turritella” eurynome Whitfield. Abbreviations as in Table 10. MD MH WN FMNH-UC 24505 syntype TT 2153 3.0 USNM 644614 17.5 85.0 из USNM 498001 11.5 52.0 12:5 spicuous but spirals seldom beaded or scalloped. Whorl profile generally straight-sided until latest whorls when lower 5›—%% of whorl becomes slightly rounded and con- vex. Lateral aspect of growth line trace slightly pro- socline with well developed upper and lower inflection points. Lateral sinus moderately deep, apex above whorl midpoint. Spiral and basal sinuses of only moderate depth. Measurements. —See Table 55. Stratigraphic and Geographic Distribution. — AL: Tuscahoma Formation (Bells Landing Marl Member), Tuscahoma Formation (Greggs Landing Marl Mem- ber) LA: Marthaville Formation, Pendleton Forma- tion; TX: Pendleton Formation; middle-upper Sabi- nian Stage, Upper Paleocene. Type Locality. — T. bellifera: Bells Landing, Ala- bama River, Monroe County, Alabama (AL-MO-3); T. eurynome: Bowles (1939) believed it to be Bells Landing (AL-MO-3); Stenzel and Turner (1942) be- lieved it to be Greggs Landing (AL-MO-5). Other Localities. — AL-CH-6 (B), AL-DA-1 (B), AL- MO-3 (B), AL-MO-5 (ST, T), AL-WI-18 (B), AL-CH-6 (B), LA-NA-14 (LB), LA-SA-19 (LB), LA-SA-20 (WW), LA-SA-21 (WW), LA-SA-22 (WW), TX-SA-4 (LB), TX-SA-19 (LB). Type Material.—lectotype of T. bellifera Aldrich USNM 644614; 3 syntypes of T. eurynome Whitfield FMNH-UC 24505. Remarks/Discussion. —Bowles (1939) states that apical whorls of “T.” eurynome are “marked by three prominent subequal revolving lirae ... a fourth lira arising just in front of the posterior suture very early in the shell growth . . . ," and that on adult whorls this fourth spiral occasionally “becomes strongly devel- oped and approximates the prominent carina of T. bellifera." Bowles also states that he did not observe the earliest apical whorls of bellifera, but that the ear- liest whorls known show “four distinct, subequal, re- volving lirae and two smaller, less distinct secondary lirae," and that three versus four spirals on the earliest whorls distinguished the two species. The variability on both early and late whorls of specimens fitting the descriptions of these two forms suggests, however, that this separation cannot be maintained, especially con- sidering the occurrence of both forms only in the Tus- 84 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 56.— Measurements of “Turritella” gardnerae LeBlanc. Ab- breviations as in Table 10. MD MH WN LSU 3145 holotype 11.0 28.0 10.5 cahoma Formation (Bells and Greggs Landing Mem- bers). A single, somewhat variable species is therefore recognized here. “Turritella” gardnerae Le Blanc in Barry and LeBlanc, 1942 Turritella gardnerae Le Blanc in Barry and LeBlanc 1942, p.94, pl.13, f.1,2; Palmer and Brann, 1966, p.988. Description. — Original description: “Spire probably high (incomplete holotype has only five whorls pre- served), tapering very gradually; apical angle 9 degrees. Whorls wider than high, marked by a subsutural band or collar and depressed areas immediately below the subsutural band above the suture. Sutures are linear, indistinct and impressed slighly on the upper part of the subsutural band. Sculptural consisting of very nu- merous subequal fine and coarse spiral lines becoming larger but not prominent, anteriorly; spirals on de- pressed area above sutures and on subsutural band more prominent than elsewhere. Incrementals are strongly reflexed at the posterior third of the whorl as in the members of Turritella humerosa group.” (LeBlanc, 1942, p.94) Protoconch and earliest teleo- conch whorls unknown. Measurements. —See Table 56. Stratigraphic and Geographic Distribution. — LA: undifferentiated Upper Midway Group; Midwayan Stage, Lower Paleocene. Type Locality. — Natchitoches Parish, Louisiana (LA- NA-6). Other Localities. —LA-SA-9 ? (LB). Type Material.—holotype T. gardnerae LeBlanc LSU 3145. Remarks/Discussion. — According to LeBlanc, this species is known from only two localities. I have been unable to locate the type locality, and the second lo- cality has been flooded by the waters of Toledo Bend Reservoir. “Turritella” gardnerae is most similar to * T." aldrichi Bowles, although its status remains un- certain without additional material. “Turritella” humerosa Conrad, 1835b Plate 13, Figures 1-4 Turritella humerosa Conrad, 1835b, p.340, pl.13, f.3; 1846, p.219; 1865a, p.32; 1866, p.11 H.C.Lea, 1849, p.107; not Aldrich, 1886, p.59 [= “T.”aldrichi Bowles]; Harris, 1894a, p.303 in part; not 1896, p.15,16,32, 36,110, pl.11, Ғ10-13; not 18992, p.75, pl.10, f.5-7; not 18995, p.308, pl.55, £5 [= “Т.” eurynome Aldrich]; Clark, 1896, p.70, pl.14, f.1; Clark and Martin, 1901, p.148, pl.27, f.1,1a; not Veatch and Stephenson, 1911, p.217,219,223,224 [= T. aldrichi Bowles]; Clark and Miller, 1912, p.92,94,95,119,120; not Stephenson and Veatch, 1915, p.68,69 [= 7. aldrichi Bowles]; Guillaume, 1924, p.290; Trowbridge, 1932, in part, pl.38, f.7, copy Clark and Martin, 1901; not pl.31, £.6,7; Gardner, 1935, p.288- 290; Bowles, 1939, p.312, pl.33, f.10; Stenzel and Turner, 1942, card 70; Vokes, 1961, p.49, pl.10, f.2. Turritella (Haustator) humerosa Conrad. Cossmann, 1912, p.118. Not Turritella humerosa Conrad “var.”. Cooke, 1926, pl.94, f.1 9 [= “T.” multilira Whitfield]; not Semmes, 1929, f.59-1, copy Cooke, 1926. Turritella (Peyrotia) humerosa Conrad. Shimer and Shrock, 1944, p.493, pl.201, f.18, copy Clark and Martin, 1901. Description. —Shell large to very large, of 20 to 25 whorls. Maximum observed whorl diameter 27.0 mm. Apical angle 17°; pleural angle often slightly less than apical angle. Protoconch unknown; earliest whorls ob- served (dia. = 0.34 mm) bearing two moderately pro- nounced spirals, probably corresponding to C and B, on lower Y. of whorl, moving slightly upward and strengthening until B just above whorl middle and C in about middle to lower У of whorl. Third spiral (= A?) appearing in middle of upper Y. of second whorl, but remaining weak. Lower two spirals subequal, C being slightly larger, forming moderate basal carina. Whorl profile above C more or less straight-sided ex- cept for spirals; profile below markedly concave to suture. Intercalaries appearing with whorl 10-12, larg- est whorls covered with numerous, evenly spaced, fine spirals. Whorl profile beginning to change well after introduction of secondary spirals, basal carina becom- ing less prominent, adapical / of whorl beneath up- permost two spirals expanding, area just below becom- ing relatively constricted. Adult whorls showing rounded adapical carina just beneath the suture, slight- ly curved whorl profile widest at about middle of lower V» of whorl. Spirals very seldom beaded although minor relief may be created by intersection of spirals and growth lines. Lateral aspect of growth line trace or- thocline to slightly prosocline, usually with upper and lower inflection points. Lateral sinus moderately deep, apex just above whorl middle. Spiral and basal sinuses relatively shallow. | Measurements. —See Table 57. Table 57. — Measurements of “Turritella” humerosa Conrad. Ab- breviations as in Table 10. MD MH WN ANSP 31388 lectotype 21.6 89.6 9.5 ANSP 31388 paratype 20.4 90.2 9.5 ANSP 31388 paratype 20.4 S9 4.0 ANSP 31388 paratype Ме 41.3 3.5 ANSP 31388 paratype 15.6 9799 4.0 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 85 Stratigraphic and Geographic Distribution.—MD, VA: Aquia Formation; Upper Paleocene. Type Locality. —probably Piscataway Creek, Prince Georges County, Maryland (MD-PG-5). Other Localities. — MD-AA-5 (B), MD-PG-7 (В), MD-PG-10 (B), VA-ST-2 (B), VA-ST-4 (B), VA-ST-5 (B), VA-ST-6 (B), УА-НА-2 (MCZ). Type Material. —lectotype and 4 paratypes of T. hu- merosa Conrad ANSP 31388. Remarks/Discussion. — As noted by Bowles (1939) the adapical carina so characteristic of humerosa group Species (and the principal character suggesting a close relationship among them) is somewhat different among the different species. In humerosa the carina is not formed by a single spiral rib, but by the swelling of a broad area just below the suture, on which the upper- most (presumably A) spiral is the most prominent. This carina may or may not be homologous with those of eurynome, multilira and praecincta, which form mainly from the expansion of a single spiral rib, spe- cifically the secondary above the uppermost primary (therefore presumably R). Although protoconchs and very earliest teleoconch whorls have not been preserved on the specimens available to me, it is almost certainly the case that “Т.” humerosa shows an apical sculpture formula of C, B, Аз, similar to that seen in members of the rina group and by “Т.” aldrichi Bowles. Spiller (1977) has noted that humerosa has three to four whorls, in its proto- conch. If this is true, and if it applies to at least the Other large Late Paleocene species with adapically car- inate whorls it would suggest 1) that aldrichi is not closely related to humerosa group; 2) that humerosa group species may have had relatively longer intervals in the plankton, greater powers of dispersal, and re- duced tendency for speciation, possibly explaining their lower species diversity compared to the basally cari- nate rina and mortoni groups which appear to have had only short planktonic larval stages; 3) many of the forms designated as “humerosa” from other areas of the New World may have been more closely related to Coastal Plain taxa than are, for example, forms as- Signed to Palmerella mortoni, which has also been used in a similar “catch-all” fashion, but which may have had lower powers of dispersal. In the Aquia Formation, humerosa occurs in the Same beds as P. mortoni, but is never as abundant. No ecological or environmental separation is detectable between the two forms. “Turritella” multilira Whitfield, 1865 Plate 12, Figure 5, 9-11 Turritella multilira Whitfield, 1865, p.266; not Aldrich, 1886a, p.59 [= “Т.” aldrichi Bowles]; not Aldrich, 1894, p.237,239 [= “Т.” aldrichi Bowles, not “T.” bellifera Aldrich as in Bowles, 1939, p.316, fide Palmer and Brann, 1966, p.997]; Whitfield, 1899, p.177; Harris, 1899a, p.75, р1.10, £.6 [as humerosa “уаг.”]; Bowles, 1939, p.316, pl.32, £.15, pl.34, £19; Stenzel and Turner, 1942, card 84; LeBlanc in Barry and LeBlanc, 1942, p.102, pl.13, f.9; Palmer and Brann, 1966, p.997; Toulmin, 1977, p.232, pl.29, ЕВ. Turritella bellifera Aldrich. Aldrich, 1886a in part, p.58; not, 1885. Not Turritella humerosa Conrad var. multilira Whitfield. Aldrich, 1894, p.246, р1.13, ЁЗ as multileia [sic] [= “Т.” aldrichi Bowles, not T. bellifera Aldrich as in Bowles, 1939, p.316]; Harris, 1896, p.111 in part, pl.11, f.10, copy Aldrich, 1894. Turritella multilirata [sic] Whitfield. Whitfield, 1899, p.177; not Adams and Reeve, 1848 (1850), p.48. Turritella humerosa Conrad “var.”. Cooke, 1926, р1.94, Ғ.1; Semmes, 1929, f.59-1, copy Cooke, 1926; Brann and Kent, 1960, p.920 in part, No. 272. Description. —Shell medium to large, of probably around 20 whorls. Maximum observed whorl diameter 20.0 mm. Apical angle 16°; pleural angle 9°. Sutures inconspicuous. Protoconch and earliest teleoconch whorls unknown. Earliest whorls observed (dia. = 1.9 mm) three prominent spirals of equal strength, ap- proximately equally spaced over whorl height. Whorl profile of these early whorls convex and evenly round- ed, maximum width in middle. Area between lowest spiral and suture excavated and smoothly concave. Weaker intercalaries beginning to appear between orig- inal spirals, whorl profile changing to more straight- sided, although area between each pair of stronger spi- rals slightly concave. Secondary spiral above upper original becoming increasingly prominent and forming sharp adapical carina on late whorls. Whorl profile of latest whorls more or less straight-sided, but with slight constriction just below carina and slight inflation at about whorl middle. Form and strength of carina vari- able; upper surface usually sloping upward to suture, but is less often horizontal from a more sharply apically bent carina. Carina rarely doubled by expansion of another more adapical spiral. Lateral aspect of growth line trace slightly prosocline, with upper and lower inflection points. Lateral sinus only moderately deep, apex above whorl middle. Spiral and basal sinuses rel- atively shallow. Measurements. —See Table 58. Stratigraphic and Geographic Distribution. — AL: Nanafalia Formation, Tuscahoma Formation (Bells Landing Member); LA: Pendleton Formation; MX: undifferentiated Wilcox Group; lower— middle Sabi- nian Stage, Upper Paleocene. Table 58. — Measurements of “Turritella” multilira Whitfield. Ab- breviations as in Table 10. MD MH WN FMNH-UC 24521 syntype 11.0 43.5 8.0 USNM 498012 133 60.0 11.0 86 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Type Locality. — Nanafalia Bluff, Marengo County, Alabama (fide Bowles, 1939) (AL-MA-1). Other Localities. —AL-DA-1 (T), AL-PI-6 (Т), AL- WI-55 (PB), LA-NA-16 (L), MX-NL-4 (B). Type Material. — 2 syntypes of T. multilira Whitfield FMNH-UC 24521. Remarks/Discussion. — Although this species is re- ported by Bowles (1939) to be restricted to the Nan- afalia Formation, I have collected a small number of specimens assignable to it from the Bells Landing Member of the overlying Tuscahoma Formation at Bells Landing. As Bowles noted, some specimens of multilira on which the primary spiral ribs are relatively well developed and the adapical carina is not may resemble individuals of “T.” eurynome Whitfield on which secondary spirals are relatively reduced and the adapical carina relatively pronounced, but such inter- mediates do not appear to be sufficiently common to suggest that only a single species is present. “Turritella” praecincta praecincta Conrad, 1864 Plate 14, Figures 1-4, 6 Turritella praecincta Conrad, 1864, p.211; 1865a, p.32; 1866, p.11; Harris, 1897b, p.24, 29,31; 1899a, p.76 in part, pl.10, f.8; 1899b, p.308, pl.55, f.6; Guillaume, 1924, p.290; 1926, p.425; Cooke, 1926, p.94, f.3; Semmes, 1929, f.59-3; Bowles, 1939, p.318, pl.33, f.11; Stenzel and Turner, 1942, card 97; LeBlanc in Barry and LeBlanc, 1942, p.103, pl.13, Ғ10; Wassem and Wilbert, 1943, p.188, 189, 193, p.31, #7; Brann and Kent, 1960, p.933; Palmer and Brann, 1966, p.1002; Toulmin, 1969, pl. 1, f.18; 1977, p.233, рі.29, £. 10,11. Description. —Shell medium sized to large, of 15 to 25 whorls. Maximum observed whorl diameter 31.1 mm. Apical angle 19°; pleural angle 15°; pleural angle not as low on younger specimens. Sutures vary from inconspicuous to deeply incised. Protoconch and ear- liest teleoconch whorls unknown. Earliest whorls ob- served (dia. = 0.36 mm) bearing two moderately prom- inent spirals of approximately equal strength on lower % of whorl, a third, weaker spiral appearing within 1.0 whorl midway between this pair and upper suture and rapidly becoming as strong. At whorl diameter of 1.0— 1.5 mm, a fourth spiral appearing between uppermost of original three and upper suture. Within seven to eight whorls this uppermost spiral becoming most prominent, forming a sharp adapical carina projecting laterally as far as the lowest spiral and producing straight-sided whorl profile. Upper surface of carina on earliest carinate whorls bearing numerous fine spi- rals and slopes up roof-like to suture. Underside of carina immediately concave. Faint intercalaries pres- ent on rest of whorl; original spirals remaining strong but not projecting. Lower margin of whorl slightly to Table 59.—Measurements of “Turritella” praecincta praecincta. Abbreviations as in Table 10. MD MH WN USNM 497991 hypotype 28.0 92.3 12.5 MCZIP 29280 311 98.4 11.5 MCZIP 29281 28.8 78.0 8.0 MCZIP 29282 28.4 82.8 9.5 markedly concave, flaring out to meet suture. Carina quickly increasing in width on later whorls, attenuating outward and often becoming undulatory and/or flared adapically, sometimes leaving suture in a deep groove. Base of whorl bears numerous prominent spirals. Lat- eral aspect of growth line trace prosocline, with well developed upper and lower inflection points. Lateral sinus moderately deep; spiral and basal sinuses rela- tively shallow. Measurements. —See Table 59. Stratigraphic and Geographic Distribution. — AL: Nanafalia Formation, Tuscahoma Formation (Bells Landing and Greggs Landing Members); LA: Pendle- ton Formation; Sabinian Stage, Upper Paleocene. Type Locality. — unknown; given by Conrad as “Dal- las Co., Alabama." Other Localities. — AL-DA-1 (T), AL-PI-2 (Т), AL- MO-3 (T), AL-MO-5 (T), AL-WI-1 (T), AL-WI-18 (B), AL-CH-6 (B), AL-CH-12 (T), TX-SA-4 (WW), LA- SA-21 (WW), GA-CL-4 (B). Type Material. —holotype of 7T. praecincta Conrad unknown (fide Moore, 1962, p.89), hypotype (Bowles, 1939) USNM 49799]. Remarks/Discussion. — Prior to the work of Andrews (1971), the only published mention of “T.” praecincta outside the Gulf coast was a passing reference (“Уа.: Aquia creek.””) by Harris (1899a, p.76). Yet the sharply adapically carinate specimens from the middle of the Paspotansa Member of the Aquia Formation (“Bed E" in the scheme of Beauchamp [1984]) (Plate 14, Figure 4) of Maryland and Virginia are definitely not assign- able to “Т.” humerosa Conrad, and fall reasonably within the range of variation of “T.” praecincta. The Atlantic coast praecincta are smaller than their coun- terparts in Alabama, have smaller apical angles (around 15°), and show more pronounced spiral sculpture on the whorl below the carina, but they show the sharp, ledge-like, sometimes undulating and posteriorly flar- ing adapical carina distinctive of praecincta on the Gulf coast. The carina is never as extreme in the Aquia specimens as in praecincta from Alabama, but it is much more pronounced than in any other adapically carinate form, such as “Т.” multilira. These qualitative assessments are supported by the | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 87 1.5 | О B4 О ) E Do G O ri 0.5 + = О a a Be | | | | | O | | S Бе | | i | | 9 mp | 5 -3 -2.5 -2 1.5 1 -0.5 ) 0.5 D 3 B 05 x -1 T x ж | x x wipe | x ux | | Е2 x Virginia Factor 1 | | С] Alabama | gu T О B 2.56 + a D PS x 1.5 + 1 albo ОБ Г] | со a LE ы > О | 5 — XX L— | een п-9 B | - | o x Deep -1.5 Е] -0.5 0 „05 15 1.5 x TE => a X 2%” 55 вн B | up dE Factor 2 Text-figure 22. — Results of factor analysis of specimens of “Turritella” praecincta Conrad. A. first and second factor axes. B. Second and third factor axes. x represents specimens from Virginia; [0 represents specimens from Alabama (the type area). See Appendix 1 for specimens used and details of analysis. results of the multivariate morphometric analyses factors (Table 61) indicate that variables on the second (Text-figure 22). A total of 118 variables was measured and ninth whorls load most heavily on the first factor, on 28 specimens from Alabama and Virginia. The first whereas variables from the sixth and seventh whorls three factor axes explain 76.896 of the variation in the load most heavily on the second factor and variables total data set (Table 60). Loadings on the first three on twelfth and thirteenth whorls load most heavily on 88 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 60.—Results of factor analysis of specimens of “Turritella” praecincta (Conrad). Variance explained by first five factor axes. cumulative proportion Factor of variance 1 0.4018 2 0.5645 3 0.6396 4 0.7077 5 0.7353 the third factor. Virginina and Alabama specimens fall into two more or less distinct clusters, separated largely along the second factor axis, in a pattern similar to that showed by Palmerella mortoni s.s. and P. m. post- mortoni (Text-figure 18, p.59). Given the geographic separation, and the observation that Virginia speci- mens are most similar to praecincta from Alabama than to any other species, it seems advisable to rec- ognize the Virginia forms as a subspecies of praecincta (see below) rather than a separate species. Although Bowles (1939) states that praecincta in Al- abama is restricted to the Tuscahoma Formation, Toulmin (1977) reports at least two occurrences (lo- calities AL-DA-1, AL-PI-1) in the underlying Nana- falia Formation, probably from the Grampian Hills Member. The specimens I have examined from these localities are markedly smaller than the average for later Bells and Greggs Landing samples (MD < 17.0 mm). “Turritella” praecincta virginiensis, new subspecies Plate 14, Figure 4 Diagnosis. —Similar to but smaller than“ Turritella” praecincta from Alabama, with smaller apical angle and more pronounced spiral sculpture below a less pronounced adapical carina. Description. — Shell medium sized, of perhaps 10- 25 whorls? Maximum observed diameter 16.9 mm. Apical angle approximately 15°, pleural angle approx- imately 12°, giving shell a somewhat domed outline. Sutures usually inconspicuous. Protoconch and early teleoconch whorls unknown. Earliest whorls observed (dia. = 9.8 mm) bearing faint adapical carina composed of several lesser spiral cords. Carina increasing in strength but remaining rounded to subquadrate in pro- file, and not expanding abapically down whorl. Whorls straight-sided below carina, surfaces covered with sub- equal spiral lines of fine to moderate strength. Base of whorl bears numerous prominent spirals. Lateral as- pect of growth line trace prosocline, with well devel- oped upper and lower inflection points. Lateral sinus moderately deep; spiral and basal sinuses relatively shallow. Measurements. — бее Table 62. Stratigraphic and Geographic Distribution. — V A: Aquia Formation; SC: Black Mingo Group (Williams- burg Fm.) (?). Type Locality. —Belvedere Beach, 1 km south of mouth of Potomac Creek, King George County, Vir- ginia (VA-KG-5). Other Localities. — V A-ST-4 (A), VA-ST-6 (A), VA- KG-5 (A), SC-BE-3 (MCZ). Type Material. —holotype of “T.” praecincta virgi- niensis n. ssp. MCZIP 29093, paratype PRI 33199. Remarks/Discussion. —See above under “T.” prae- cincta praecincta. “Turritella” sp. Plate 13, Figure 5 Torquesia prehumerosa Govoni, 1983, p.114-118, pl.5, £.11-15. Description. — Govoni (1983) provides the following description: *Shell medium-size, long, slender, gently tapering turriculate. Suture narrowly but distinctly im- pressed in earlier whorls, becoming indistinct ad- pressed to nearly flush in mature whorls. Protoconch erect, homeostrophic, turbinate, of about 2 smooth, rounded, rapidly expanding volutions; first volution minute, initially depressed; protoconch merges into teleoconch without perceptible break. Whorls of teleo- conch wider than high; total number of whorls un- known. Primary spirals appear on juvenile whorls in order a4 B, C, d; spirals B and C quickly develop into strong, sharp-edged cords (C remaining slightly stron- ger than B) which dominate sculpture of juvenile and earliest adolescent whorls; spiral a initially forms low but distinct, rounded thread midway between B and posterior suture, but increases rapidly in strength rel- ative to concurrently weakening cords B and C until, by about tenth to twelth whorl, it forms a strong thread equal to b and c; d appears very early (probably prior to B) and forms a distinct though increasingly obscured spiral equal in strength to a. Over same interval, whorl sides become very slightly convex above gentle an- gulation formed by primary spiral c; by about eighth or ninth teleoconch whorl, fine secondary threads de- velop, first between spiral a and posterior suture and then in other interspaces below. With continuing ad- dition and strengthening of threads in adolescent whorls, sides become more or less flattened except for very slight inflection between anterior suture and rem- nant of anterior angulation, and a posterior swelling which, in all but most mature whorls, forms a broad, low, rounded, though usually readily apparent subsu- | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 89 Table 61.— Results of factor analysis of specimens of “Turritella” praecincta (Conrad). Rotated factor loadings of each variable on each of | the first three factor axes. Variables as described in Text-figure 3. | Variable factor 1 factor 2 factor 3 Variable factor 1 factor 2 factor 3 | УН! — 0.789 0.144 0.046 W3-7 0.044 0.921 —0.007 SWI —0.789 0.145 0.047 CR7 —0.004 0.915 0.028 cwl —0.789 0.146 0.047 CD? —0.256 0.834 —0.058 | W1-1 —0.789 0.144 0.046 WW? 0.119 0.904 0.044 | W2-1 —0.788 0.144 0.146 WHS8 0.700 0.538 —0.150 | W3-1 —0.787 0.143 0.045 SW8 0.712 0.575 —0.099 | CRI —0.788 0.146 0.047 CWS8 0.701 0.603 —0.089 | CDI —0.697 0.138 0.046 W1-8 0.708 0.568 —0.116 WWI —0.789 0.144 0.046 W2-8 0.710 0.565 0.27 WH2 —0.908 0.158 — 01097 W3-8 0.701 0.561 —0.129 Sw2 0.906 0.158 01029 CR8 0.701 0.555 0:075 CW2 —0.908 0.159 =0.027 CD8 0.596 0.614 —0.002 W1-2 —0.908 0.159 0.097 WWS 0.710 0.574 —0.117 W2-2 “(ОУ 0.160 —0;021 WH9 0.931 —0.028 0.001 W3-2 —0.916 0.160 —0.020 SW9 0.940 0.069 —0.082 CR2 :20:917 0.161 —0.020 CW9 0.921 0.233 —0.051 (3192 —0.920 0.162 —0.005 W1-9 0.946 0.098 0.091 | WW2 0.903 0.158 —0.030 W2-9 0.947 0.073 —0.085 | WH3 0/52) 0.337 —0.309 W3-9 0.941 0.058 —0.095 SW3 2207/20 0.344 —0.330 CR9 0.924 0.011 0.067 CW3 —0.734 0.343 — 0825 CD9 0.733 0.486 —0.013 | W1-3 08129! 0.344 0829 WWO9 0.940 0.072 —0.068 | W2-3 SOs), 0.344 2204825 WH10 0.819 —0.310 0.248 | W3-3 —0.749 0.341 —0.318 SW10 0.852 —0.134 0.251 | CR3 —0.768 0.337 —0.305 CWI10 0.852 —0.014 0.254 | CD3 —0.541 0.351 0375 W1-10 0.854 —0.143 0.221 WW3 079 0.340 | W2-10 0.851 —0.162 0.229 | WH4 —0.583 0.536 —0.402 W3-10 0.850 —0.167 0.228 | Sw4 E0505 0.550 —0.416 CR10 0.824 —0.179 0.349 CWA —0.514 0.547 —0.412 CD10 0.745 0.188 0.280 W1-4 = 0.585 0.545 —0.411 WW10 0.856 0.15 0.226 | W2-4 SD 0.544 —0.409 УН! 0.805 – 0.367 0.234 | W3-4 —0.547 0.545 —0.411 SWII 0.849 —0.150 0.225 CR4 —0.546 0.543 —0.409 Cwll 0.850 —0.061 0.221 | CD4 —0.481 0.521 —0.385 WI-11 0.850 —0.164 0.219 | WWA 0/518 0.549 —0.415 W2-11 0.849 —0.169 0.211 | WHS —0.285 0.826 0.047 W3-11 0.846 0.191 0.223 SWs —0.234 0.837 0.064 CR11 0.823 0.2 0.316 CWS USO 0.837 0.067 CD11 0:753 0.101 0.185 WI-5 —0.254 0.835 0.071 wwii 0.852 S04 37, 0.228 W2-5 —0.249 0.835 0.071 WH12 0.494 —0.070 0.576 W3-5 —0.264 0.834 0.077 Sw12 0:535 0.038 0.546 CRS ISS 0.832 0.062 Cw12 0.522 ӨЛҮ? 0.550 | CD5 —0.292 0.796 0.040 WI1-12 0.540 0.051 0.544 | WWS 0.219 0.833 0.071 W2-12 0.538 0.044 0.548 | WH6 ОС 0.924 0.005 W3-12 0.536 0.040 0.546 | SW6 —0.045 0.927 0.056 CR12 0.514 0.021 0.607 | CW6 —0.062 0.930 0.039 CD12 0.547 0.227 0.528 | W1-6 —0.053 0.926 0.061 WWI2 0.543 0.066 0.548 | W2-6 —0.043 0.923 0.051 WH13 —0.047 0.003 0.817 | W3-6 —0.044 ШӘП? 0.059 SWI13 0082 0.055 0.832 | CR6 —0.084 0.921 0.017 CW13 —0.026 0.076 0.833 | CD6 —0.189 0.849 —0.108 W1-13 = 0.035 0.054 0.833 | Wwe —0.026 0.920 0.077 W2-13 — 0:032 0.054 0.833 WH7 —0.019 0.923 2010 DT W3-13 01035 0.047 0.832 | SW7 0.069 0.914 0.061 CR13 0.037 0.039 0.831 | CW7 0.026 0.916 0.019 CD13 —0.016 0.107 0.807 | УУ1-7 0.040 0.914 0.021 WWI13 20:029 0.063 0.830 | W2-7 0.042 0.916 0.029 90 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 62.—Measurements of “Turritella” praecincta virginiensis. Abbreviations as in Table 10. MD MH WN MCZIP 29093 holotype 16.9 >03 9.0 MCZIP 29094 15:9 58.1 9.0 tural collar. The numerous fine, subequal spiral threads, numbering about 40 in largest preserved whorls, cover entire whorl surface including subsutural collar. Last whorl lacks obvious subsutural collar and exhibits a moderately strong, spiral thread-covered carina at the basal whorl angulation that apparently extends back from aperture onto penultimate whorl just above an- terior suture; whorl base covered by numerous spiral threads. Growth lines indistinct, sinuous; acutely pro- socline in upper third of whorl and steeply opisthocline below, forming a moderately deep spiral lateral sinus with vertex above midwhorl; slope increases in ante- rior third of whorl until, in lower fifth, slope increases very rapidly so that lines intersect suture at nearly right angle; recurved slightly on base angulation to form very shallow spiral arc on base. Aperture incompletely known; parietal region calloused.” Measurements. —See Table 63. Stratigraphic and Geographic Distribution. — MD: Brightseat Formation; Lower Paleocene. Type Locality. —bluff on Southwest Branch, just W of Capital Beltway, Prince Georges County, Maryland (MD-PG-12). Other Localities. — MD-PG-13 (GO), MD-PG-14 (GO). Type Material. —holotype and 20 paratypes of Tor- quesia “prehumerosa” Govoni USNM. Remarks/Discussion. — Of all the adapically carinate species of Midwayan age, this form most closely re- sembles the Sabinan-age “T.” humerosa Conrad. Both species show a similar apical sculpture formula and form of the adapical carina, i.e., rounded and including more than a single spiral rib. Together with its geo- graphic position, these characters make prehumerosa from the Brightseat a good candidate for a direct an- cestor of humerosa from the Aquia (cf., Govoni, 1983). Table 63.—Measurements of “Turritella” sp. (“prehumerosa” Govoni). Abbreviations as in Table 10. MD MH WN holotype USNM 11.1 43.8 11 paratype USNM 9.1 29.6 >) paratype USNM 5.6 14.3 5 paratype USNM 1.4 4.4 8 “T.” prehumerosa is very similar to biboraensis Gard- ner, their geographic separation being perhaps the chief distinguishing feature. Govoni’s form differs from “T.” toulmini n.sp. in having a relatively straight whorl pro- file below the carina. See Govoni and Hansen (in press) for further discussion. “Turritella” toulmini, new species Plate 12, Figure 6-7 Turritella humerosa Conrad. Toulmin, 1977, pl.8, f.6; not Conrad, 1835b. Diagnosis. — Medium sized turritellid with a broad, rounded adapical carina, moderate, even spiral sculp- ture, and convex, slightly inflated whorl profile, widest around midwhorl, Description. —Shell medium to large; total number of whorls unknown. Maximum observed whorl di- ameter 18.0 mm. Apical angle 105; pleural angle may be slightly less than apical angle. Sutures inconspicu- ous. Protoconch and earliest teleoconch whorls un- known. Earliest teleoconch whorls observed on which sculptural detail is visible (dia. = 3.0 mm) showing at least three spirals, approximately evenly spaced over whorl, one at upper suture already forming an incipient adapical carina, one weaker near whorl middle, one in about middle of lower Y. of whorl. Whorls at this size straight-sided, approximately as high as wide. Spire attenuate, many-whorled. Numerous intercalary spi- rals appearing on succeeding whorls; sculpture of latest ' whorls consisting of approximately evenly spaced, equal strength spirals over upper % of whorl, intercalated with one or more finer lines, and two to three stronger spirals on lower У. Whorl profile of late whorls convex over lower % of whorl, widest point at about whorl middle, with a moderate constriction just below the pronounced, rounded, broad adapical carina. Lateral aspect of growth line trace orthocline, with well de- veloped upper and lower inflection points and mod- erately deep sinus with apex just above whorl middle. Basal sinus unknown. Measurements. —See Table 64. Stratigraphic and Geographic Distribution. — AL: Clayton Formation; lower Midwayan Stage, Lower Pa- leocene. Type Locality. —type section of Clayton Formation, Table 64.— Measurements of “Turritella” toulmini n. sp. Abbre- viations as in Table 10. MD MH WN MCZIP 29274 holotype 14.6 55.0 8 MCZIP 29275 paratype TAS 62.0 6 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 91 RR cut E of Clayton, Barbour County, Alabama (AL- BA-1). Other Localities. —GA-CL-6 (T). Type Material.—holotype of T. toulmini n. sp. MCZIP 29274, paratypes MCZIP 29275, 29276, PRI 33197. Etymology. — Named for the late Lyman D. Toul- min, pioneer in Lower Tertiary stratigraphy and pa- leontology in the eastern Gulf Coastal Plain. 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Digitized points and calculated measurements for these five species are shown in Text-figure 3. These Measurements were divided into separate computer files, corresponding to separate samples (Table 65), and analyses began with these files. For each species, sub- sequent analysis followed these steps: 1) Each file was run separately through the computer program HISPIRE3 (Morris and Allmon, 1994), which aligns whorls of multiple specimens by comparing whorl heights. HISPIRE3 aligns incomplete shells in three steps. In the first step, the program creates an array of whorl heights, in which rows are specimens and columns are whorls. It then sorts this array in order of increasing size of first (smallest) whorl. In the second step, the program steps through the rows (specimens) of the ar- ray and compares the heights of the first whorl of each pair of specimens. If a specimen has a first whorl of more than 0.8 of a whorl larger than the first whorl of the previous specimen, then blank whorls are added Resources and Development Commission, Division of Geology Bulletin 19, 125 pp., 2 pl. Wiley, E.O. 1981. Phylogenetics. John Wiley and Sons, New York, 439 pp. Wilmarth, M.G., compiler. 1930. Tentative correlation ofthe named geologic units of Texas. U.S. Geological Survey Chart, 1 sheet. Wilson, B. 1993. Australian marine shells. Prosobranch gastropods. Part one. Odyssey Publishing, Kallaroo, Western Australia, 408 pp. Woodring, W.P. 1928. Miocene mollusks from Bowden, Jamaica; Part II, Gas- tropods and discussion of results. Carnegie Institution of Washington Publication 385, 564 pp. 1957. Geology and paleontology of Canal Zone and adjoining parts of Panama. Geology and descriptions of Tertiary mollusks (Gastropods: Trochidae to Turritellidae). U. S. Geological Survey Professional Paper 306-A, 145 pp. 1972. Zoogeographic affinities of the Tertiary marine molluscan faunas of Northeastern Brazil. Anais Academia Brasileira Ciencias 43 (supplement), pp. 119-124. Woodward, S.P. 1851. A manual of the Mollusca part 1. viii + 158 pp, pl. 1-12. Wu, R.S.S., and Richards, J. 1981. Variations in benthic community structure in a sub-trop- ical estuary. Marine Biology, vol. 64, pp. 191-198. to its small end, and it is moved to the right in the array until it comes close to the size of a corresponding whorl in the previous specimen. This creates an array in which specimens are sorted in order of increasing size and are aligned by the sizes of their first whorls. In the third step, HISPIRE3 iteratively a) finds the average height of each whorl in a composite average specimen in the array, b) applies up to three methods to move specimens to reduce the total error in the data set, and c) calculates the change in total error. The three methods are as follows. The specimen with the largest single error from the composite average is moved by one whorl to reduce its error (Method 1), then the specimen with the second largest errot is moved (Meth- od 2), then the program steps down throught the spec- imens (from largest to smallest), and moves each spec- imen that is more than one whorl in error (Method 3). These iterations are repeated until the corrections pro- duce an increase in the total error. This process is repeated until the number of passes is equal to 25 the number of specimens (a proportion arrived at empir- ically by experimenting with model data sets). One round of such iterations is called a “pass.” The three methods are used differently in different passes. Method 1 is used in each iteration of all passes. Method 2 is used in all but the last two passes. Method 102 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Table 65.—Summary of samples used for morphometric analysis. no. of по no. of no. speci- of speci- of Species/sample locality horizon mens whorls CV Species/sample locality horizon mens whorls CV Palmerella mortoni CARINA6 AL-CL-4 GS 9 6 4.57 MORTON2A AL-MO-3 TSBL 4 8 7.43 CARINAS8 AL-MO-1 GS 4 8 4.78 MORTON2B = AL-MO-3 TSBL 8 8 6.23 CARINA9 AL-MO-11 GS 7 8 B MORTON2C AL-MO-3 TSBL 11 9 7.63 CARINA10 AL-CL-4 GS 7 6 Su» MORTON2D AL-MO-3 TSBL 6 9 6.08 CARINA11 AL-CL-15 GS 17 9 5.73 MORTON2E AL-MO-3 TSBL 9 10 6.23 CARINA13 AL-CL-4 GS 11 9 6.94 MORTON2F AL-MO-3 TSBL 32 9 7:82 CARINA14 AL-CL-4 GS 11 7 6.71 MORTON2G AL-MO-3 TSBL 17 9 6.87 CARINAI5 AL-WA-6 GS 8 7 6.83 MORTON2H AL-MO-3 TSBL 15 8 8.71 CARINA17 AL-CL-14 GS? 10 8 4.99 MORTON3A AL-MO-3 TSBL 23 9 7.61 CARINA18 AL-CL-14 GS? 12 9 6.49 MORTON3B AL-MO-3 TSBL 17 11 9.74 CARINA19 AL-CL-14 GS? 8 8 4.27 MORTON3C = AL-MO-3 TSBL 12 10 7.26 CARINA20 AL-CL-15 GS Y 7 558 MORTON4B AL-MO-3 TSBL 18 9 9.32 Haustator perdita MORTON4C AL-MO-3 TSBL 23 10 7.96 PERDITI MS-HN-6 MB 15 9 8.10 MORTON6B AL-MO-3 TSBL 3 8 7.54 PERDIT2 MS-CL-3 MB 8 9 5.24 MORTON7 AL-MA-1 NF 4 10 5.14 PERDIT4 MS-HN-6 MB 16 7 8.17 MORTONS AL-MA-1 NF 9 8 6.40 PERDIT5 MS-HN-6 MB 4 6 4.94 MORTON9 AL-MO-5 TSGL 10 9 8.46 PERDIT6 MS-CL-15 MB 8 7 5.73 MORTONI11 AL-WI-1 TSBL 23 10 7216 PERDIT7 MS-CL-3 MB 8 8 5.74 МОЕТОМ 12 АТ-СН-7 МЕ 5 10 5.59 PERDIT8A MS-CL-15 MB 15 9 ЕЈ МОЕТОМ 13 AL-DA-1 NF 8 11 4.52 PERDIT8B MS-CL-15 MB 14 10 6.77 MORTONI15 AL-MO-3 TSBL 6 q 6.92 “Turritella” praecincta МОЕТОМ 17 VA-KG-5 AQ 4 10 5.14 PRAECI AL-MO-5 TSGL 9 9 7,29 МОЕТОМ18 VA-KG-5 AQ 17 10 6.94 PRAEC2 AL-MO-3 TSBL 11 12 5.92 MORTON2O VA-KG-6 AQ 18 10 6.37 PRAEC4 VA-ST-2 AQ 9 Т) 4.74 MORTON21 VA-KG-6 AQ 14 10 7.32 Turritella terebra MORTON22 MD-PG-6 AQ 8 10 5.84 TEREBR І R-1 У 9 6.05 MORTON?23 MD-PG-6 AQ 15 8 8.83 TEREBR2 R-2 3 [2 3.10 MORTON24 MD-PG-6 AQ 7 9 4.83 TEREBR4 R-4 10 14 5.93 MORTON26 VA-KG-6 AQ 10 9 5.93 TEREBR9 R-5 9 13 Sev y Haustator carinata TEREBR10 R-6 8 13 6.43 CARINA2 AL-CL-4 GS D 12 7.94 TEREBR18 R-2 8 14 7223 CARINA4 AL-CL-15 GS 12 8 4.15 ТЕКЕВК19 R-8 34 13 DAI 3 is used in passes 2 through n where n= the number of specimens in the sample. Variations of this default alignment method work better in some data sets. The user can change the num- ber of passes and thus the methods (1-3) used. If, how- ever, the number of passes has been forced (by the user) to less than / the number of specimens, Method 3 is not used; if the number of passes has been forced (by the user) to some greater number of specimens, then Method 3 is used only in the second iteration. These iterations produce a matrix of aligned speci- mens. In the final step this matrix is used to create the final data file from the original input data file, restoring specimens to their original order (but numbering them consecutively from 1; original numbering is lost unless specimen number is included as an additional vari- able). Missing whorl measurements are filled in this matrix with zeroes. HISPIRE3 assays the accuracy of whorl alignments by checking the variation of heights for each whorl in the sample, and warns the user if this variation suggests poor alignments. The coefficient of variation (CV) is calculated for each aligned whorl and these values are then averaged across all whorls and again across all but the first and last whorls. Experience with the pro- gram suggests that if these two CV’s are greater than 7-8%, the alignments of one or more specimens are likely to be incorrect by more than a full whorl. In the initial runs, the default parameters of the pro- gram were used and CV’s recorded. 2) For those files whose CV’s exceeded 7%, a se- quence of new analyses was performed. First, the num- ber of passes was forced to Y the number of specimens; then the number of passes was forced to less than Уз the number of specimens; then the number of passes was forced to twice the number of specimens. 3) For files whose CV’s still exceeded 7%, graphs were made of aligned whorl number against whorl height. Specimens that still clearly lay outside the range of variation of the others were noted and these spec- PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 103 imens were manually adjusted by one whorl in either direction in the data file. 4) For one file, the range of variation in the whorl number/whorl height graph suggested that the speci- mens were all poorly aligned. This file (PERDIT1) was run through the program HISPIRE2, which aligns whorls by whorl expansion rate (W of Raup, 1966). See Morris and Allmon (1994) for further discussion. 5) Each data file had been run separately up to this point, yet single species were represented by many data files, and all of these now had to be aligned to each other. To do this, each separately aligned file was run through BMDP program 2D to calculate average val- ues of whorl height for each whorl. The resulting values were assembled to form a composite average specimen representative of each file. 6) All composite average specimens for each species were grouped together and were now themselves run through HISPIRE3 in order to obtain alignment among the separate data files (i.e., samples) within each spe- cies. The results of this HISPIRE run were examined and the number of “empty” whorls (i.e., zeroes) added to the beginnings and/or ends of each composite was noted. 7) All files for each species were strung together in a new file, and the appropriate number number of empty whorls added to each file. The result was a single large data file for each species, in which specimens within samples and samples with each other had both been aligned by HISPIRE. APPENDIX 2 Locality Register Listed here are all localities given as occurrences for the turritellid species described in this paper. Some localities are redundant because of imprecise descrip- tions or changes in place names. One or more refer- ences are given when these have been available. In- formation on lithostratigraphic units from which fos- sils were collected is also included if known. When full descriptions of localities can be found in readily avail- able sources such as Barry and LeBlanc (1942) and Toulmin (1977), I have only included their locality designation. References to collections following each locality are as follows: USGS—U.S.Geological Survey localities, listed in registry books deposited in U.S.National Museum; PRI—Paleontological Re- search Institution, Ithaca, NY; MGS—Mississippi Geological Survey; WA —locality from which I have collected personally. Page numbers on which each lo- cality is cited are listed after the locality description. ALABAMA Barbour County AL-BA-1 type locality of Clayton Fm., RR cut approx. 300 m north of Co. Hwy. 28, approx. 0.3 miles east of Clayton (86WA23; USGS 12164). pp. 83, 91 Butler County AL-BU-1 Toulmin (1977) loc. Abu 1. pp. 68,81 AL-BU-2 Toulmin (1977) loc. Abu 6. pp. 68,81 AL-BU-3 Toulmin (1977) loc. Abu 11. pp. 68, 81 AL-BU-4 Toulmin (1977) loc. Abu 3. pp. 73 Choctaw County AL-CH-6 Tuscahoma Landing, Tombigbee River; Tuscahoma Fm. incl. Bells Landing Marl (Toulmin, 1977, loc. ACh-2; USGS 5472). pp. 63, 83, 86 AL-CH-9 roadcut on Co. Hwy 21, approx. 1.3 miles south of Bar- rytown, 2.9 miles north of jct. with Hwy. 84, approx. 0.1 miles south of bridge over Souwilpa Creek, Coffeeville Lock and Dam 7.5 min. quad., T.10 N, R.3 W, sec 24; Upper Lisbon Fm. and Gosport Sand (?) (Toulmin, 1977, Loc. ACh-8; 85WA115). p. 69 AL-CH-11 road cut on Ala. Hwy 17, 6.7 mi N of Butler courthouse, sec 22 T 14 N, R 2 W; Bells Landing Mbr, Tuscahoma Fm. (Toulmin, 1977, loc. Ach12). p. 63 AL-CH-12 Toulmin (1977) loc. Ach20. pp. 63, 86 Clarke County AL-CL-1 Choctaw Corner. p 73 1b 1% mi WSW of Choctaw Corner. p. 73 1с headwaters of Beaver Creek, approx. % mi E of Choctaw Cor- ners (USGS 7153, 7154). p. 73 1d “gully on land of Henry Jackson, near middle of sec 7, T 11 N, R 2 E, about 3 miles west of Choctaw Corners" (Bowles, 1939, p. 303) (USGS 7155). p. 73 AL-CL-2 Woods Bluff, left bank of Tombigbee River, sec 15, T 11 N, R 1 W; Bashi Fm. (Toulmin, 1977, loc. Acl2; USGS 262, 2667, 3099, 3100, 5470, 6205, 6206, 6207, 7482). p. 73 AL-CL-4 Little Stave Creek, approx. 3 mi. north of Jackson, approx. 0.75 mi. west of Hwy. 43, Jackson 7.5 min. quad. T.7 N, R.2 E, sec 20,29; Marianna Ls., Yazoo Fm., Moody’s Branch Fm., Gos- port Sand, Lisbon Fm., Tallahatta Fm. (83WA07,08, 85WA84). pp. 46, 69 AL-CL-7 road cut on east side of Ala. Hwy. 69, south side of Bashi Creek, 2.5 miles south of Marengo Co. line, 1.3 miles north of Camp- bell, T.11 N, ЕЛ E, sec 9 (Toulmin, 1977, Loc. Acl-1). p. 73 AL-CL-13 Coffeeville Landing, left bank Tombigbee River, Coffee- ville 7.5 min. quad., R.1 W, T.9 N, sec 8,17 (Toulmin, 1977, Ас13) (85WA103, 86WAS9; USGS 12181). pp. 77, 79 AL-CL-14 faulted blocks downstream from Coffeeville Landing, left bank Tombigbee River, Coffeeville 7.5 min. quad., section 17; Lisbon Fm. (83WA03,04, 85У/ 108, 86WA60). рр. 51, 54 AL-CL-16 between Alabama and Tombigbee Rivers, 1 mi SW of Rockville, 8.5 mi SE of Jackson (USGS 6157). p. 54 Coffee County AL-CO-1 Elba Dam on Pea River; Bashi Fm. (Toulmin, 1977, ACof- 1) (85М А114, 86WA45; USGS 10013, 10780). pp. 51, 72 AL-CO-2 near mouth of branch flowing into Pea River at Church- wells Branch, 5.5. mi S of Elba (USGS 10012, 10781.) . p. 73 Conecuh County AL-CN-1 E side of Sepulga River on land of J.G.Robinson, sec 13, T4N,R 13 E (USGS 6737). pp. 54, 77 104 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Dale County AL-DA-1 Becks Mill (Munn’s Mill), bed of Pea River, just upstream of Veterans Memorial Bridge (Hwy. 231), sec 7, T 7 N, R 23 E (Toulmin, 1977, Loc. Ada-2; 86WA41; USGS 10769). pp. 63,83, 86, 88 AL-DA-3 “hillside at swimming pool, near Atlantic Coast Line RR Station, Ozark” (Bowles, 1939,. p. 303); Bashi Fm. (USGS 11093). p. 73 AL-DA-4 shell bed in bank of Hurricane Creek, a southward flowing tributary of the Choctawatchee River, beneath bridge on Ala. Hwy 134 to Enterprise, approx. % mi W of jct with old US Hwy. 231, sec 2 T 4 N, R 24 E; Bells Landing Mbr. p 63 Henry County AL-HE-1 Toulmin (1977) loc. Ahe3. p.68 AL-HE-2 Toulmin and LaMoreaux (1963, p.399) bed 8 (Clayton Fm.), 80.7-80.4 mi above confluence of Flint and Chattahoochee Rivers, approx 8 mi above Ft. Gaines. p. 61 AL-HE-3 approx. 7 mi above Ft. Gaines, near Morris Landing (Veatch and Stephenson, 1911, p.220); Clayton Fm. p.61 AL-HE-4 Toulmin (1977) loc. Ahel. p. 63 Marengo County AL-MA-1 Nanafalia Landing, Tombigbee River, Pennington 7.5 min. quad., T.14 N, R.1 E, section 31 (USGS 271, 5641) AL-MA-2 Toulmin (1977) loc. Ama-1; Porters Creek Fm. Monroe County AL-MO-1 Claiborne Landing and Bluff, left bank Alabama River, downstream from Hwy. 84 bridge at Claiborne, outcrops at water level and up steep bluff mostly overgrown except for occasional slumps, Claiborne 7.5 min. quad. T.7 N, R.5 E, section 25 (85WA95,96,97, 86УУА57) la = Gosport Sand in bluff (USGS 263, 2391, 2867). pp. 46, 51, 68 1b = Upper Lisbon Fm. exposed at base of bluff on river bank (USGS 2395, 2396, 12171). pp. 54, 77, 79 AL-MO-3 Bell’s Landing, left bank of Alabama River, Hybart 7.5 min. quad. T.10 N, R.6 E, sec 36; Bells Landing Marl Mbr., Tuscahoma Fm. (85WA85, 86WA55; USGS 260, 2669, 3098, 5593, 5594, 5595). pp. 63,83, 86 AL-MO-4 Lisbon Landing/Lisbon Bluff, right bank Alabama River, approx. 1 mi S of Claiborne Lock and Dam, Claiborne 7.5 min. quad., T.7 N, R.5 E, sec 3; Lower Lisbon Fm. (fide Toulmin, 1977) (85WA94; USGS 3105, 5511, 6086). pp. 68, 77 4b Lisbon Bluff, Bed 7 of Cooke section (USGS 13431, 14215). p. 68 4c Lisbon Bluff, Bed 4 of Cooke section (USGS 13433). p. 68 4d 1 mi S of Lisbon Landing. pp. 54, 79 AL-MO-5 Gregg's Landing, right bank Alabama River just downstream of island, Hybart 7.5 min. quad., T.10 N, R.6 E, sections 20, 29; Greggs Landing Marl Mbr., Tuscahoma Fm. (85WA86, 86WAS6, USGS 268, 2670, 3117, 3118, 3604, 5642). pp. 63, 83, 86 AL-MO-11 Rattlesnake Bluff, left bank Alabama River, Claiborne 7.5 min. quad., T.7 N, R.5 E, section 31; Gosport Sand (83WA09, 85WA98). p. 68 Pike County AL-PI-1 Three Notch Road., middle of sect. 17, T 8 N, R 20 E, 1⁄4 mi NE of Henderson; Nanafalia Fm. (USGS 10765). pp. 63, 88 AL-PI-2 4.6-4.8 mi. north of Henderson on Co. Hwy. 21 (not marked — first right on entering Henderson from east on Hwy. 6) (Toulmin, 1977, Loc. Ар1-27) (86WA42). p. 86 AL-PI-6 County hwy 3 in abandoned road bed on N side of paved road 1.0 mi NW of Henderson sec 7 T 8 N, R 20 E; Nanafalia Fm. (Toulmin (1977) loc. Apil). pp. 63, 86 Washington County AL-WA-2 Hatchetigbee Bluff, right bank Tombigbee River at sharp bend, Tattlersville 7.5 min. quad., T.8 N, R.1 W, sections 17,38,20,21; type section, Hatchetigbee Fm. (Toulmin, 1977, Awa- 1; 83WA05,06, 85WA100, 86WA61). р. 73 AL-WA-6 outcrop on Tombigbee River near Lone Star Cement Co. quarry, Tattlersville 7.5 min. quad., T.8 N, R.1 W, section 32; Gosport Sand (85WA101). p. 69 Wilcox County AL-WI-1 Yellow Bluff, right bank Alabama River at sharp bend, Pine Hill 7.5 min. quad., T.11 N, R.6 E, sec 18,19 (Toulmin, 1977, Awil9; 85WA87; USGS 3121, 7480, 10782). pp. 63,86 AL-WI-2 1 mile west of Oak Hill (USGS 3107). p. 81 AL-WI-3 Matthew’s Landing, right bank of Alabama River at bend, Lee Long Bridge 7.5 min. quad., T.12 N, R.6 E, section 2 (85WA91, 86WA53; USGS 3116, 2671, 5596). pp. 68, 81 AL-WI-9 Prairie Creek; Clayton Fm. (USGS 264). pp. 53,61, 68, 81, 83 AL-WI-12 Dales Branch, near Oak Hill. p. 53 AL-WI-13 2 mi W of Oak Hill. p. 53 AL-WI-16 “Black Bluff’ beds, surface of prairies at Graveyard Hill, 3 mi W of Oak Hill Post Office; Clayton Fm. (USGS 3102 ?). p. 68 AL-WI-18 Lower Peachtree Ferry, Alabama River. pp. 83, 86 18b Lower Peachtree Ferry, “2nd fossil horizon, 60 ft. from top, 80 ft. from bottom of bluff’ (Bowles, 1939, p. 299) (USGS 5602). p. 63 18c descent to Lower Peachtree Ferry; Tuscahoma Fm. (Cooke 1926, p.264) (USGS 10779) (= AL-WI-34). p. 63 AL-WI-21 Black Creek branch of Prairie Creek (USGS 283). pp. 68, 81 AL-WI-24 downgrade of Hwy. 21 toward creek, just north of jct. with Hwy. 28; type locality of Pine Barren Mbr.,MacBryde Mbr., Clayton Fm. (Toulmin, 1977, Loc. Awi-3; 85WA78) р. 68 AL-WI-25 outcrop on east side of road, approx. 2.0 miles north of jct of Hwy 41 and 10 at Camden; Lower Mbr., Porters Creek Fm. (Toulmin, 1977, Loc. Awi-27; 85WA79). p. 68 AL-WI-36 Clayton Fm. (USGS 281). pp. 68, 81 AL-WI-37 Clayton Fm. (USGS 284). pp. 68, 81 AL-WI-38 top of hill 0.8 mi W of J Lee Long Bridge in ditch and field near jct Ala Hwy 10 and 28 sec 7 T 13 N, R 7 E; Clayton Fm. (LaMoreaux and Toulmin, 1959, p.54) (Toulmin (1977) loc. 5). p. 68 AL-WI-39 Clayton Fm. (Toulmin (1977) loc. 31). p. 68 AL-WI-40 Clayton Fm. (Toulmin (1977) loc. 34). p. 68 AL-WI-41 Clayton Fm. (Toulmin (1977) loc. 37). p. 68 AL-WI-42 Porters Creek Fm. (Lower Member) (Toulmin (1977) loc. 8). p. 68, 81 AL-WI-43 Porters Creek Fm. (Lower Member) (Toulmin (1977) loc. 10). pp. 68, 81 AL-WI-44 Porters Creek Fm. (Lower Member) (Toulmin (1977) loc. 26). pp. 68, 81 AL-WI-45 Porters Creek Fm. (Matthews Landing Mbr.) (Toulmin (1977) loc. 11). pp. 68, 81 AL-WI-46 Porters Creek Fm. (Matthews Landing Mbr.) (Toulmin (1977) loc. 12). pp. 68, 81 AL-WI-47 Porters Creek Fm. (Matthews Landing Mbr.) (Toulmin (1977) loc. 13). pp. 68, 81 AL-WI-48 Porters Creek Fm. (Matthews Landing Mbr.) (Toulmin (1977) loc. 14). pp. 68, 81 AL-WI-49 Naheola Fm. (Coal Bluff Marl Mbr.) (Toulmin (1977) loc. 16). pp. 68 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 105 AL-WI-50 Clayton Fm. (Toulmin (1977) loc. 3). p. 81 AL-WI-51 Camden—Oak Hill Hwy [Hwy 10] road cut 9.9 mi E of Camden NE % sec 10 T 11 N, R 9 E, at school near Rosebud Porters Creek Fm. (Lower Mbr.) (Toulmin (1977) loc. 9; 85WA80; Siesser, 1983, loc.A15). p. 81 AL-WI-52 Porters Creek Fm. (Lower Mbr.) (Toulmin (1977) loc. 35). p. 81 AL-WI-53 Naheola Fm. (Coal Bluff Marl Mbr.) (Toulmin (1977) loc. 20). pp. 53, 81 AL-WI-54 Naheola Fm. (Coal Bluff Marl Mbr.) (Toulmin (1977) loc. 25). р. 59 AL-WI-55 “new hwy to Camden, 9.7 mi W of Pineapple"; Nanafalia Fm. (USGS 10766). pp. 63, 86 AL-WI-57 Toulmin (1977) loc. 23. p. 63 AL-WI-58 Toulmin (1977) loc. 19. p. 63 AL-WI-59 "state Hwy 96, 10.3 mi NE of Kimbrough" (Stenzel and Turner, 1942, card 46) p. 53 ARKANSAS Bradley County AR-BR-1 well ~ 30 yds NW of Lee Hammaker's house, middle of sec. 8, T 12S, R 9 W, approx. 18 mi w of Rison (USGS 2404). pp. 47, 49 Cleveland County AR-CL-1 Vince Ferry, Saline River (PRI Loc. 897) 1b % mi above Vince Bluff (USGS 2403). pp. 47, 49 AR-CL-4 well at Rison, NE Ya, sec 1, T9 S, R 11 W (USGS 2231). рр. 47, 49 4b well at Rison, SE М, SW М, sec 1, T9S, R 11 W (USGS 2413). p. 49 AR-CL-7 well at the Orton place, Toledo (USGS 2410). p. 49 AR-CL-8% mi N of Crossroads Church, 4.5 mi NW of Kingsland T9S,R 12 W, SW % sec 22 (USGS 2420). p. 47 Drew County AR-DR-1 Wadsworths well, Long Prairie. p.47 Jefferson County AR-JF-1 White Bluff, left bank of Arkansas River — 3.5 mi E of Redfield, W Y», SE М, sec 19, T 2 S, R 10 W and W ‘2, NE М, NW и sec 30, T 3 S, R 10 W [fossil bed covered by backwater from dam— 1985] (USGS 2220). pp. 47, 49 Lonoke County AR-LO-1 near Cabot (Harris, 1894, p.25-6). p. 61 Pulaski County AR-PU-1 near Olsens switch, a few miles SW of Little Rock. p.68 AR-PU-4 Fourche Cr., near mouth of Crooked Creek, 8-9 mi SW of Little Rock (Stephenson and Crider, 1916, p.49). p. 61 St. Francis County AR-SF-1 Hwy 70 over Crow Creek, approx. 3.0 mi. east of Forest City, approx. 0.5 mi. west of Madison, Madison 7.5 min. quad. R.3 E, T.5 М, section 25 (85УУА56; PRI 1046; USGS 41577). pp. 47, 49 AR-SF-2 upstream of RR bridge over Little Crow Creek, southwest of Madison, Madison 7.5 min. quad., T.5 N, R.3 E, section 36 (85WA57; USGS 42667; PRI 1048). pp. 47, 49 AR-SF-4 Crowleys Ridge, Big Crow Creek (USGS 4156; PRI 894). pp. 47, 49 White County AR-WH-1 Bradford (Harris, 1894, p. 25; Stephenson and Crider, 1916, p.51). p. 61 AR-WH-2 cut of St. Louis, Iron Mountain and Southern RR , % mi S of Grandglaise (Stephenson and Crider, 1916, p.52). p. 61 FLORIDA Citrus County FL-CI-1 Large quarry around 5 mi.SE of Crystal River. (MacNeil, 1946.) p. 74 FL-CI-2 Suwannee River at Ellaville. (MacNeil, 1946.) p. 74 Leon County FL-LE-1 “Road metal pit 2.9 mi S of the northern limits of the town of Gulf Hammock, just SW of state road 55." SW % Sec. 34, T. 14 S, R. 16 E. p. 72 FL-LE-2 “About % mile below the Florida Power Corporation Plant at Inglis, Florida, on the right bank of the Withlacoochee River." SE 4 NW 4 Sec 3, T17S, RI6E. p. 72. Marion County FL-MA-1 Martin Station, old railroad station, 10 mi N of Ocala, at junction of Hwy 441 and Martin-Anthony road. p. 74 GEORGIA Bibb County GA-BI-1 Brown's Mountain, near Bond's Store, 9 mi S of Macon (USGS 7696?). p. 69 Burke County GA-BU-1 5.0 mi W of Waynesboro on Hwy. 24, E side of bridge over Rocky Creek (86WA03; USGS 3935?). p. 47 GA-BU-3 Sloan’s Scarp, McBean Creek, between McBean Station and Savannah River, on Burke-Richmond Co. line (Veatch and Stephenson, 1911, p. 239) (USGS 5296). pp. 54, 69, 77 GA-BU-7 “МсВеап Creek, on land on Henry Byn" (Bowles, 1939, p. 287) (USGS 3288). p. 54 GA-BU-8 4.5 W of Hancock Landing, Savannah River (Veatch and Stephenson, 1911). p. 69 GA-BU-9 spring on Percy Boyd farm, 3.5 mi SW of Shell Bluff Post Office (Veatch and Stephenson, 1911). p. 69 GA-BU-10 “Dukes Bridge" —4 mi E of gouge on Waynesboro Ка. (*Barnwell" — ANSP collection). p. 69 Clay County GA-CL-2 left bank of Chattahoochee River at approx. mi 71.3 above confluence with Flint River; Nanafalia Fm. (Bed 16, Toulmin and Lamoreaux, 1963) (86WA27). p. 63 GA-CL-3 right bank of Chattahoochee River, near Morris Landing, approx. 7 mi above Ft. Gaines; “collected at waters edge during a low level of the river" (Bowles, 1939, p.298) (USGS 5478). p. 61 GA-CL-4 old road, approx. 2 mi E of Days Crossroads, /-/ mi W of Fairview School, 9 mi N of Ft. Gaines; Tuscahoma Fm. (USGS 12018). p. 86 GA-CL-6 exposure on east bank of Chattahoochee R. at mile 80.5 (above confluence with Flint R.); Clayton Fm. (bed 8 of Toulmin and LaMoreaux, 1963) (Toulmin, 1977 loc. Gcl2). pp. 61, 91 GA-CL-7 just above mouth of Sandy Creek, Chattahoochee R. (ANSP collection). p. 63 Glascock County GA-GL-1 English Plantation, 1.5 mi N of Gibson (Veatch and Ste- phenson, 1911, p.273). p. 69 GA-GL-2 “ss ledges at base of Barnwell near Rocky Comfort Creek” — Cooke, 1943, p.63. p. 49 106 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Henry County GA-HE-1 Chattahoochee River, Franklin Landing boat ramp; Nan- afalia Fm. (Toulmin, 1977, Loc. AHe-1; 86WA26). p. 63 Houston County GA-HO-1 Mossy Hill, 4.5 mi S of Perry (Veatch and Stephenson, 1911, p.258). p. 69 GA-HO-2 Perry-Elko public road, 4.5 mi S of Perry (Veatch and Stephenson, 1911, p.259). p. 69 Jefferson County GA-JF-1 Old Town rd., E side of Boiling Spring Creek, 7.5 mi SE of Louisville (USGS 52477, 74287). p. 69 Jones County GA-JO-1 cut on Ga RR 1 mi E of Roberts (Veatch and Stephenson, 1911, p.281); “McBean Fm.”. pp. 46, 47, 49, 69 Macon County СА-МА-1 Flint River at the county bridge, 1 mi NW of Montezuma; Clayton Fm. (USGS 3994). p. 61 Randolph County GA-RA-1 approx. 1.8 mi S of Stewart-Randolph Co. line, approx. 9 mi N of Cuthbert; MacBryde limestone exposed as blocks and in bed of dirt road on west side of hwy. (Toulmin, 1977, Loc. GRa-1; 86WA21). p. 68 GA-RA-3 "Green's Branch”, exposures along stream at small wa- terfall approx. 500 m W of Hwy. 266, 1 mi S of jct. with Co. Hwy. 15, access behind old farm barn, top of hill Bashi Fm. (GSA '80, stop 13, p.455) (86WA24). pp. Sl GA-RA-4 old quarries 1 mi NW of Cuthbert (Veatch and Stephen- son, 1911, p.266). p. 69 Schley County GA-SL-1 5 mi W of Ellaville (USGS 12092). p. 81 GA-SL-2 Walls Crossing, 4-5 mi NW of Ellaville. p. 68 GA-SL-3 Lumkin Farm on Goldendale Creek, 6 mi SE of Ellaville (Veatch and Stephenson, 1911, p. 235). p. 61 Sumter County GA-SU-1 50 foot bluff R side of Flint River, 16.5 mi E of Americus, near old Danville Ferry (Veatch and Stephenson, 1911, p.260). p. 69 Twiggs County GA-TW-7 Gallemore Station (Veatch and Stephenson, 1911, p.256). p. 69 Washington County GA-WA-4 2 mi S of Chalker (Veatch and Stephenson, 1911, p.276). p. 69 GA-WA-5 Little Keg Creek, 7 mi NNW of Sandersville (Cooke, 1943, p.55). p. 69 Webster County GA-WE-1 Lime Spring, Cole property, 2 mi SE of Preston. p. 81 Wilkinson County GA-WI-1 “farm of JW Huckabee, 1.5 mi N of Lewiston” (Veatch and Stephenson, 1911, p.278). p. 69 LOUISIANA Bienville Parish LA-BI-1 Holstons Well, 5 mi SE of Gibbsland (USGS 2033). pp. 51, 65 LA-BI-2 Mt. Lebanon (USGS 2911). pp. 51, 65 LA-BI-3 well at Rabun's place, sec 19 T 17 R 5 W (USGS 2037). prot LA-BI-4 sec 17 T 18 R 5 W (USGS 2034). p. 65 LA-BI-5 bluff on Hammetts Branch, sec 30, T 18 R5 W (USGS 2035). p. 65 LA-BI-6 well #1, sec 17 T 18 R 6 W (USGS 2045). p. 65 LA-BI-7 Hammetts Branch, 2 mi E of Mt. Lebanon (USGS 2400). p. 65 LA-BI-8 Hammetts Branch, SW 4 sec 30 T ISNR6W (USGS 2917). p. 65 Caldwell Parish LA-CL-1 Gibson Landing, Ouachita River 1b,1d 1 mi above landing. p. 46 1c at landing. p. 46 le % mi below landing. p.46 1g % mi below landing. p.46 1h 1 mi below landing (PRI loc. P8). p. 46 LA-CL-2 Bunker Hill Bluff and Landing, right bank of Ouachita River at bend, Columbia 15 min. quad., R.5 E, T.13 N, section 19 (85WA60, 86WA75). p. 46 LA-CL-3 Grandview Bluff, Ouachita River . p. 46 LA-CL-6 Wyant Bluff (PRI loc. P2). p. 47 Catahoula Parish LA-CA-1 Danville Landing, right bank Ouchita River at Catahoula- Caldwell Parish line, Harrisonburg 15 min. quad., T.11 N, R.5 E, section 15, ТЛО N, R.5 E, section 22 (85WA52). p. 49 LA-CA-2 Carter Landing, Ouachita River. p. 49 Claiborne Parish LA-CB-1 Pittmans Mill, SW М, SE Ys sec 19 T 19 R 7 W (USGS 2038). p. 51 LA-CB-2 secs 1,2,11,12, T 19 N, R 6 W (USGS 10794). p. 65 Franklin Parish LA-FR-1 well at Winnsboro (USGS 10456). p. 46 Grant Parish LA-GR-1 (Creole Bluff) Montgomery Landing, bluffs below ceme- tery, E side of Red River, Montgomery 15 min. quad, R.5 W, T.8 N, sec 20 (PRI locs. 883,10,15; 85WA49, 86WA76). pp. 46, 47, 49, 50, 75 la Montgomery Landing. p. 46 1d Y, mi below ferry (PRI loc. 883) . p. 47 Jackson Parish LA-JA-2 “from gullies north of old Quitman-Liberty Hill Road on Mrs. JM Turners place west of Madden Creek, sec 15,T16N,R 3 W, 3.6 mi W of Quitman RR xing” (TBEG loc. La-7) (Stenzel and Turner, 1940). p. 51 La Salle Parish LA-LS-1 “Urania well no. 1, drilled by the Simms Oil Co. at Urania” (Bowles, 1939, p. 307) (USGS 8931). p. 46 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 107 Natchitoches Parish LA-NA-5 Road cut along local road in center of NE М Ne М sec 19, T9N,R 8 W; Pendleton Fm. (Le Blanc (1942) loc. 59). p. 63 LA-NA-6 Road cut in NW М SE М sec 33, T 10 N, R 10 W, along dirt road leading west from L. Hwy. 404; Midway Group (Le Blanc (1942) loc. 8). pp. 81, 84 LA-NA-12 RR cut 75 yds E of depot at Marthaville; Marthaville Fm. (LeBlanc (1942) loc. 21) p. 63 LA-NA-14 about 50 yds N of La Hwy 1 in Roe's field, NW М, NE Y SW / sec 26, T9 N, R 10 W; Marthaville Fm. (LeBlanc (1942) loc 16). p. 63 LA-NA-15 road cut along local road, SW Y» NW М SE М sec 24 T 9 N, R 10 W; Marthaville Fm. (LeBlanc, 1942, loc. 15). p. 63 LA-NA-16 Pendleton Fm. (Le Blanc, 1942, Loc.55). pp 63, 86 LA-NA-17 Pendleton Fm. (Le Blanc, 1942, Loc. 56). p. 63 Rapides Parish LA-RA-1 well 2 mi NW of Boyce (Bowles, 1939, p. 277) (USGS 7743). p. 47 LA-RA-2 well of Cotile Oil and Gas Co., near Boyce (Bowles, 1939, p. 277) (USGS 7944). p. 47 LA-RA-3 Boyce Oil Co. Well, 6 mi SW of Boyce (1535-2766’) (USGS 7247). p. 75 Sabine Parish LA-SA-9 Undifferentiated Midway Group (Le Blanc (1942) loc. 1). p. 84 LA-SA-19 Marthaville Fm. (Le Blanc, 1942, Loc.37). p. 83 LA-SA-20 Pendleton Fm. (Wassem and Wilbert (1943) loc. 4). pp. 63, 83 LA-SA-21 Pendleton Fm. (Wassem and Wilbert (1943) loc. 5). pp. 63, 83, 86 LA-SA-22 Pendleton Fm. (Wassem and Wilbert (1943) loc. 7). pp. 63, 83 Winn Parish LA-WI-1 St. Maurice, Saline Bayou (USGS 2005, 2916, 4272). pp. 46, 51, 69, 78 LA-WI-2 c. 10 mi NW of Winnfield (USGS 2919). p. 51 MARYLAND Anne Arundel County MD-AA-5 South River, 2.5 mi from Annapolis (USGS 3934) (=? MD-AA-3). pp. 55, 85 Charles County MD-CH-2 Popes Creek. p. 65 MD-CH-7 Glymont, bluff on E side of Potomac River, 0.5 mi N of Potomac Heights. p. 55 Prince Georges County MD-PG-3 Upper Marlboro. p. 55 MD-PG-5 Piscataway (USGS 13451). pp. 55, 85 MD-PG-7 E bank of creek 100 yds NNW of reservation parking area A-A at Fort Washington Park, % mi NNE of Fort Washington (=? USGS 2370) p. 85 MD-PG-10 ravine, 1 mi E of Fort Washington (USGS 2029). pp. 55, 85 MD-PG-12 south bluff of Southwest Branch, just west of Capital Beltway and jct with Central Ave. (no longer exposed); Brightseat Fm. (Govoni (1983) loc. A). pp. 64, 90 MD-PG-13 banks and stream bed of small tributary to Cattail Branch, just south of Sheriff Road and east of Раса School and west of jet of Brightseat Rd with Sheriff Rd. (no longer exposed); Brightseat Fm. (Govoni (1983) loc. B). pp. 64, 90 MD-PG-14 banks of small tributary to Cabin Branch, just south of Central Ave (Md Hwy 214) just east of jct with Addison Rd.; Brightseat Formation (Govoni (1983) loc. C). pp. 64, 90 MISSISSIPPI Clarke County MS-CL-1 cuts on both sides of the Southern Railroad north of Wat- tubbee, T.3 N, R.14 E, sec 3 Archusa Marl Mbr, Cook Mtn Fm. (MGS Loc. 61,62; 85WA29,30) [these localities are probably at or near locality usually called **Wattubbee" or “Wattubbee Hills” by earlier authors—fide Dockery, 1980:14; USGS 338, 2616, 3092, 6479). pp. 51, 54, 69, 77, 79 MS-CL-2 in wash of Rose Hill Road, 8 mi W of Enterprise (USGS 2621; PRI 729). pp. 51, 54, 77 MS-CL-3 high bank of Garland Creek, about 1 mi. upstream along right fork, T.1 N, R.16 E, sec 28 (MGS Loc. 9; 85WA70; USGS 6471) [Garland Creek loc. of authors = USGS 330, 2630, 6471]. pp. 46, 47, 75 MS-CL-5 2-3 mi E of Shubuta. p. 54 MS-CL-7 McLeod’s mill, Suwanlovey Cr., 6 mi W of Desoto sta. on RR (USGS 14220). p. 54 MS-CL-15 left bank of Chickasawhay River, at bend just below old hunting lodge, Т.1 N,R.16 E, section 30 (MGS Loc. 16; 85WA72). pp. 47, 75 MS-CL-16 “wash of Old Enterprise road, at residence of Mr. J.Harrison Johnson, 8 mi S of Hickory and 12 mi NW of Enter- prise" (Bowles, 1939, p. 279) (USGS 2622). pp. 51, 54, 69, 77 MS-CL-17 “in a washed off place in Mr. J.H.Johnson’s farm, south of the creek and about 9 У miles south of Hickory" (Bowles, 1939, p. 279) (USGS 2623). pp. 51, 54, 69, 77 MS-CL-18 S bank of Chickasawhay River at bridge on Hwy. 45, 1 mi S of Quitman (USGS 14218). pp. 47, 54 MS-CL-19 bluff on old bridge site on SW bank of Chickasawhay River, De Soto (USGS 14229; MS-CL-4?). p. 54 Hinds County MS-HN-1 Jackson. p.47 1b “Greens marl bed” (USGS 54, 58). pp. 46, 75 1c “Jackson marl bed" (USGS 64, 3665, 13398). p. 46 1d “south of pump house” (USGS 734). p. 46 le “Bailey’s Marl Bed" (USGS 59). p. 75 MS-HN-2 Dry Creek, Jackson. p. 46 MS-HN-3 Town Creek. pp. 46, 75, 78 MS-HN-4 Moody's Branch (USGS 60, 63, 3735, 3746, 6462, 10377). pp. 46, 47, 75 4b Moody's Branch, first bluff below first bridge, east of the Institute for the Blind (USGS 4250). p. 46 , MS-HN-6 Riverside Park, Jackson, ravine behind old Riverside swimming pool, T.6 N, R.1 E,sect 36; Moody’s Branch Fm. (MGS Loc. 2; 85WA24). pp. 46, 75, 78 Lauderdale County MS-LA-3 McKays Marl Bed, Souwashee Creek, 2 mi S of Meridian; Bashi Fm. (USGS 2105) p. 73 MS-LA-4 “cut on Memphis and Meridian RR, 300 yards south of Seymours Hill, 1 % miles south of Meridian” (Bowles, 1939, p. 303); Bashi Fm. (USGS 7267, 7734, 10055). p. 73 Newton County MS-NE-1 “Indian mound railroad cut on the side of Pottoxchitto 108 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 swamp, 4 Y. miles east of Newton” (Bowles, 1939, р. 279) (PRI loc. 726; USGS 2620, 14058; MGS 67). pp. 51, 54, 69, 77, MS-NE-2 Hickory. p 51, 69, 79 MS-NE-3 2 mi NE of Newton on Rt 15 (PRI loc. 1803). p. 69 MS-NE-7 jct. of I-20 and Hwy.15, outcrop in grass behind gas sta- tion, southwest corner of intersection; Cook Mountain Fm. (85WA41; MGS 65). p. 69 MS-NE-9 “‘in wash of road from Newton to Decatur, about 3 miles northeast of Newton, in hill going down to Pottoxchitto Creek” (Bowles, 1939, p. 279) (USGS 2618). p. 77 MS-NE-10 “in wash of old Enterprise road, going down the hill to the Pottoxchitto swamp, 2 miles southeast of Hickory” (Bowles, 1939, p. 279) (USGS 2624; MS-NE-5?). pp. 51, 54, 69, 78 MS-NE-11 “in wash of the old General Jackson road from Nashville, Tennessee to New Orleans, 1 mile south of Hickory" (Bowles, 1939, p. 279) (USGS 2625). pp. 51, 77 Tippah County MS-TI-8 type section of Owl Creek Fm. (Cretaceous), Ripley 7.5 min. quad., approx. 2.75 mi NE of Ripley; Clayton Limestone exposed upstream from best Owl Creek exposures (86 WA69; USGS 64972). p. 80 MS-TI-9 ravine just N of Booneville Rd., 3 mi NE of Ripley (USGS 5586). p. 80 MS-TI-10 bluff on small stream approx. 1.5 mi S of Ripley (USGS 6496). p. 80 Yazoo County MS-YZ-1 Sims Siding, 8 mi N of Yazoo City, “roadside on hill at Sims Station, Yazoo & Miss. RR 10 mi N of Yazoo City, 2 mi S of Eden” (USGS 7674). pp. 49, 78 MS-YZ-2 Tcheva Creek, just upstream from Hwy. bridge, T.13 N, R.1 E, section 32 Moody's Branch Fm. (MGS Loc. 11; 85WA65). pp. 49, 50, 75 MS-YZ-4 Locust Grove Plantation, 1.5 mi S of Sartartia (Bowles, 1939, p. 277) (USGS 7393). p. 47 MS-YZ-5 Miss Lite Clay Pit at Cynthia, T.7 N, R.1 W, sec 25; Yazoo Clay (MGS Loc. 15; 85WA64; =MS-HN-7). p. 47 MS-YZ-6 near head of ravine, % mi SE of Free Run on line betw se Y sec 26 and sw М sec 25 T I3N, R 1 W, 10 mi NE of Yazoo City (USGS 10118). p. 78 SOUTH CAROLINA Aiken County SC-AI-1 Hixon Branch, Tinkers Creek, public rd. 200 ft. E of bridge, 12.5 mi W of Williston (USGS 4603). p. 78 Berkeley County SC-BE-3 Wilson's Landing, R bank of Santee River just below Santee dam; Black Mingo Fm. (only visible at very low water, 1985) (86WA115). pp. 55, 88 Clarendon County SC-CL-2 Kingstree Rd. E of Deep Cr., 5 mi E of Manning; Black Mingo Fm. (USGS 10403) p. 55 Lexington County SC-LE-1 “property of Eugene Senn, north of Kennerly road, Lex- inton-Calhoun county line, 18 mi north of house, 7 miles east of Swansea" (Bowles, 1939, p. 287) (USGS 7731) . pp. 54, 78 SC-LE-2 "south of old Bethel Church and east of Lexington-Or- angeburg road, about 2 miles southeast of Edmund" (Bowles, 1939, p. 287) (USGS 7732). p. 54 Orangeburg County SC-OR-1 3 mi WNW of Orangeburg. pp 77, 79 SC-OR-3 6 mi WNW of Orangeburg. p. 79 SC-OR-5 Old Columbia Rd., south of Early Branch, 5 mi N of Orangeburg (USGS 7722). p. 54 SC-OR-6 Poosers Hill, 5.1 mi N of Orangeburg (USGS 4580). pp. 54, 69, 80 SC-OR-7 Caw-Caw swamp, 2 mi W of Orangeburg (USGS 2009). pp.54, 69, 79 SC-OR-10 Stroman's Mill, 5 mi SE of Springfield (USGS 7816). p. 54 SC-OR-11 “at old brickyard in Orangeburg” (Bowles, 1939, p. 287) (USGS 2021). p. 54 Sumter County SC-SU-2 approx. 3.5 mi W of Pinewood; Black Mingo Fm. (USGS 10401). p. 55 Williamsburg County SC-WI-1 roadside, 1.1 mi S of Salters. p. 78 TENNESSEE Hardeman County TTN-HA-7a 1.5 mi Eof Middleton, 200 ft W of milepost 481 (USGS 6091). p. 80 TN-HA-7b 200 ft. E of milepost 481 (USGS 6495). p. 80 TN-HA-12 McDonald's Mill, 4 mi SW of Middleton (USGS 2148). р. 61 TEXAS Anderson County TX-AN-2 1.5 mi SE of Palestine on the Boston Rd. (USGS 10852). Do TX-AN-3 .5 mi E of Robinson on Centerville Rd. (USGS 13266). pool Atascosa County TX-AT-2 1 mi E of Jourdanton on the Pleasanton Rd. (USGS 13231). p. 50 TX-AT-3 % mi NW of Jourdanton (USGS 8870). p. 50 Bastrop County TX-BA-1 Colorado River, ~ 4 mi below Webberville; Kincaid Fm. (USGS 5280) 1c 5 mi below Webberville. p. 53 TX-BA-2 Colorado River at mouth of Dry Creek 2b Y. mi above Dry Creek (USGS 10526). p. 53 TX-BA-3 bluff on right bank of Colorado River, approx. 200 m downstream from Hwy 71 bridge at Smithville; Viesca Mbr., Weches Fm. (TBEG loc. 11-T-2; USGS 6088, 10386) [outcrop overgrown and inaccessible 1985, 1986]. p. 78 TX-BA-15 1.5-2 mi below Travis-Bastrop Co. line; Wills Point Fm. (USGS 11890, 11913, 12113). p. 53 TX-BA-30 Cedar Creek approx 4 mi SE of Williams Store (USGS 11909). p. 68 TX-BA-31 roadside gully Y. mi E of Williams store, Austin Quad. (USGS 11911). p. 68 TX-BA-32 approx. 3 mi SE of Williams store (Gardner, 1935) (USGS 11910). p. 68 Brazos County TX-BR-1 Little Brazos River, upstream from Hwy. 21 bridge; Whee- lock Mbr., Cook Mountain Fm. (85WA07; USGS 14210). p. 50 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 109 Burleson County TX-BU-1 Stone City Bluff (“Moseley’s Ferry), south bank of Brazos River just downstream of Hwy. 21 bridge, ~ 11 mi W of Bryan; Stone City Beds (85WA06; TBEG loc. 26-T-1; PRI loc. 723; USGS 5473). pp. 50, 73 TX-BU-2 [see TX-RO-10]. p. 73 Caldwell County TX-CA-1 5.5 mi N of Lockhart; Kincaid Fm. (= E.A.Moser farm approx У mi NW of Sims’ oil derrick, USGS 11707) (= ТХ-СА- 3). p. 68 Falls County TX-FL-7 roadside gully, % mi NW of Stranger School (USGS 11932). p. 68 Houston County TX-HO-2 left bank Trinity River, just upstream from old Alabama Ferry; access via FM 132 SW of Porter Springs (TBEG loc. 113- T-9; 86WA78; USGS 10738, 10735, 10736). p. 79 TX-HO-13 “Jose Procela League, from water well dug at village gin in 1916, Percilla” (USGS 9257). p. 77 Hunt County TX-HU-1 approx 14 mi SE of Greenville on Lone Oak pike approx. 50 yds from pike on E side of Cowleach fork just as road climbs hill on NE side on pike (USGS 10264). p. 68 Kaufman County TX-KA-1 Water Hill, 5 mi NE of Kemp; Kincaid Fm. (USGS 11665). p. 68 TX-KA-4 4 mi NE of Kemp (USGS 2440). p. 68 TX-KA-5 old Flat Rock Quarry (now flooded), approx. 5 mi S of Ola (85WA20). p. 66 La Salle County TX-LA-1 south side Hwy. 97, 2.8 mi. east of jet. with Hwy. 81 at Cotulla (USGS 14296; 86WA116). p. 72 TX-LA-2 2 mi SW of Shelton Ranch House, 2.5 mi SW of Millet (USGS 14298). p. 72 TX-LA-3 shingle below Brinkley Ranch house, 4 mi NE of Cotulla (USGS 14293). p. 72 TX-LA-4 1 mi SW of windmill near Altita Ranch house, 3.5 mi NE of Cotulla (USGS 14297). p. 72 TX-LA-5 № mi SE of Rockwood on Cotulla-LaMotta Ranch Rd. (USGS 14295). p. 72 Lee County TX-LE-11 between Orells and Price’s Crossings, Elm Creek. p. 51 TX-LE-12 “2 mile creek, first ford above 2 mile Negro Church”, 3-4 mi S of Middleton (USGS 14213). p. 79 Leon County TX-LO-12 ditch on N side of Concord-Centerville Rd, 0.6 mi SE of Robbins crossroad in S corner of JMPowell 100 acre tract, S corner of RMTyus survey (Stenzel and Turner). p. 51 TX-LO-13 bluff on Navasota River, near old iron bridge on E.C.Watson 800 acres, J.M.Viesca survey, SW part of county (Renick and Stenzel, 1931, loc. 8). p. 51 Limestone County TX-LI-2a 2.5 mi NW of Groesbeck p. 82 TX-LI-3 limestone quarry of D.P.Frost Construction Co., south side of Hwy. 84, west of jct. of Hwy. 84 and Hwy. 14 in Mexia; Te- huacana Limestone (85WA19). p. 67 Maverick County TX-MV-1 Bibora Tank, 18 mi S of Eagle Pass (USGS 6583, 11753). p. 82 TX-MV-4a land of Indio Cattle Co., 1 % mi below White Bluff, ~ 4 У mi SW of Jacal Ranch house 4b 3.5-4 mi SW of house, Las Islas crossing rd. (USGS 11763). p. 82 TX-MV-6 Indio Wells, 29 mi S of Eagle Pass. p. 82 Nacogdoches County TX-NA-7 road cut 4.8 miles east of Melrose (85WA55). p. 48 TX-NA-8 road side ditch on road from Chireno to St. Hwy 21 North of town (Stenzel and Turner). p. 48 Robertson County TX-RO-6 Dunn's Ranch. p. 51 TX-RO-11 Montgomery's well, У mile E of Wheelock (USGS 2048). p. 50 TX-RO-12 Taylors well. 5 mi SE of Franklin, 6 mi NW of Wheelock (USGS 2050). pp. 51, 78 Sabine County TX-SA-2b Y mi below Robinson's Ferry, center sec 6, T 3 N, R 12 W (USGS 10516). pp. 47, 50 TX-SA-3 Sabine River (otcp. 22 of Veatch, 1902). p. 79 TX-SA-4 bluff ~ % mi below Sabinetown ferry landing, ~ Y mi up from bridge (LeBlanc, 1942, loc. 40). pp. 83, 86 TX-SA-5 Pendleton, М mi down from bridge over Sabine R. on LA Hwy 6. p. 63 5c М mi above Pendleton Ferry. p. 63 TX-SA-19 (Le Blanc, 1942, loc. 51). pp. 63, 83 Starr County TX-ST-1 Alta Vista Ranch, 5.8 mi W, 35° N of Rio Grande City (USGS 13799). p. 47 Zapata County TX-ZA-2 1.1 mi NE of benchmark 400, Zapata-Hebbronville Rd. (USGS 14290). p. 72 TX-ZA-3 scarp on S side of Arroyo El Tigre above and below bridge on Laredo-Roma Hwy. (USGS 141165, 66, 67, 68). p. 72 TX-ZA-4 hillside У; mi ENE of Fordyce Ranch house (USGS 14291). рота TX-ZA-5 first bench above Rio Grande above Rio Grande south of Ramireno Ford (USGS 14292). p. 72 VIRGINIA Hanover County VA-HA-2 right bank of Pamunkey River, 0.5 mi. E of Wickham railroad crossing; Piscataway Mbr., Aquia Fm. (86WA106; USGS 26337). pp. 55, 65, 85 VA-HA-3 right bank of Pamunkey River, small ravine 0.63 SE of mouth of Totopotomoy Creek; Woodstock Mbr., Nanjemoy Fm. (86WA107; USGS 26403). p.66 VA-HA-4 right bank of Pamunkey River, small ravine 3-400 m downstream from VA-HA-3; Piney Point Fm. over Woodstock Mbr., Nanjemoy Fm. (86WA108). p. 66 King George County VA-KG-3 Woodstock. p. 65 VA-KG-5 Belvedere Beach, low bluff approx. 0.6 mi S of mouth of Potomac Creek. p. 88 Stafford County VA-ST-2 “Aquia Creek". p. 85 VA-ST-4 high cliff on western shore of Potomac River between mouths of Aquia and Potomac Creeks; Aquia Fm. (86WA99) (=? USGS 2347). pp. 55, 85, 88 VA-ST-5 Stafford Courthouse (USGS 12934). pp. 55, 85 VA-ST-6 bluffs at mouth of Aquia Creek (USGS 2030, 13809, 13810). pp. 55, 85, 88 MEXICO Nuevo Leon MX-NL-3 1200 feet N and 1300 feet E of Rancho la Rosita, in arroyo, Zacate district. (USGS 13512.) pp. 47, 72 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 MX-NL-4 275 m from Rancho Paredila toward Ranch San Agustin. (USGS 13763) p. 86 Tamaulipas MX-TA-1 15.9 km S of Cividad Camargo. (USGS 13504). pp. 47, 72 MX-TA-2 1 km N of Villa Neuva or Neuvo Camargo, Camargo Seder, La Mision. (USGS 13506). pp. 47, 72 RECENT LOCALITIES R-1 NW of Samar Island, Philippines. 17 fms. (MCZM). p. 101 R-2 Manila Bay, Philippines. 20 fms. (MCZM). p. 101 R-3 Philippines. (MCZM 111526). p. 101 R-4 Philippines. (MCZM 224596). p. 101 R-5 Near Cavite, Manila Bay, Philippines. (MCZM). p. 101 R-6 Darwin, Northen Australia. (MCZM). p. 101 R-7 Manila Bay, Philippines. MCZM 43097, 141712). p. 101 R-8 East Point, near Darwin, Northern Australia. (MCZM). p. 101 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 111 PLATES PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 1 Figure Page 1-3. ОВИЕ moriorni вер C Premon oni Govan; 1999). oer sy cer eese Е ee gue ar Fidei ees eite буы к» кэз 64 1. Scanning electron micrograph of apex. Unnumbered USNM specimen. Brightseat Fm., Locality MD-PG-12. x85 2. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. Same specimen as Figure 1. x 200 3. Holotype unnumbered USNM specimen. Brightseat Fm., Locality MD-PG-12. Height — 39.9 mm. ELO E E ОИ ШО ОЮ SO o DA E UNE dace One destin Wa che жар кели Sa Ей ote eel A O le Cee ev aei eo 54 4. Holotype ? ANSP 15508. Aquia Fm., Locality MD-PG-5 ? Height = 59.9 mm. 5. Scanning electron micrograph of apex. MCZIP 29320b. Aquia Fm., Locality VA-KG-5. x26. 6. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. Same specimen as Figure 5. x 140. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 1 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 2 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 113 EXPLANATION OF PLATE 2 Figure 1,9. Palmerella mortoni mediavia (Bowles) 1. Latex cast of ANSP 11884. Clayton Fm. ?, Locality GA-CL-7. Height = 71.5 mm. 9. Holotype USNM 495146. Clayton Fm., Locality AL-WI-9. Height = 41.0 mm. =). IPabmerellas ЛОНО РОУ ЕТ ОШО ЛЧ НЧА УУ ES AER ы зу уке Т" 61 2,3. Unnumbered GSA specimens. Nanafalia Fm., Locality AL-DA-1. Heights, Figure 2 = 91.2 mm., Figure 3 = 31.9 mm. (From Toulmin, 1977, pl. 29, figs. 12,13). Note double carinae. 4. MCZIP 29308. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 74.2 mm. 5. MCZIP 29329a. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 51.2 mm. Note double carina. 54 658. PalMerellaanoOMONIINONOHAKSCHTA) Ree с ОТУТ гел Л у ЫМ ГТ О И е е. 6. MCZIP 29099. Aquia Fm., Locality VA-KG-6. Height = 104.4 mm. 7. MCZIP 29307. Aquia Fm., Locality VA-KG-6. Height = 110.0 mm. 8. USNM 497988. Black Mingo Group, Locality SC-SU-2. Height = 57.0 mm. 114 Figure 10-12. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 3 Page IA UL RAI КС 5... en А оао a бин 33 1. Holotype USNM 498009. Lisbon Fm., Locality AL-MO-4d. Height = 39.5 mm. (From Stenzel and Turner, 1942) 2. MCZIP 29311. Lisbon Fm., Locality AL-CL-14. Height = 29.8 mm. Perl ЭЛ ЛО SUO IZ). c coi de ee ne eni en re re т 51 3. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. MCZIP 29322a. Weches Fm., Locality TX-RO-5. x 110. 4. Scanning electron micrograph of apex. Same specimen as Figure 3. x 32. 5. MCZIP 29309. Weches Fm., Locality TX-RO-5. Height = 25.9 mm. 6. MCZIP 29320. Bashi Fm., Locality AL-CO-1. Height = 24.6. „- РЕ НСК ШОЛ ea. (Stenzel-and ев) e Red нн АН aint am idees RI Cote a 47 7. Scanning electron micrograph of apex. MCZIP 29323. Weches Fm., Locality TX-NA-7. x 36. 8. Holotype TBEG 20960. Weches Fm., Locality TX-NA-8. Height = 19.5 mm. 9. Scanning electron micrograph of apex, showing protoconch and initiation of spiral sculpture. Same specimen as Figure 7. x 210 ТИКО even ШЫНЫЛЫ О quM qu one MM ne ee a 50 10. Hypotype PRI 2912. Cook Mountain Fm. ?, Locality TX-RO-14. Height = 26 mm. 11. Scanning electron micrograph of apex. MCZIP 29324. Lisbon Fm., Locality AL-CL-14. х 35. 12. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. Same specimen as Figure 11. x 180 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 3 PLATE 4 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 115 EXPLANATION OF PLATE 4 Figure Page 155, ДИЛЕ ДОСАН (Palmen). оо E 0 2. 49 1. Holotype PRI 4579. Moodys Branch Fm., Locality LA-GR-1. Height = 32.5 mm. 2. MCZIP 29315. Moodys Branch Fm., Locality LA-GR-1. Height = 24.5 mm. 3. Scanning electron micrograph showing protoconch and initiation of spiral sculpture. MCZIP 29325a. Moodys Branch Fm., Locality LA-GR-1. x 220. 4. Scanning electron micrograph. Same specimen as Figure 3. x 110. 5. Scanning electron micrograph of apex. MCZIP 29326a. Moodys Branch Fm., Locality LA-GR-1. x21. 6,7. -Palmerella. apud Cex OECROLIO) зы rio ныса Кз - 46 6. MCZIP 29290. Gosport Sand, Locality AL-CL-4. Height — 25.2 mm. 7. MCZIP 29347. Gosport Sand, Locality AL-CL-4. Height — 20.8 mm. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 5 Figure Page 1,2, ИЕР СВИСТОК costes Bae E UL СС к ллы аз RUN UR S ry eere ee 48 1. Holotype USNM 498010. White Bluff Fm., Locality AR-JF-1. Height = 24.1 mm. 2. Scanning electron micrograph of broken apex showing early teleoconch sculpture. MCZIP 29327b. Moodys Branch Fm., Locality LA-GR-1. x 140 3-6. Palmer еШ are (codon ООШ mu ee OR ITUR Pe rue а ее ti ie 46 3. MCZIP 29314. Moodys Branch Fm., Locality MS-CL-3. Height = 32.3 mm. 4. Scanning electron micrograph of apex. MCZIP 29328. Moodys Branch Fm., Locality MS-CL-15. x 30 5. Scanning electron micrograph of broken apex showing initiation of spiral sculpture. Same specimen as Figure 4. x 180 6. MCZIP 29316. Yazoo Fm., Danville Landing Mbr., Locality LA-CA-1. Height = 34.2 mm. NENA аа ON Cora EI LCD) sc A A A A A en КТ ОЕ re 45 7. Lectotype ANSP 13213. Moodys Branch Fm., Locality MS-HN-4. Height = 48.1 mm. 8. Scanning electron micrograph of apex. MCZIP 29325b. Moodys Branch Fm., Locality LA-GR-1. x32. 9. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. Same specimen as Figure 8. x 250. | | | PLATE 5 PLATE 6 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 147 EXPLANATION OF PLATE 6 Figure Page 123. Palmerellampleboides (Уай у Ее 65 1. Scanning electron micrograph of apex and early teleoconch whorls. USNM 138264. Cook Mountain Fm., Locality LA-BI-1. x 16. 2. Scanning electron micrograph of apex showing protoconch and initiation of spiral sculpture. Same specimen as Figure 1. x 230. 3. USNM 138264. Cook Mountain Fm., Locality LA-BI-1. Height = 17.3 mm. d64:Parlmerellasdumblei las) ЕКЕН ee lau ee 50 4. MCZIP 29317. Stone City Beds, Locality TX-BU-1. Height = 28.1 mm. 5. Scanning electron micrograph of apex. USNM specimen. USGS 1/742. Cook Mountain Formation ?, Locality TX-SA-2b. x32 6. Scanning electron micrograph of apex, showing protoconch and initiation of spiral sculpture. USNM specimen. USGS 13231. Cook Mountain Fm.?, Locality TX-AT-2. x 190. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 7 Figure Page TESSA АРА ONE ee сема а Т 22... 67 1. Syntype FMNH-UC 24522. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. Height = 35.0 mm. 2. Scanning electron micrograph of apex and early teleoconch whorls. MCZIP 29343. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. х 15. 3. Scanning electron micrograph of damaged apex showing initiation of spiral sculpture. MCZIP 29344. Porters Creek Fm., Matthews Landing Member, Locality AL-WI-3. x 120. ES ЖОШ ОПИС ПАРИС РУ Seed N ee eva, atl О л ао... 72 4. Holotype USNM 494993. Bashi Fm., Locality AL-CL-2. Height = 35.5 mm. 5. MCZIP 29345b. Bashi Fm., Locality AL-CO-1. x 150. ON SANS OR VERMESSCEHSB IUD deseo) ee ы, ee ee C t Rw ee 79 6. USNM specimen. USGS 6495. Clayton Fm., Locality TN-HA-7b. Height = 23.2 mm. 7. USNM specimen. USGS 6495. Clayton Fm., Locality TN-HA-7b. Height = 22.8 mm. 8. Scanning electron micrograph of apex and early teleoconch whorls showing initiation of spiral sculpture. USNM specimen. USGS 6495. Clayton Fm., Locality TN-HA-7b. x58. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 7 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 8 | | Figure 139. аитат ala (CONTAR ма E еко быт о AS $: ІС PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 119 EXPLANATION OF PLATE 8 MCZIP 29312. Moodys Branch Fm., Locality MS-CL-15. Height = 42.0 mm. 2. MCZIP 29313. Moodys Branch Fm., Locality MS-HN-6. Height = 25.2 mm. 3. Turritella lowei Cooke. Holotype USNM 353945. Moodys Branch Fm., Locality MS-HN-4b. Height = 23.0 mm. 4. Turritella perdita Conrad. Paralectotype ANSP 13232. Moodys Branch Fm., Locality MS-HN-1. Height = 37.7 mm. 5. Turritella perdita Conrad. Paralectotype ANSP 13232. Moodys Branch Fm., Locality MS-HN-1. Height = 35.0 mm. 6. Scanning electron micrograph of damaged apex, showing initiation of spiral sculpture. MCZIP 29340, Locality MS-CL-15. x 150. 7. Scanning electron micrograph of apex. USNM specimen. USGS 1466. Moodys Branch Fm., Locality MS-HN-4. 8. Scanning electron micrograph of apex. MCZIP 29342d. Moodys Branch Fm., Locality MS-CL-15. x28. 9. Scanning electron micrograph of apex showing initiation of spiral sculpture. MCZIP 29341a. Moodys Branch Fm., Locality MS- HN-6. x 110. 10. Haustator infans (Stenzel and Turner). Syntype TMM/TBEG 20957. Stone City Beds, Locality TX-BU-1. Height = 9.0 mm. ... 73 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 9 Figure Page ЕОС О VA ode 2. ee ne DY eee 68 1. Turritella carinata I.Lea. Syntype ANSP 5662. Gosport Sand, Locality AL-MO-1a. Height = 28.1 mm. 2. Turritella carinata 1.Lea. Lectotype ANSP 5661. Gosport Sand, Locality AL-MO-1a. Height = 27.9 mm. 3. Turritella carinata 1.Lea. Hypotype PRI 2893. Gosport Sand, Locality AL-MO-1a. Height = 48.0 mm. 4. MCZIP 29293. Gosport Sand, Locality AL-WA-6. Height = 48.5 mm. 5. MCZIP 29292. Gosport Sand, Locality AL-CL-4. Height = 36.9 mm. 6. MCZIP 29294. Gosport Sand, Locality AL-MO-1a. Height = 43.9 mm. 7. Turritella carinata 1.Lea. Hypotype USNM 497995. Gosport Sand, Locality AL-MO-1a. Height = 39.5 mm. 8. Scanning electron micrograph of apex. MCZIP 29339a. Gosport Sand, Locality AL-MO-la. x24. 9. Scanning electron micrograph of damaged apex showing initiation of spiral sculpture. Same specimen as Figure 8. x 160 10. Turritella carinata palmerae Bowles. Holotype USNM 497997. Lisbon Fm., Locality AL-MO-4. Height = 29.0 mm. 11. MCZIP 29295. Gosport Sand, Locality AL-CL-4. Height = 35.6 mm. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 9 PLATE 10 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 121 EXPLANATION OF PLATE 10 Figure Page 154557 Haustator: АРА Тет) СЕ ек АНАСЫ а К 76 1. MCZIP 29335. Upper Lisbon Fm., Locality AL-MO-1b. Height = 50.2 mm. 4. Scanning electron micrograph of apex and early teleoconch whorls. MCZIP 29336. Lisbon Fm., Locality AL-CL-14. x14 5. Scanning electron micrograph of broken apex showing initiation of spiral sculpture. Same specimen as Figure 4. x150 2. Haustator subrina (Palmer). MCZIP 29304. Upper Lisbon Fm., Locality AL-MO-1b. Height = 58.7 mm. ................... 78 3. Haustator rivurbana (Cooke). Unnumbered MGS specimen. Moodys Branch Fm., Locality MS-HN-3. Height = 30.3 mm. .... 78 6-8. F'urFUelldsoXOIe NUES) er ero centenis Ено а уз s NE aia do 26 6. Unnumbered specimen in the collection of the Rijksmuseum van Natuurlijke Historie, Leiden. Recent. Guyana Shelf. Height — 46.7 mm. 7. Scanning electron micrograph of apex. MCZIP 29337. Recent. Gulf of Mexico. x 20. 8. Scanning electron micrograph of apex showing initiation of spiral sculpture. MCZIP 29338. Recent. Gulf of Mexico. x60. ОАО COSER BON SS N. сансара не dat d dL RR лы МО О орке UL а 70 Holotype USNM 495172. Cook Mountain Fm., Locality MX-TA-1. Height = 30.0 mm (from Stenzel and Turner, 1942). ТО Haustatorfisehenakalmen) ee ont а лы nl PER EMEN NN a 25... 72 FLMNH 5411. Inglis Fm., Levy County, Florida. Height = 19.6 mm. 122. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 11 Figure Page 1. “Turritella” plummeri Stenzel and Turner. Holotype FMNH-UC 57373. Kincaid Fm., Locality TX-KA-7. Height = 36.5 mm.. 37 2. “Turritella” ola Plummer. Holotype FMNH-UC 31281. Kincaid Fm., Locality TX-KA-7. Height = 36.0 тт................. p 3. "Turritella" kincaidensis Plummer. Lectotype FMNH-UC 31280. Kincaid Fm., Locality TX-KA-5. Height = 25.0 mm.............. 37 4. Palmerella levicunea (Harris). MCZIP 29277. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. Height — 37.6 НЫЙ aca i IAS Ku MEME I oe c ТТТ DL сл UON. OUR e y uba US 513 5. Palmerella sp. MCZIP 29280a. Nanjemoy Fm., Woodstock Mbr., Locality VAGH AGS. Herit: 202 IHE 222222222222 22. 66 6. “Turritella” polysticha Stenzel and Turner. FMNH-UC 57371. Wills Point Fm., Solomon Branch Mbr., Locality TX-BA-34. Height Of arees spee ioni = 12 2 TUI 22... ee a 00. UE ue nice 57 7. “Turritella” saffordi Gabb. MCZIP 29289. Clayton Fm., Locality MS-TI-8. Height = 32 mm. Latex cast. .................... 3 8. Palmerella hilli (Gardner). Holotype USNM 373054. Kincaid Fm., Locality TX-BA-2b. Heisht = 35. ои 52 9.10. РИО еер пея species: ONE сызу M а en nee па бан 66 9. Holotype MCZIP 29270. Kincaid Fm., Tehuacana Limestone Mbr., Locality TX-KA-5. Height = 33.4 mm. 10. Paratype MCZIP 29273. Kincaid Fm., Tehuacana Limestone Mbr., Locality TX-KA-5. Height = 44 mm. PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE 11 PLATE 12 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 ақышы А! A Y \ м \ TON Figure 1-4. 5, 9-11. PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 123 EXPLANATION OF PLATE 12 TITTEN BON SS КО Жл. id 81 1. Scanning electron micrograph of apex, showing protoconch and initiation of spiral sculpture. MCZIP 29333a. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. х 140. 2. Scanning electron micrograph of apex and early teleoconch whorls. MCZIP 29334a. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. x 14. 3. MCZIP 29305. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. Height = 38.8 mm. 4. Turritella aldrichi Bowles. Paratype USNM 495149. Porters Creek Fm., Matthews Landing Marl Mbr., Locality AL-WI-3. Height = 41.0 mm. Мили МО ао О о ЕРОТ ME 85 5. Syntype FMNH-UC 24521. Nanafalia Fm., Locality AL-MA-1? Height = 43.5 mm. 9. MCZIP 29334a. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height=47.3 mm. 10. MCZIP 29334b. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height=45.8 mm. 11. Unnumbered GSA specimen. Nanafalia Fm., Locality AL-MA-1. Height = 105.9 mm. РОЛЕН еу реве ene ee en 2 are M co yi 90 6. Plastoparatype MCZIP 29276. Clayton Fm., Locality AL-BA-1. Height = 56.6 mm. 7. Paratype MCZIP 29283. Clayton Fm., Locality AL-BA-1. Height = 32.9 mm. . “Turritella” claytonensis Bowles. Holotype USNM 131637. Clayton Fm., Locality AL-WI-9. Height = 68 mm. (from Stenzel a ae оа ча. 82 124 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 EXPLANATION OF PLATE 13 Figure Jc TE УУ ООО ООШ ООШ S ТТ аи кыйкйшк RI Бов eg PE | о е 1. Paratype ANSP 31388. Aquia Fm., Locality MD-PG-6. Height = 90.2 mm. 2. Lectotype ANSP 31388. Aquia Fm., Locality MD-PG-6. Height = 89.6 mm. 3. Scanning electron micrograph of broken apex showing early teleoconch sculpture. MCZIP 29332. Aquia Fm., Locality VA-ST-4. x 240. 4. Scanning electron micrograph of broken apex. Same specimen as Figure 3. x 80. 5. “Turritella” sp. (“prehumerosa” Govoni, 1983). Paratype, unnumbered USNM specimen. Brightseat Fm., Locality MD-PG-12. LHe dace dicii) у до O, ГЕНЕТИК НК КИС ал та O ос 63. “ЛОПЕ КОШО SNA NUT AUC Ca а т AM ee ао ПОЕ 6. Syntype FMNH-UC 24505. Tuscahoma Fm., Bells Landing Marl Mbr. 2, Locality AL-MO-3 ? Height = 27.5 mm. 7. Unnumbered GSA specimen. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 86.4 mm. 8. Turritella bellifera Aldrich. Holotype USNM 644614. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 83 85.0 mm. PLATE 13 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 PLATE PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 125 EXPLANATION OF PLATE 14 Figure 1-4,6. “Turritella” praecincta Conrad КЕБЕ RM M ҒТА ТС ы. СТУ ОРЫ А UU NE 86 1. “Turritella” praecincta praecincta. USNM 559308. Tuscahoma Fm., Greggs Landing Marl Mbr. Locality AL-MO-5. Height = 76.0 mm. 2. “Turritella” praecincta praecincta. MCZIP 29349. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 67.7 mm. 3. “Turritella” praecincta praecincta. MCZIP 29280. Tuscahoma Fm., Bells Landing Marl Mbr., Locality AL-MO-3. Height = 98.3 mm. Note double carina. 4. “Turritella” praecincta virginiensis n. ssp. Holotype MCZIP 29093. Aquia Fm., Locality VA-KG-5. Height = 48.3 mm. 6. “Turritella” praecincta virginiensis. MCZIP 29330. Black Mingo Group, Williamsburg Fm., Locality SC-BE-3. Height = 48 mm. Latex cast made from specimen in collection of USGS. 5. “Turritella” biboraensis Gardner. Holotype USNM 370989. Kincaid Fm., Locality TX-MV-1. Height = 42.4 mm. (from Stenzel and Turner T942 uou er LINK Уз ncn. ho Lu 2 000 205065422 82 7. “Turritella” nasuta Gabb. MCZIP 29318. Lisbon Fm., Locality AL-MO-1b. Height = 41.3 mm. ........................... 37 126 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 INDEX Note: Page number in italics indicates a Table or Text-figure; page numbers in bold indicates a Plate; page number underlined indicates principal discussion; a lower-case “n” following a page number refers to a footnote on that page. Localities are indexed in Appendix 2 (p. 103). abrupta Spieker (1922 | Turfiiella a оз. nee 55 Academy of Natural Sciences of Philadephia (ANSP) ........... N... Е 7,44,45,46,55,69,75,84 acropora Dalt (0889) tele... ee DO Pd) cp аР о ИН 67 CHIU ОНЕ ТОЕ ск 10 adabionenesis Oppenheim (1915), Torquesia ............. 33,34 Adams and Reeve ӨЗА. cese uneven an 68,85 НОВО (ТОБТО чл. ils bc eee e en oe ERR 10,12,24,26 еро То” Иа НЕЕ en eter n ae 10,33,34 aeguisiriata Conad (1856), Turritella e die 18 Sello, IE оа ato NU ORE s 7 alabamiensis Whitfield (1865), Palmerella ..................... beset те ин чек 16,18,26,31,37,38,39,40,41,67-68,73,82,7 alcida Dt НО О) ТИЛЕЙ s. uev. dte ETUR 18 Altero SEED ықы B ирк 83,85 AEE (Байы Е а eren 54,61,76,81,83,84,85 Vd iul Bee DEMNM C e: 83 Altesch СВО C ree анинин 67,72,78,81,83,85 ЗЕ (О ВЕ cys E E MM c Күсе ы 79 aldrichi Bowles 1939) PUT” y coo vee oes ea 2. NA Ха” с + ск 16,18,27,31,37,38,39,40,41,68, 80,81-82,81,82,84,85,91,12 Allen. Jo lr Wen ae 7,44,50 ARDO (IIO Е 9,10,12,22,24,27,44,45,46,47 Alison КОЛ Е ne 9,12,13,17,21,22,24,26,30,44 Allison and Adegoke (1909) ee o et ba. ds Ame S dcs: 9,12,13,16,24,26,27,40,42,44,70,72,74,76,78,79 АИО ООВ ev Lco edu OR кл си 8.13.32 ООО е re 8,9,17,19 rs lan can ОВИ Pare бн ы rar ap De E an SU 8,15,16 RE MOS ON) лғы МЕН rg аа Mere 15,16 Algo е) еле С 200 13,14,15,34,35 Atom СУВИ АЈ 22 22. КЫН ee ei 516 Almom Ор: o 5 spe nn ене woo 9 Аи (19945) лл wn О зз е» 8,27,30,32,33 Allmorksind Dockery ООД) eso mis УУ 8 Almos and Ко (1993 И оон тнай 8,9 А тоне Jones, and. Vaughan (1992) Ta... 8,9,17 Allmon, Jones, Aiello, Gowlett-Holmes, and Probert (1994) .. 8,9 Alto. Niel and Norris (ПОЗОВИ оо оао анн 33 alowensis Hodson (1926), Ти ШеЙа со ,.................... 575) И LI Le eur dx НЕ = 12 alticostata; Conrad: (1834), БИРОА u.s rr ener he TTE 18 amiro CORR (1857), THETA 20222222222... 5,30, Supo Фасотавніт Оо. 55 mmama Hodson (109 o) een, orca ee eek kc ok а rr en 55 mrumacoensis Hodson ООВ Jo аттана эрекет (1922), еа ooer ion Soo ERI Do diveata Conrad (1854), Ralmereln zer... cn oer 16,18,25,31,33,34,36,37,38,39,40,41,45—46,50,76,5 amaras Woodring (1937) PO a a 35 AED ICONS A CD оаа ча 10,21 andersoni Dickerson (1916), Turritella ..................... 36 andreasi Williston, in Hodson (1926), Turritella ............. 302 ао ао иеа 55,61,86 ООО Е ilr 29 angulliana. Cooke (1919), Turritella ....................... 55) apita ав бтевопо (890), Balmerela nn nn н 16,18,31,37,38,39,40,41,46,47,52,54,4 Aquia Formation ............ 16,38,41,55,58,60,65,81,85,88,90 АРОН ТЕО e Sac nA от 10.1229 arenicola Conna (1965); Lalmerellane т И тео 16,18, 25,31,37,38,39,41,46—-47,49,50,54,5 a pranneri АО or a e e 37,47 a. danvillensis Stenzel and Turner (1940) ......... 18,31,37,47 БАО РУЛЫ een aca BTE e 10,12,29 Baird(1870) m Pe m merecen e BE E vr 10,12 Bara well FORMA. а. 74 Bashi Маш Formation ................... 16,38,41,48,51,52,73 bayovarensis Olsson (1932), Turritella ...................... 35 Beauchamp 934) tt RUN TES s 86 Beerbowenmel968) o cce ss c EO REN Certes 13 bellifera Aldrich (1885), Turritella .............. 31,40,83–84,13 Bell’s Landing Marl Member (Tuscahoma Formation) .......... ne Abo Ce 0-02-54 38,63,83,84,85,86 DENA Eu ee. 7 Borgere аце Molister ОГОЛА) es 34 Berggren, Kent, Flynn, and van Couvering (1985) ........ 16,38 berjadinerisis Hodson (1926) Turritella ann. 33) bibotaensis Gardner (1945), < Turritella er AT M ER о Те ае 16,37,40,41,80,82,82,90,14 Bieleraudokladiielde ООО) ст еке ee 17 DASS AAN Onis OTURA с ки Sio) b. cartagenensis Brown and Pilsbry (1917) ................ oo) PROTEIN c 35 Diiira ер Нено пи (ПОЧ ТИ ШЕКЕ en er. 18 Black Ming: Group... rn ee 38,41,55,57,88 Гауе СТАУ Е АИ NE d E 12 DIO Wie НТК... y Boardman, Cheetham, and Cook У Он ds ig BOSSE, долана ЕЛАНА кл DEAE es aro Y oy if ОИЕ 99у а. EET STD EORR dn een З AD 12,18,21,22,30,37,40,42,44,45,46,49,50,51,52,53, 54,55,57,61,63,65,67,68,69,70,72,73,74,76,78,79,80, 81,82,83,84,85,86,88 Peano ana Ront (4960). A пара SEN. nn vu NO 45,46,47,49,50,53,54,61,63,65,68,72,73,74,75, 76,78,79,85,86 ВУ (01192 0). ы. ТЕ 61,72,81,83 brazita (see nasuta) Тена а у етл Кс... 38 A 79). 77S A ee io И 39 Варе Formation ои ce ни 16,38,53,55,64,81,90 broderipiana d’Orbigny (1840), Turritella ................... 36 buchivacoana Hodson (1926), Turritella .................... 39 0 canonensis Hodson ТОО 35) ЛАОС ИО ОВО ОТЕ... 35 ОЕЕО бО Соб) ТЕ So b. socorroensis Hodson ТГ. 55 a CLO ZO). Е 35 bunkerhillensis Palmer in Harris and Palmer (1947), Turritella ... PAE MM Ee Rae ee Mn 37 burdeni Tuomey and Holmes (1857), Turritella ............. 18 buwaldana Dickerson (1916), Turritella .................... 36 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 197 Gans SA na I и ee 14 Caldo RE TA 32 Caleta Olsson (198 Zunrnella EA EORR EE PER 35 Gall (189i Rena SHE MESE e 46,48,49 Gallos ao RR АРЫ О ds 10,12 Garin bell О КИ ee МАС НИКИ 44,46n Campbell (1995)p- te rar a Bee re 46n Cane Rıyeribonmalion a... o een И 48 canonensis (see buchivacoana) caparonis Mautya(l92S)lürntella ee see 35 carinata Beas( ТОТО S ran A . 16,18,20,23,26,31,37,38,39,40,41,68—70,70—71,78,101,9 сораппелае ВО Ле Зо e no аи је 37,68–69 @avarepraccarimata tars. Cooke: Озо 76 carinifera claibornensis de Gregorio (1890), Turritella ........ 68 Саптап Mii. EEE сат КӨКҮЛ, 9 carota MacNeil in MacNeil and Dockery (1984), Turritella ... 49 cartagenensis (see bifastigata) Carte (ШО неи ле AT ME КЁ АК. А ЖЫН o аад 41 Gastle Hayne Тшей INN e 38,79 cauredalitoensis Hodson (1926), Turritella .................. 35 Geriihideas Ken ee een 20 (бекішенее A de nee 20,21 Eat толты 06 0 8L 1 а 13 chiapasensis (see rivurbana) chicoensi та ВИШУ Od) ы... 0: 36 (hin(194 уа А ОН oret) aa ioo RM 57 chipolana ОР ИУ ОЈ) Жей a a A 18,35 chirena Stenzel and Turner (1940), Palmerella ................. STE Um ver, АЕ 18,25,31,37,38,39,40,41,47-48,3 chiriguiensisOlssonid 922) и Turrel m t van ee 35 оО) С ОО ЕЕ c S ДЕ 57 cladistic analys ist a A аа detiene 36—40,40 claibornensis (see carinifera) Glarke dsi Soke Brit el tappe xecdatea AA 45n САВО e ee бине EET 83 КЫЧ И 02 а A A EUA 54 ыды с р дуу и хы Жеты 84 Clark and Martin (1901) .................... 37,54,55,61,65,84 Лака Mer IN па асет ODER хан 54,65,84 Clayton Formation ........... 16,38,52,53,55,61,67,80,81,83,90 ClayionensisBowles (1939) “Turritella wenn... ARA O 16,37,38,39,40,41,80,82,82,83,12 cleveland Нах (LS) IN EUA O ES A SAE dr 16,18,25,31,37,38,39,40,41,48—49,50,5 ОЕ ТЕ Т ЛЫ а EE ERE 20 cocoditana (see buchivacoana) (Са Mejttow iter: o. tote Vere ee 10 COUPE NR e E а Okt NAE 10,18,29 ӨСІП ЕНТ СӘЛ ТЕ es 1.012,29 (СООЛ А u ne. ae e RS 10,29 colinensis (see buchivacoana) columbiana Weisbord (1929), Turritella .................... 35 COMMUNISSRISSOHASZ2O) ПЛ. 18 conquistadorana Hanna and Israelsky (1925), Turritella ...... 35 (волта 8 AO erene a 18,37,49,53,54,55,61,67,68,76,79 Conrad CIS 8d) Sis на АЕС 18,54 ова ОВ и r er aa 37 Gonrad (ISS Sana NEAN E ооо ИЕ 54,68 Сойган ОО) алав. 18,37,63,81,82,83,84 ЛЕТТІ ТТТ ne ee 18,74 Солта (S46) pk сы лы Nasa 84 Conradin Waes (LIIR И МДЕ 18,37,45 олха боо) Toon du Е Ы Еа a c 18 Cond ааа on clit eR 45 C ИВЕ ОЕ NE s cor ee uri an EN RUNI DAR 18 Gontadkl 800) en ei. Ia LL Ыы Т, 18 (ола Е dE GR SE ЖА КЕ. чу es LL ы 18 Gone (364) зет ызы ыт aime ad 37,86 «ола Босат, лесе. conan EX 00. 45,54,68,84,86 (волта Оо) соны салатын ете 18,37,46,74 Conrad 566) қы заты Йа. 46,68,74,84,86 GookeMountamsbomnationy east. nee ar 16,38,41,46,48,50,51,52,54,65,68,69,72,73,77,79 соке) dud Tocem cL een Sa а 74 EOOKE-MI2H) oo ii o 37,54,61,74,75,78,84,85,86 A а и Oo. A a, 61,76 EOI) Bra ossi Raute 44 cooper Carpenter (1864) Turritella 252110222... 36 GODER ЕТ re т 33 cornellana Hodson (1926), Turritella н м, 35 cortezi Bowles (1939), Haustator ....... 16,37,38,39,40,41,70-72 коју па (ISOS) ea oir Le dU sco oe ER 54,67,68,83 Cossman (12) eaae 10,12,33,45,49,67,68,83,84 ООО сы жасы Жән ee ae ыы 10 costaricensis Olsson (190 2) Tortella екен sense. 35 Go ront ЭВ аа E S ылды 11 СОНО У CAS IS a ae A GOs (LOGO) Re ge eee се ee a T каш 30 СОБІ а Il бе UI d 10 creola Palmer in Harris and Palmer (1947), Palmerella .......... О Е а 16,18,25,31,37,38,39,40,41,49-50,50,52,4 аво О ee 36,37,40,43,43-44,48 CASA ht пат сва cv eR nae ЖЫ њи 5 10 GHOCH OO KG 010): ТҮЛЕН oc. ctu cs uc RB Men 35 eruzianaOlsson (1932), Turritella nn... 35 GiystalaRi ver FORMON E eu рынка hah a 74 GienocoilDissme ue Ve A ee St Te Da E T ЕС 10,12,29 КОШО НН teas here н А ee smithvillensis Harris, in Van Winkle and Harris (1919) ..... 48 lisbonensis Harris, in Van Winkle and Harris (1919) ....... 48 Cummins, Powell, Stanton, and Staff (1986) ................ i5 curamichatensis Hodson (1926), Turritella .................. 25 C Ы ee 45 DI СУВИ ек nr er ee 46,48 а (18923 ER anne bns nt 18,37,74 ШЕ GUESS о ates cal ens 222226226 18 МӨШ( E e ү ES 11 Шапан азат Tult ПЕНЫ ae a 68 Шана в Фо СЕО. uu А ee ae ЕУ eee 68 Danville Landing Member (Yazoo Formation) ........... 47,49 danvillensis (see arenicola) Davies: Baines, and Savage (1975) rr Puedes. en ee 34,35 declivis (Adams and Reeve, 1848), Gazameda ............... 21 ЧЕКИШ O A о еН iot y ЧӨН АХ ЗН ӨЗӘ nn. en uu па О е 7 68 dobyensis Dockery (1980), Turritella ............ 27,37,38,39,46 ШОРУ КБИ longues 502. са ал or ue 38,41 ERORE T сы аға EIS etae аа 7,78 Dackel O7 mes at. 44,45,47,49,74,75,78 Dockery GOSU у. eee ree m 27,37,40,44,46,47,68,69,73,76 DICKE ШОВ). ое маса EDS DONT: 35 IDO CSS) тес LES d РЕВА 222224 36 Dackerysand Nystrom (1992)... o ae 46n Doms О ерли REN. аа kr IN 12,28,30,42 domingensis Brown and Pilsbry (1917), Turritella ........... 35 она Аа ms ccc АН хе лл» СЫЛАЙЫН 10,12 128 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Dow (DA verein 76 ТРОНЕ (0929) e naar 10,30,32,33,37,42 аренах (1895); Palmerella--v-y a Bocce ta на ae A AAA 2274 16,18,25,31,37,38,39,40,41,48,49,50,6 duplinensis Gardner and Aldrich (1919), Turritella .......... 18 Eee en ER Тар 7 пиа Hartıs (1895), Барпекецарғе 2. ж. те аз ни та нил eg as на 16,18, 25,31,37,38,39,40,41,47,50-51,51,52,54,3 а ШЫ SDA erat, ea ese cee 10 АЕБ CUO OU) ee re 10 a ы P Ine РУ 48 BE ОНУ ИТ аи НЕ ен ee 33 Bidredgesand: тастан (1980) 2... 8 22. вики уже eee ne bee 14 Беке ап Gould (1972) mm Ae ae ene 13 elemensis Hodson (1926), Turritella |... eee 35 (MC gea ceu cede RITE een 48 БЕРЕД н, o CM пе а m CE ГҮ, 7 Гаре И сода (Ci OE aa wen cine cre eo T ee 5 ЭЛЕ ИЕККЕ ЕЧ a I M RR aR c rs 14 eterina de Gregorio (1890), OUO воо tS 68,69 etiwanensis Tuomey and Holmes (1857), Turritella .......... 18 eurynome: Withheld: (1565); “Turritella тя mo ET EN 16,37,38,39,41,81,83-84,83,85,86,13 THO LOK er СЕР ARE сес ку ра 10 exoleia.Emndeus1758), ОРО оноо и 26,33,10 Jalconensıs Hodson (1926), Turritella a. vehe teeta nn 35 БӘЛЕ EA IU SUUS cr LN ERIS NUR TER ел ы 7-8 jaw add Ртсе( орта алыл EAS AREE he e А? 46n TOSS o Ол ТТТ е O е TU 57 felli (see nasuta) femina Stenzel in Renick and Stenzel (1931), Palmerella ........ Тек өзі 16,18,25,31,37,38,39,40,41,47,50,51-52,54,66,3 f. oligoploka Stenzel in Renick and Stenzel (1931) ...... 57751 Л јен ШОВ) RUE eS eA ULT S eem ГМ 35 Field Museum of Natural History (FMNH) .... 7,44,67,68,83, 22 filicarmenensis Hodson (1926), Turritella ................... БЕУ ШУ ОЕ, A аана er ness 10, iiio Elia 192... ci sete ТТЫ РСЕ NS 21 O D. chu e UL ME EcL е 10:21:22: 74 Finlay and Marwick (1937) ан нА TUS DIST fischeri Palmer in Richards and Palmer (1953), Haustator ....... Verum A O сс 16,37,40,41,72,72,74,78 НО ШӨЛДІ O PO MT зе АЙАЛЫ, 45 EIER IST ee rennen EENAA 19 КЕШ YO nt ЫЗ nee N 19 О ПОО) АЕ REN eno, SI 19 A E ee a ОРЕВ 19 Eloada Geological Survey (FOS) . 208 РИБ Ран 7,44,72 Florida Museum of Natural History (FLMNH) .......... 7,45,72 Ones MSG SANA ne o RAE ROE 74 FONOS BONOA ЗУ Еа. ann er UO Bone ишы climes: 992) e. mer LLLA re 15 Freiter- aie. Graham. 01962)* cido EV ees АДУ, 17 Prener and Mans ШЫЛЫ en am ee EE NS 18 Mena Kelta COLO) ена S A ERI 18 Jerome Sdevoke (1977) HOWSIQUOT. ar een ECT ee 34 ОЛО Mawwick (1931), ZZ eAGOIDWSU ы на ои RE Re EI 21 СЕ КООШ E LL NET 18,37,53,61,79,80 Sanes NBO CLO SL), Turrel Vs nn. een Aven ee 35 Garden 00928): a ea ЕРА ERIS NINE 18 Barden ОЗ tata en eC er o АН A 34 Gardner (1995) тои 34,37,44,52,53,67,82,84 Card (E545) ақысын ы tod sexe atte bre UA Rm wn NETT HRS 57 Gardner (од) ie eee ce Pt wae SETS 10 GardBerand»Aldrich(19 10) 4 өзе pet en ea n tu 18 gardnerae LeBlanc (1942), “Turritella” ....... 37,40,41,80,84,84 Gut Are (ДӘУ уе c cenae tete en на Е 2130,57 САУТ QE N AE) 7 gatunensis Conrad (1857), Turritella б GI OTTe SIS Mans teld (1925) сл MT ғ” ED ШУ ШОЛОЙ СО ААД ЕЗУ Nune Y s B EA AOUSOM CL O MEN EN 55 gaweaveri Hodson (1926), Turritella ....................... 35 GEIGER UE OA A Ne 10,12,29 PEUS CEMO A ER 13 Geological Sutvey of Alabama wen. ns 7,44,45 ghigna de Gregorio (1890), Turritella ................... 68-69 ИРЕНА Hodson (Тоу ела TT 33) РИБЕ Bowles (1939); Ж ОННО ТТТ oe ae 16,18,26,31,37,38,39,40,41,49,52,72-73,74,7 CEA ee ТӨН ОППА 39 gittosinus (Powell and Bartrum, 1929), Tropicolpus (Amplicolpus) Mim dor A E re АЕ tM УЛ А сыы. 21 (УСНЕ (IO SON eG EA ste wna epos К eee NIE 35 СВОИ) ИЕ ОАА 46,49,68,76,78 Ghberbelo79) тое ee ОА 33,39 ОТРОВА тан о a RE ES ENTE 71510729 gonostoma Valenciennes (1832), Turritella .................. 18 (БОРО S АЛ PUSH 16,38,41,46,51,54,68, a da ELLE А аа ЭРЕ; (ХОШОЕТУӨБУР NC UR а КЕШУ элт EN AN SPA ES ULIS: Y Govoni (1983) .......... 18,30,37,42,44,53,55,57,64,81,88,90,91 Govoui and hansen Gi Pross) r rri cd O A ERREUR UMEN ИК 30,37,42,44,53,55,57,64,81,88,90,91 (Старао а а Shimer (1909) mine... па quu ТИИ 61 Grampian Hills Member (Nanafalia Formation) ......... 63,88 PUn REEVE (1849); Turritella na Ned aan ee 18 ВОСИТАЕ Е Bear (1841); Turritella iiare Тт 68,69 ОНИ Merry ad Ынта oe ne 10,11,12 Че SICPEOBIQ ВОО) оо 18,37,46,54,67,68, = аа тео ОО О m NES туз teh vec oe Gregg's Landing Marl Member (Tuscahoma Formation) ........ ENE Ұл te oe es MUT Mene cae NEN 38,63,83,84,86 guarirensis (see machapoorensis) Canne 1994)» ee en, 12,28,30,42,54,61,84,86 ОЧА ШӨ (1996) талл T Eee ул 30,54,61 guppyi (see altilira) HAUSE CLO н) Lo S oer UA uS ERO SS ДОМ Riad EUR лед аза 38 Hansa pde ТОЕ) Sr е NS a ак уз У ва По ои (049) ан etate RP CN ҚА ата а 18 АШТЫ Беру eo io RD een EROS e TADS (ОҚУ acres AW Aene A RETO АЕ 9% Уа и ОЦЕ А e ЈАВА esse bci E e л Ne pee 37,54,61,84 Нап ШӘ рлар ш.м EE CT 44,45 ,47,48,49,61,67,72 Шай ISO SAEs а e ECT tne, Wid SS ои тт 572507911 КЕШ ШЕЙШ ум, Павела NE 46,68 НАМ (И) o ка 18,37,52,53,54,61,67,79,81,82,84,85 Јо а ДУ о) ер њи и e ote куз M od RENTUR 72,86 Ја пр ESS) вв ан аат 49,54,63,72,83,84,85,86 HATS (USD) NN О ee HR а 54,61,83,84,86 Haris anad Palmier (ЗА) ел Е 12,22,37,44,50 VA ыы ҚОНЫСЫ RER, 10,27 | | | PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 129 ИА О ge Т oe 10,12,26,29,30,37,42,43,44,67,67n СЕТИО CIO) te M M CR wr . 16,18,20,23,26,31,37,38,39,40,41,68-70,70-71,78,101,9 chpalmeraeiBowies ов ha. и 37,68—69 С. várs praecarinata Harris. Cooke (1986) nn. 76 сое. Воз). 16,37,38,39,40,41,70-72 fischeri Palmer in Richards and Palmer (1953) ............... ОЕ БАА КЕЛЕЕ. 16,37,40,41,72,72,74,78 Шпон Adegokesu mu en ee 34 Саон МОО О) на ebur c EE 67 gilberti Bowlesid 939) ics nee EN Das ottica ТТТ e d, SUR 16,18,26,31,37,38,39,40,41,49,52,72—73,74,7 imbricatarıuskamarck «Е 42,67 infans:Stenzelvand Turner (0940) e reme dad gu ap M ан u 16,26,31,37,38,39,40,41,73—74,73,8 martinensis Dall (1892) ........... 16,37,39,40,41,74,74,78,79 nigeriensisedepoke (19 TE TER seen 33,34 оуамоуеллаввоке Әт 34 DerditaxGontadadiS65) азастан ее ge 18,20,31,33,37,38,39,40,41,46,52,74-76,75-77,101,8 PjacksonensissGooke: (оле cid acre es си 264/15 DeloweunCooke: @0@2(У) 5 4 е Pos BS rina Palmer (198) са алп do OSEE MET ap OT Ue AEN ELI e Ren 16,18,31,32,37,38,39,40,41,69,76-78,77,78,79,10 K carolina ае вес 37,76-78 tering Байтен (OST) wee ais ccc id 26, 76-78 к. wechesensis Bowles (1939) .................... 37,76-78 rivurbana Cooke (1926) .... 16,31,37,38,39,40,41,78,78,79,10 r. chiapasensis Allison in Allison and Adegoke (1969) .... 78 r. mexicana Allison in Allison and Adegoke (1969) ...... 78 subrina Palmer (1937) ... 16,31,37,38,39,40,41,72,78-79,79,10 subtilis Ке 37,38,74,79,79 tennesseensissGab biG SOO) Fear зала een ee er n 16,18,26,31,37,38,39,40,41,79–80,80,7 vausham BONES (L939) ЕГ 37,38,80,79 Mecht OTOA «o tdem erster od eee 38 Вер GES SA) а а ара nee 46 Hp USUS eS. ner eee igh ve A eer ok hak eek, Que cette ea 18 Lleiprmitls90) ni ee A qas ey, 68 henkeri (see martinensis) Hennig (Обод ыы атт 17 СКА dOS S entre ен ере EU ED ABEL E 31 hilli Gardner 1989) AA ш ee re Жаа. 16,31,37,38,39,41,52-53,55,61,62-63,67,11 ЗІН те ИЛ ЕН 97 РЕВЕ аи 19,100 EOS MIL que ceni iia id E AR 34 O Е 7,8 TIO UD TICK GUO T/A) ohm Sc Decal iss AM tos сл" 18 Eloubrick О Na ne on dU CP пе Ы АТ 9 Houbrick9544) a og ae as А ar ficu 9 Houme k вар as a adn uos оох 20 Houbriek C985) ten s War ноль dep bdo S Tuo vot Ott 14 ЕСИН Des cos М CN 14 HOLD CRISS 22 2-2 20 houstonia (see nasuta) Howell Marbo (1945) Liela nn. 18 hubbardi Hodson ОР ОЛЕН 200. 95. Huddlestumand есе (OSS) ar an. nen 46n Ж о Фома ne ido dl oa E SERE RS e 44 humerosa: Contad во МАН 72. 7 Т 16,18,21,27,31,33,36,37,38,39,40,41,42,43, 68,81,82,84-85,86,90,13 humerosa group . 32,34,36,37,38,39,40,42,43,43—44,80—81,82,84 hybrida Deshayes (1832), Turritella ................ 29,31,42,68 hybrida group (growth limes): 2.2222. 2.2725 26,30,42 Ida КОБО s tm icu oe Edit ca ode AT 10,19:22:24.31,32 En e RNC Зе хасан сы ML И 10 ПОСАО ОИ ОЗ ВИЦ e О ЊЕ e te ae 35 imbricataria Lamarck (1804), Turritella ................. 31,42 imbricataria group (growth lines) ..................... 28,30,42 incertae sedis (Coastal Plain species) ....................... 37 шеш Contad ЗА) Turritella a 18,74 шата Stenzel and Turner (1940); Аана... а.м азаа. ІЛ 16,26,31,37,38,39,40,41,73-74,73,8 infracarinata Grzybowski (1899), Turritella ................. 35 Inglis Member (Moodys Branch Formation) ................ Ta Institut Royal des Sciences Naturelle de Belgique (IRSNB) ... 33 International Commission on Zoological Nomenclature (ICZN) (QS Nec A са ie RU NET LET 45n ДОДАО не театара a аза ee A c 67n EA SPA а: ы по as 10414247 Мейд 199 5) 9 50:35 te cd ee ана 12.17 Деп MOM p ee EE eer Но NAME 42 абое ки KA) ro uo REGE IE аа. 18,21 Jablonski анаа 983) coo recs ues pb RN 17,24 jacksonensis (see perdita) ПОНОВО (ВО) an. e к e tae a ы 18 UNE OSEE ee A Meu Lor usen cR. re 9 KO ts га ата а sean а екен 35 Ке 920) pui ТН 18,37,79 КЕШ ОЧУ о) to nacura oS ye dca peto O Nees 50,67 КАССОВОЕ а s Se n a e do ы 15 UA e Ты 0 UE 11 Kincaid Formation essi.. ooe re i ei 16,38,41,53,55,66,67,82 kincaiuensis Plummer (1933) еа n c en, See EE КОЕ ee 16,31,37,38,39,41,67,11 Kollmann and BEL (T983) нн REST ab a en 34 ROE EO а nr 10,12:23,27 ЈОДА КА ОЕ) Е а E 8,13 Јо а LIGO 22 eave. ve qui Ser 45n KURS IH) mj aw Se am SS ЊЕ ДЕ E са кали зе 57 {КООМ n NR раја en nes EINE drei ІТ 10,29 ПРАВА ЦЕ ЧЕ OS 6) А и 5-2. A acies 32 Јата то ke 9 9)EO uf ЕЕЕ uo ране вани ы сона Een 12 lanensis Набор IPOS) ПП... 27277 ЗЭ Eazanısrand,Protheroi(l 984) rent ре 15,39 Boar АОЛ ОСИ c Uc MP 46,68 [tea Ке SI А ee 68,84 [ioa ОИ ТТТ” 18,37,46,68,72,76 LeBlanc in Barry and Le Blanc (1942) ..... 37,61,81,83,84,85,86 JECHOUE (1939) «usto ce ы она Rc PME e db 18 ОРО О Поинта rl copia der Regis ЕСШЕ itin 10,12,29 levicunea Harris (1896), Palmerella .. 16,31,37,38,39,41,53,67,11 Lieberman, Allmon, and Eldredge (1993) ............... 917,21 Eins TMP OFAN ON 222222222224 8 ызы 38,81 imonni Olsson (022) пите а е ise saver босану Bia cua 45 MEIC US (Ш/ ОВ лук мс a аа 12,18,33 Lisbon Formation .............. 16.38,41,51,52,54,68,69,77,79 usbonensissBowles (1939), Paimerela 2... көнесі cn СОТ Ие 16,31,37,38,39,40,41,47,52,53-54,3 litripa de Gregorio (1890), Turritella an un... 68,69 lloydsmithi Pilsbry and Brown (1917), Turritella ............ 35 Logansport Formation. +... ta recreat о eai ia an 38,81 130 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 Louisiana State University, Museum of Geoscience ....... 45,84 ООШ GIB AT) er e E ОН НИИ 45,45n Ve eee ete ee ИЕ eis ПОВРАТИ Y E 79 lowei (see perdita) machapoorensis Maury (1925), Turritella ................... 35 Ti а а Ку Fods or (1926) КА АИ ИИВ i reds DD m. РАНЕ У НОВО (1926) 95 Л er SUITES 35 m. wiedenmayeri Hodson (1926) 7.2.7... eB 59 NENA ne аа 74 МСЕ СОО) mn РИИ СО РА КОКА 74 MacNeil aud Dockery (1984) rn ne na 8,27 Manoti ond Scherer (1972) Ba. e en 34,35 maiquetiana Weisbord (1962), Turritella ................... DO Tete (SS cee ne Se Net IT Ent ES 12 ДЕЛИ ОРО ОЕ N EN aU ce Рина 10,12,18,29 ИЕНА A седан sette Ce P Lotta ORS Fe 15 Martltaville Formation 4.55 6 ev ere mes 36,41,63,73,83 martinensis Dall (1892), Haustator ...... 16,37,39,40,41,74,74,78,79 m. henkeri Adegoke in Allison and Adegoke (1969) ........ 74 на 80515 ap heii tvs ара NE И or быс атлас иы ЛОЛЕ 12:27 Re a dE rire ele ia do м РО ВР кескені Bobo COLONIA 8,10,11,13,17,18,22,24,26,27,29,30,32,33,36,42 Marwick (19576) odo sciet dues etes 12713.17522,23:242277 NERVEN I) sinc aww item canine ы АТОН 10 harm d (oor D И tes лл. 222. 10,11,19,21,24,26,31,32,44 TEE TOR RS а ааа tne 24-27,26,44 masasensis Marks (1951), Zumtellan cic end ee nenn. 35 matarucana Hodson (1926), Turritella ..................... 39 Matthew’s Landing Marl Member (Porters Creek Formation) .... RET MN а ЕТТЕ 38,41,53,58,67,81,81 jd ner di) и ee eo ae on ee кал es ca 34 iyu ОО e Da а О АВАРА АИ БИРА ne 34 T"Rannpsbtodson (1926); лела za ee 55 PE с RN Re O 14,17 MeBean Formation uan 38,46,47,54,68,77,79,80 mcbeanensis Bowles (1939), Turritella ................... 37,38 ИО ЛЕ ООО Е ale 61,74 ОКЕАН у ee ae ee 34 ieyoensiv Olson 1931). Turritella e sus crier att eider ke 95 Merriam (1941) ..... 8,9,12,13,17,21,27,28,29,30,32,34,36,38,40 merriami Dickerson (1913), Turritella ...................... 36 Mies EURO SURE RC а ОНА АСА ТАЈ 8,10,29 mexicana (see rivurbana) ӘУЕН СЕ ee MERE AU AAT 46 NEEDS ead Leda ot vea bsc RYE E E IE 14 Midway Group. undificrentiated.... 752222220002 20227 84 Miniter scl ШЕ (LSO ss wink mas mese cane heeds АРУ) 45 mimetes Brown and Pilsbry (1917), Turritella ............... 30 mmmigoensis Bowles (1939), Итеа н. лела ee 37 mirandana (see altilira) Mississippi Office of Geology (MGS) .................... 45,78 mixta Gardner (1928); Turritella asire en er Y 18 momljera H.C. bea (1841); Turritella i lI 68,69 montanitensis Hodson (1926); Turritella Lis 35 od Е 272.2. аған 10,12,26,30,37,67 Moody’s Branch Formation ... 16,38,41,46,47,49,54,72,74,75,78 hilos су а тола ел Т TN RE ANNE I 86 МИКК КӨН C PES ДЫМЫ ызаны РИОЯ, РАДО morphomemie analysis s.i. 222 си ин 55,100-102 КОН EN nennen ТІ Moss and Allo (1993). «01s ЫТ 19,100,101 DiI LEI. A E UR 18 moton Conad (1830), POWErelitt- e iau t dre ARR Peete tee 18,20, 23,27,31,32,36,48,53,53,54-64,66,68,85,101 mortom mediavid BOWLES (1939) ee VET a E ee Pda би 37,39,40,41,48,55,57-61,61,2 ТОРОНТО ОПАО ВАСИ CLG SO) scene erase A E oto A сте, SED Es 25,37,38,39,40,41,54—61,65,88,1-2 тогіопі postmortoni Harris (18942) ......................... LOL ЖАЛАҚ ae rap ka dua 16,37,38,39,40,41,55,57-64,59,61-64,88,2 mononi premorom” GOvont (LOSS) meme ern, ee; To 2 ке ее 16,18,30,31,37,38,39,41,48,53,55,57-64,64,1 ЙОРЛО ОШО neta ser а есек ее 36,37,40,43,43-44,66 MOS ИЛМ ТЕЛА ҚАЗЫ Сет ар НИП ARTE erg NETS 10 multa Whitheld-(1869), Turritella inre eet e ee: ESPOT CA SEMEN 31,37,38,39,40,41,81,85-86,12 Miter ay (О О) o E NE Т 38 Museum of Comparative Zoology, Harvard University (MCZ) ... a Нилу 7-8,44,45,46,53,63,66,67,69,77,81,86,88,90,91 Nalicola Formation е, Senat 38,41,53,58,67,81 Nanatala Formation О ои е 16,38,41,55,85,86,88 INAumjemoysEormationme еее at к ЛҮ ee eee tae 65.66 пази OD ВОО) еа ay artes 000 37,38,14 оргаша тепсе апае (5940) er ТОЕ 37 METE Вб ев (ОВО) Е ЕО ОИ MES 37 ПОПТОН е ee en O) Ne STRUPVULCTISIS. BO WIESEEL OSD) re er p КӨШ (9 OR esa ee ee 14 Тео апд Ра е (ӨӨ шылу Ser iate re T TUE E 7 IN COMA OMAR А URS RAG DRT CRETE Te E L ан а 11,29 ЖООШ ПОР ЕТЕ EX E er EE CEPR COREL aE NWN 10,29 nermexd Rates CSOD) Turela mi ROTE 37,38 ING ОНО Aetna Pe ee 13 nicholsi (see vistana) NN ee БУА 35 Nicol Shaak and нораво (1970) een 74 ПРОТОН AREO KE 9 A ЈУ ОАО 33 INDI ОШОО DUS tetra ЛИ Nc. ЕМИ ee ТЕТІ! 10 omna Conrad (1833) Turritella ООА 37/36 Ocala ETOO tea. eee о вах и. 74 ocoyana Coma (1590) не. eee ees 36 ola Plummer (1933), Turritella ccoo.» 37,38,41,48,52,11 oligoploka (see femina) IESO CLO ZO aaa ai once teva ce USER TRAD CREDERE ERE T 11 DIM RAP. 10 ОР ШИШ CLO) Eee Fev T VS meee 33,34 Oppenheim Adegoke (1977), LOrquesi@ i ss cree eee 34 (OTERO TUN Wt eck N А CEA I oe ae 11 oreodoxa (see bifastigata) Ostrea thirsae beds (Nanafalia Formation) .................. 63 ССОО) а от И Aa 46 öyawoyei Adegoke 1977), Haustator о т 34 Pachecoensis Stanton (1896), Turritella ern 36 Paleontological Research Institution (PRD o МИТ валови ee 7–8,45,46,49,50,51,53,63,69,77,78,79,88 Ја УБА ДА ERRANG 45 Је роу O Ае Mima teal halle ie either IRR Қап ah КЕРИ: 12,18,21,30,37,40,42,44,46,47,49,50,51,53,54,61, 65,68,69,76,78 Palmen (maras ad Palmer ЭА И ЕТА ОБРИ ss ts 18,45,46,49,50,51,70,73,74,75,76,78 Palmer Gn Richardsand Palmer, 1953) me 12 PA (ОШО en ee TEE EE EEA EN 35 P О) M keke me SN RT 35 Pamor anma Brann (DICO). и nannten oa 44,45,46,47,49,50,51,52,53,54,61,63,65,67,68,69, 70,72,73,74,75,76,78,79,80,81,82,83,85 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 131 palmerae (see carinata) Бартегецав Е A na ahnen 37,44,45 alabamiensis Ме бЭ) И ИКО TM Nc те 3. 16,18,26,31,37,38,39,40,41,67-68,73,82,7 Alvear Comrad (IR ee RENNEN 16,18,25,31,33,34,36,37,38,39,40,41,45—46,50,76,5 арпа де Cregomo вет Ст 22000 dremooidiGonrad Во) en tne d tT AAN 16,18,25,31,37,38,39,41,46-47,49,50,54,5 свота тен алт ВО ee ee 37,47 a. danvillensis Stenzel and Turner (1940) ....... 18,31,37,47 Chirena-Stenzel and bummer (940) ОЕ SR NENNE, 18,25,31,37,38,39,40,41,47-48,3 clevelandia Harms: (1890) МА тата та тн A Жалы РҚА ira че 16,18,25,31,37,38,39,40,41,48-49,50,5 creola. Palmer in Harris and Palmer (94 7)" 22222... M ee ча 16,18,25,31,37,38,39,40,41,49-50,50,52,4 dumblei Harris (1895) ..16,18,25,31,37,38,39,40,41,48,49,50,6 dütexata Harris (895): arm O ТТС ИРЕ; 16,18,25,31,37,38,39,40,41,47,50-51,51,52,54,3 femina Stenzel in Renick and Stenzel (1931) ................. IT ER 16,18,25,31,37,38,39,40,41,47,50,51–52,54,66,3 f: oligoploka Stenzel in Renick and Stenzel (1931) .... 37,51 Aili Gardiier (MOSS) rr rate ee ae T EI EN 16,31,37,38,39,41,52-53,55,61,62-63,67,11 levicunea Harris (1896) ........... 16,31,37,38,39,41,53,67,11 lisbonensis Bowles (1939) . 16,31,37,38,39,40,41,47,52,53-54,3 MOTION COLAC Mss OE cete теа | Ses 18,20, 23,27,31,32,36,48,53,53,54—64,66,68,85,101 mortoni mediavie SOLE ClO SO Tm О ОАО И 37,39,40,41,48,55,57-61,61,2 MOTION MONON CONTAC Әә Ше тет cL MEN e ЫА 25,37,38,39,40,41,54-61,65,88,1-2 ЛОЙ ОШ ОУ ОНО ТАТА У 394a) Е КОЕ reir aban Тее 16,37,38,39,40,41,55,57-64,59,61-64,88,2 mortoni DIEMONONE ОУ ОООО eR к нта Ou ТЕТ 16,18,30,31,37,38,39,41,48,53,55,57-64,64,1 pleboides Vaughan (1895) ....... 1.8,25,9.1,97538,39,71,92409,0 potomacensis Clark and Martin (1901) .......... 37,38,65-66 Spree ee paa EUN 66,11 WICNZELIMEWSPECIES eos SNR МИЕ Л 37,38,39,41,66-67,11 paraguanensis (see machapoorensis) paraguanensis (see variegata) LONDON N Re E OE TI 1229 PAGOLA ете ee ТИТ. 10 Paspotanza Member (Aquia Formation) .................... 33 Pattetson ClO Site CHE conor AU oce c cc RAS qn 9 PAUP (Phylogenetic Analysis Using Parsimony) ............ 15 Pendleton FOMO от 38,41,63,83,85,86 perattenuata Heilprim (1387) тие = r cnn сы ДА 35 perdita Conrad воен Шоты Е паша а 18,20,31,33,37,38,39,40,41,46,52,74—76,75—77,101,8 Di JACKSONCHSIS COOKE UDO) ТТ” 20197575 Dr Ower COOR ТОО ee c pM et СС 37 TS ERAS И ee 67n Pent and Ce Kenari ооо a 22022 67n Penrand Le елата 0:990) c9 eee eee EET 67n LEV TOLD E E Сы NEC TRN 10,12,29 DINOS CU ШИНЕ E ne EN ETE 8,15 DUSOTY Garn CIL ОЛЕИН S E ee 18 Piscataway Member (Aquia Formation) .................... 3m Pisgah Member (Kincaid Formation). Е 67 РЇ Ла Guppy (ИВО LUNEA 222222222 222 39 PNV EOE ТТТ 22022 G 10,29 pleboides Vaughan (1895) Р текей nun... пе ен ЖЫҒА MRR LIB А DEN 18, 25,31,37,38,39,41,52,65,6 Pommer (OSs) err ke nate ann nes AREA ERES 37,44,51,52,53,83 plummeri Stenzel and Turner (1940), Turritella ..... 37,41,67,11 polysticha Stenzel and Turner (1940), Turritella ........ 37,38,11 Onde ID A) RS LUCA RR ee 31 Јо Arner А het ee cick ДАУА леан ae 7 PonersiCreckbormmation 27772 Е MER: 38,41,53,67,81 postmortoni (see mortoni) IPotamididdo er re EO С 20 potomacensis Clark and Martin (1901), Palmerella .... 37,38,65-66 ОУ СЦО ЕС Ао о EI EEE 10 praecarinata (see carinata) praecellens Brown and Pilsbry (1917), Turritella ............. 35 prdecincia:Gonrad (еба) иена? кеі! qu nr ea Duc ieri o. 20,31,36,39,40,41,64,81,85,86-88,87-89,101 p. praecincta Conrad (1864) ......... 16,27,37,38,86-88,86,14 D. virginiensis new subspecies .............. 16,37,38,88,90,14 LOVENUM CT OSA COVO OS) 0e son HRS Pra de 16,18,31,37,38,39,41,42,81,82,88-90,90,91,13 “premortoni” (see mortoni) Drenunciasspieker (1922); епа zn... AAA TOREM 35 YONG. da оаа құлан ала Ser 12 NOVO NE ео RUNE NE 4012,29 ЈЕ ОНО etter mue er БАЁ 10,29 Тао ACs eec cer mad d Gane NE dn 10 quadrilira Johnson (1898) Тикпей зао ен. те 18 Queen уво та О Ота co Т ee 38,41,48 quirosana (see venezuelana) ¡RAI ALO A docte ie SE os о OR 38 RPA аита ELEM M | nase Bs ла ee 32,101 Raup and stanley (MTS а dedu. ur rie 13 Та CLS dO ROSE E nee 18 КЕК БОДО 99 e M EHE Sce c po ee Ы 38,41,48 Remok аше Оа) en er sU ILIUM 37,44 revera Warme (1917); TUUM re SRL TN 36 RE U E а LLL алақол к A 10,33 A Аа сроке о) о RT Л 34 КСО ИЕ аи о EM с. 7 Richards and PREC ISI ЕН ИЕА 37,44 Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands . 10 nna Pr A ISN TALOUN ust ae RUNE 16,18,31,32,37,38,39,40,41,69,76—78,77,78,79,10 P CORNU RAEE (9S) res cotes eee RR 37,76–78 МЕРЕ ка AC OU)... . a ТТІ Б 26,76-78 ғ. wechesensis Bowles (1939) ...................... 37,76–78 РОО ОПО И Е 24,36,37,39,40,43,43-44,74,78,80,82 Risso! 0182 GR ee аға Res S icd TIVUrDAna-Cooker( 1926), аиан ТА ЕИ EE. P AM ER 16,31,37,38,39,40,41,78,78,79,10 ғ. chiapasensis Allison in Allison and Adegoke (1969) ...... 78 r. mexicana Allison in Allison and Adegoke (1969) ........ 78 Roberts, Soemodihardjo, and Kastoro (1982) ............... 97 robusta Gryzbowski (1899), Turritella ...................... 31 Rogor ап Кос (19 37) е н Eod Ea Ld PUE 18 ING MUR SO) Mc c ec са 15 ROSEMA О РАКА AA e no CLEA 210752 34 КОЗ А О. Eh ОН ОРН een ал па а ње а y ПОШАО DAS) sis es ARR RA Ае АЛАК MENU 33,34,38,42 NA VMCLO Whe PORATION ccc veo а qe eT 41 SECC CIS OO) Lex Duce ere Ou УТД АНА JU 20,12 sajordi Gabo (1860), Turritella ner. 37,38,41,11 SOIRS) аа РРА ИЕ лм) TA SER ERBE O МЗ 46 КОШОК TE ee een EEE 59 ЗАМЕ Не тј о а МЕ с T anto зен ram m 7 Selleltena (Ое Балым 2 МАТА тына tie ence o a 10 СОАО ОИ ee Bec Pelee анон RE REL RERBA 2l DOSE) EN dr ыры ақысы = AIF 82 Schuchert, Dall, Stanton, and Bassler (1905) ........ 47,49,65,74 SCENE DEREN ыы өле deeem qe or vei ege d 10 MECN o ОВО 5). TUPTOllG ua Ls snes iad eorr E 18 SEINES (0929). amas o Dnus studs 54,61,84,85,86 Sitka Riven PORO UNS sero os ек or ERA RUIN 38 Sae СЕРЕ ОЕ А ауан ВАР CDS 14 ЭАК А ООО Ы EHEN Bree el 14 Shimer and Shrock (1944) ea). 2 IEEE 54,55,61,84 ООО Аа ae а 1% 19 ОООО De en RE 102571 PLS BOER OPUS РЫНА Luar inso ао um. a be SU 38 MOTOS AR NE A ee TEUER. О ЫЫ КЫШЫ 11,12,29 Signor (1982)« 3 605 d. ni а Lac ta le а 31 ЭНИРБӘНИШУӨШЕ555 eee loe s Bedv dup: ro emn 17 SEHE S EDUC. o етене Саты быт m 2/2) SOTTO ARE 7 BEINE EA OO и eer ee ТОЛ? Өш mnd ОБО (1887) ie een 54,67,83 Sin Johnson, and: Langdon (1894) Fr оно 72 D А DR ee cd o CERT ry les Se pc ES REA RO NE с кууы» 12 smithvillensis (see nasuta) socorroenesis (see buchivacoana) БОНУ СО ee 28,32,36 SO Тј ee 8,17,19 RSET acis ren СИН BAR 66,11 SPCC ВА СЕО een od pet never mew ee 13 SPST РОВ у ace is ae 34 UNICI OUO EOS ne eens anes 18,21,38,43,85 ОО Е aan nee 52 10,12,18,29 Mog) A EN ea 48 ЖОЛШЫ A a EA 33 SSNS ЭН, ee sas ee ee I 32 Binley ОВ К os le 31 SOON Cea er E A E E E 7 Өе EOS c Reus ааа een 66 Stenzel in Renick and Stenzel (1931) .................. 18,51,73 ӘЛӘНРЕР AN ne en 8,44 SIEHE De e M LN M UM MC C PT 44 Stenzel and Turner (1940) ..... sess 18,37,44,47,51,73 ЗЕКЕН СІР(ШӘД Lei. ee, ан Lad 44,45,46,47,49,50,51,52,53,54,61,65,67,68,69, 70,72,73,74,75,76,78,79,80,81,82,83,84,85,86 stenzeli new species, Palmerella .......... 37,38,39,41,66-67,11 SIEDHENBONELEIS TER e d NE La at 79 пен о А ЕВЕ ЕТ А РА ВИНИЛ ЗАНИМА 19 Орто (ВОЯ Er имена ET 18 SIEDHENBON COSE qu Re an. A d 10 Stephensomand-Erider ое) 5,69 oon co vd beet om tO 61 Stephenson and Veatch (1915) 9 2 Her 61,81,84 МЕТАН aA ус) врта БАН С ПИО ee а па ee IS 42,46 ПИО TS те AA ee O AR E A т 10,12,29 ОИСЕ НЕШЕ A inch deen 2224.20 16,38,41,48,50,73 a BES nn oop corse Moe МЕ RE 39 o cs lA SORS nn ee 15 SUDOHSWIGTO-Brocchn (1814), Turritella eee renr ive 42 УНОС ОРИ group (Growth: lines). 0... ads 28,42 subannulata kleilprin (1887), Turritella .. ....... ia зз 18 subgrundifera Dal (1892), Turtle a ess 18,35 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 subio Ramen (DIS) СИТА eee ctt сезаз RUE Жы. 16,31,37,38,39,40,41,72,78-79,79,10 SUbspEeciesdetinitionm een S DEUS ыла тар da 13 subtilis Kellum (926) EA CUSCO от 37,38,74,79,79 өпртаѕресоаха AN тал Батар Ды AS 8,10,13,40-44 Swamsomtls 40) SAA CA A 20 ШӘ ШИЙ re a tds дыр 15 БЛОА а ОО) ТИЛ ИЯНЫҢ а A 55 a л» 55 rc SAW OCHO NE ee Rc 11 RGGI T/L) ПОН НА REA it ETE 1112.29) Allah Auta Or a On кен» ee ER IE 41 taratarana (see gatunensis) Megane Be ea nern 7 Tehuacana Limestone (Kincaid Formation) .............. 66,67 tennesseensis Gabb 86060 Haustatone men nenn dele ee ИЉА 16,18, 26,31,37,38,39,40,41,79-80,80,7 terebra Linnaeus (1758), Turritella .. 9,20,31,56,58,56-58,61,101 terebralssleam ато о) Uca и. 42 terebralis group (growth lines) о 28,42 terstriata Rogers and Rogers (1837), Turritella .............. 18 Texas Memorial Museum/Bureau of Economic Geology (TMM/BEG) ынша қары ORA UTC Л ЖО ee 7,44,45,47,48,50,51,73 ТОЗОН О Ts aren Са uix on на tisse А Ат ed a 17 ОТСО a ae a o EN es ER RS 17 rele CO Sco Ee cw era E Dod rco mu ok otek ds 10 тоа de Gregorio (1890) Zurritellä: 222221022100. 68,69 тј етике Conrad заа) еа пл ва lee 18 MILO А (ШОРА ET ы Ала сз MISERE n cep can 24 ШОУ АН О AD) АЕ RU I a en Wil tolenasensis Merriam (1941), Turritella ..................... 36 ОНО lel аз Ben Dame Mesa 10,12,29,33,36,44,74,78,80 ХОШ ТА СВА о) Е ыйы Жака йн Тан 26,33,10 папе о (ИЕ ли ава ККК Te OC eR ee 10,12,29 ON тте сы ыт 10,30,32,33,42 adabionensis Oppenheim, LI I) nn na. 33,34 ОРрейле її Adegoke (ШО Ук. Ines jar er 34 Ра авва У САО И АИО 91 corp p ШУ Ө ДШ К лу ee тла DEO а 8,63,73,81 Toulmin (1977) ...... 8,15,16,38,41,44,46,47,53,55,61,63,64,67, 68,72,73,74,76,78,81,83,85,86,88 Оаа LaMoreaux (1963) iii: Come eR ы жо 44 VOI MS MS “Turritella t нет RE ute V Ee DEUS Un SANE кш ы nid eue 37,38,39,41,80,82,83,90-91, 90,12 DU CAO AA (LS OU) MUNG ad 18 ALODIUM Des Ut eis e LE 1.0, 21521229 ONDAS Hua: tad pts ee 63,84 MRO RTE A (CUS SS) etre ака aan ee 54 Toomey amid Holmes (Usd) и ran a 18 VEA A ET ТАҒ құ ы re 9 ROUES UD EE ed LT ren 1 turneri Plümimer (1953), Turritella 19920. 42.2 эы хш DIO AO PUTTIN ASTRO н 8209) Turritella Trn ts Л A лата 42 ААУ отоо (оом 28,42 “Turritella” (humerosa group) aldrichi Bowles (1939) ........ 16,18,27,31,37,38,39,40,41,68, 80,8 1-82,81,82,84,85,9 1,12 biboraensis Gardner (1945) ....... 16,37,40,41,80,82,82,90,14 claytonensis Bowles (1939) .. 16,37,38,39,40,41,80,82,82,83,12 СШ HOMO ТЕЛА BO S а ПЫ АТЫН Аы cA I s 16,37,38,39,41,81,83—84,83,85,86,13 gardnerae LeBlanc (1942) a asa 37,40,41,80,84,84 ДПНОТОЗОКЕОТША ОЙ (165 0). ркен таса еее anne aN КК” 16,18,21,27,31,33,36,37,38,39,40,41,42,43, 68,81,82,84-85,86,90,13 PALEOCENE AND EOCENE TURRITELLID GASTROPODS: ALLMON 133 multilira Whitfield (1865) ...... 31,37,38,39,40,41,81,85-86,12 Draecineia:Gontadi(lS OF) jaa та ee RE ee По 20,31,36,39,40,41,64,81,85,86–88,87–89, 101 p. praecincta Conrad (1864) ....... 16,27,37,38,86-88,86,14 p. virginiensis new subspecies ............ 16,37,38,88, 90,14 e prehumerosausGiovoni (LOSS) aier c EM I ee dip d AEE 16,18,31,37,38,39,41,42,81,82,88—90,90,91,13 toulmini new species ....... 37,38,39,41,80,82,83,90-91,90,12 Turritella ын re a DIESE. 8,9,10,13,29 abrupta Spieker 01922)... o. taa Фаз D EN 35 асторота Wall (ТЕЗ) ААД PERTINENCE RISE SENDER ROUND 212) alöwensis Hodson (926). C ne 0“ 35 пита Coma И БОЕ е 35:90 a. guppyi Cossmann AO 2 Pu: S9 АТИНИ ТИ НО SOLIS ee 33 а. urumacoensis Hodson О онлине en 95 andreasi Williston, in Hodson (1926) >:.................5 35 alturana:Spieken(1922) we a AA ин 35 алата ооа ОРЕЛ De ОПИРАО а NA шко олы E ОС DR ERR ETT 36 aneuillanacGoo ke: ОО) M A E 30) avovarensis OSOS ee ИЕ 95) bellifera АТО З 8384,13 Denda dinensisi ds OO EE c ae ees Bo) bifastigata.Nelsen (USO) aru 27... 89 b. cartagenensis Brown and Pilsbry (1917) .............. 55 Drore odo aiO УВО НО e Rennes Е S broderipiana д Ойлау (TSAO) e ne 36 buchivacodna EOS ONU eects 2 35 biscanonensis О ВОЛИ IZO) er Ем ga bi СОЛИСТ HOON (HOD o) Е Е в 35 ВИСЕ НО ЗОО О aces een 55) БЕОСОПКОСВУ И AO SOM IZO) a etd Meu Бо р. сосодпапа носо Ша. E 35 bunkerhillensis Palmer in Harris and Palmer (1947) ....... 37 buwaldand Dickerson: (1916) men а КЫСА Бан 36 Caleta ОЗО GOS) nn E E ыш туло BS Сара тоташа. 22 38 carota MacNeil in MacNeil and Dockery (1984) ........... 49 cauredalioensis AOS ONO LO 1 ЕТТЕ 32 CHICOCNSIS: Gal bs У ОД) ЖТ ee a ee 36 САРОЙ Walls ШАУ ТТЫ es 3 СИОН ОЗО OL ue c EU 35 CIGVIONENSISEBOWIES ШОВК еи. 37 columbiana Weisbora (1929ER И ПИЕ Е Ие 515) conquistadorana Hanna and Israelsky (1925) .............. 39 cooper Carpenter Шо 22220222222 00 36 tornellanasklodson ASS e tie to ыдысы ақ 35 COSTAR CENSOS SOLE EN NO NOS а TEE SEEDS: 515) crocus Cooke GUIS) heat rt cae us con A re 35 СИДОН БОЛ ОЗ ee 3 curamichatens is ОЗО 22 GOUVETISIS Dockery USSO ооо 27,37,38,39,46 domingensis Brown and Pilsbry (1917) ................... 35 eClemensis Hodson 9060) ee ee erh 35 VAIGUTIEASISBBIOdSOH ОСО Е 35 AM SD) ee em ар зарае 35 RliearmenensisHlodsom ОКОН ВА 35 Tarei BION (ШОЛ И Т е S5 ФОТОН Olsson hos Е eM N ahs 89 ДУ вуста ЗУ ee а а ү 3e g. caronensis МАне nn ann 35 Er ЁШ ИЛО ОШАДЫ So ee a 35 Еа Те НӘ ton Папа (ола A re ee So Јаве Коле ома ОВ. 313) арена посо 1926). 2 ие 35 Она атана (ауа sony +2 a AA к шшш; 35 рано Conrad ЗОО Е 2200 - 8,37 TOSES ODIOS) A Te Om н ME 35 пасата т уров. 35 kingaidensis РӨ (199 3) ey. oe ты ТҮ 37 (ОТОНА Во (ОВ) И 35 limonen TOES on KDD) Т Ge es ушШ... 35 lloydsmithi Pilsbry and Brown (1917) .................... 35 Mmacnapoorensis MAUS О) не ene Өсе рды 35 ПИО ЕН, Hodson (1926) а. 35 m. DaraguanensiseHlodson (MOZO) ов 35 Mawiedenimayert Hodson Өре 2.2222 52) maiguetiana УМЕО 62) ete A A У.Е Bo nigscsensissNTanks (AIDS) Е aor Ones deir 35 matatucatasblodson 20) 2,222 dins 35 mautyrtodsonitloP 6). re арада us 35 MEDEa nens BONES LOBO) ент 02-9 0 37,38 ЙОКО ӨЛЕ SOMOS) ca аи 777777 35 МАНА И t КЕЗО (ШІ Т a CUTS EA 36 МАО АЛБИ које кан SDE (1917). а 32 ПИО OESS Оез CO SO) очне на cu 23. ысы. 37 Mmontanıtensıssklodson(1926) о Swen 35 HOS NICMOS ADO ЕСО) ere а ва ee Mu 37 n brazita Stenzeland lumen (94022222... 37 ПАЈЕ ПИВО Ез (Во) e ересен ме. 37 HARO OMA QS 95) 2 242422 37 nsmuchvillensisBowles(1939) н. 37 nerinexa Harris (1895) алоо, 37 OCOV ANGE ORTA SO) e а 36 ODrHLGACOnradi 339), NEUE Евы 37,38 dla Plummer озо 37,38,41,48,52,11 Pachecdensis Stanton (ӨӨ e ea 36 еа елата а Ке они е ои н 35 DIE Nau СОО ан а ee 30 plummeri Stenzel and Turner (1940) ............. 37,41,67,11 polysticha Stenzel and Turner (1940) ................ 37,38,11 praccellensibrowneandubilsbry (1917) 22.2... 33 ТОЕ ОАР ЗКО (UO PO E сн UL c P uM 33 VEVEN TEES ae Mec DOT VM II D T. 36 балалары ола o ue mI T 22 SUDAR aaa Brocchi Шы... % 42 Sud de do» Dy SCI G IMD ALIS ODEN C M Ду. 33 terebra Linnaeus (1758) ......... 9,20,31,56,58,56-58,61,101 Поа Ое О ОНО (ТӨӨ а cs 22 quem 68,69 WANDER (SS) Л. о ен e 18 rolenasensisaviermanm (OAM) ete ОКЕ 36 MORIA 2722-2-25 c 2 37 Pas ana Conradi i99) ке vende NE ee s. 20” уалсотавзрлекен рол uM I. 35 Тезата Тае СЕН. 18,33. ViDAraguanensis Elodsom(1926) o 2 u. 35 Veezuelanadodson Шо. 2222. 35 ОТОО ПОО БОЕО) are 35 У ДУО nia ааз ISO) с са ен NL a UU D 35 СИЛА мала тара ТТТ 22. 33 AMV ОЛИ (1926) ne, 333) AA 2222-2220 2-0 8,44 ЛИН ЕАУ ОБ ЛИСТОМ OS ee. 40 Rubee (Chics: nn m и ну OmU: 8,9,10,13,33,36,45 ШИН Шинаны ee E 8,10,42 ПЕШ О О Ба вв Mey 22 са EE 11 ТООС Угу жалуу у лр шатка те oe UM с с: 11,12 Tuseahoma Formation ............. 38,41,52,55,63,83,84,85,86 134 PALAEONTOGRAPHICA AMERICANA, NUMBER 59 urumacoensis (see altilira) University of California, Museum of Paleontology (UCMP) ..... SIN LEE ИМИ ое SPM сле 45,74,78 University of North Carolina, Chapel Hill, Department of Geology r en еу суз НИ Еу суук OMS 45 U.S.Geological Survey (USGS) ............................ 45 U.S.National Museum (USNM) .............................. Ele sie ПА 7,44,46,47,49,50,52,53,54,63,64,65,66, 72,73,74,75,78,79,80,81,82,83,85,86,90 uvasana Conrad (1855), Turritella ......................... 36 Valenciennes (1832) + ae en RT 18 Van Nieuwenhuise and Colquhoun (1982) .................. 44 Van alert (0965) nn NE REED 61 variabilis Conrad (1830), Turritella ........................ 18 а ТН” аа ааа а 20 varicosta Spieker (1922), Turritella ......................... 35 variegata Linnaeus (1758), Turritella .................... JOD v. paraguanensis Hodson (1926) ......................... Do Vaughan’ (1895) ........ QE ME LA сс C T 18,37,65 МОНА (606) аа ИЕЛ ATH ИНЕ ed 44,49,65 vaughani Bowles (1939), Haustator ................ 37,38,80,79 Veatch (1906) и ази аа 54,61,63 Veatch and Stephenson (1911) .................... 44,61,81,84 venezuelana Hodson (1926), Turritella ..................... 85 у. quirosana Hodson (1926) ............................. 35 EPI A VA Ен И НР ERRAT а [1 Vermienlanitae ame o UU MEER HET FEIN pa 11 vermicularis group (growth lines) .......................... 28 NEMO Бар ee aa ааа аа 74 vertebroides Morton (1834), Turritella ................... 18,36 Vialov and Soloun (1936): — ... m. re a i 42 Viesca Member (Weches Formation) ....................... 9 virginiensis (see praecincta) vistana Hodson (1926), Turritella .......................... Uo v. nicholsi Hodson (1926) ............................... 85 Moker (LOGI) o> coepere ил. л Оз ОРИ Е 54,84 O БОМ ОС ДОО 17 Wal odia ee ROT СМЕНА ESTO 13 NATI NV tin A А АСЫМ ne 7 АО О ИТЕ Е EEE 44,66 warfieldi (see buchivacoana) Wassen ma Wilbert (TIAI) 9 АЕ. НЕТО 44,86 Ао and Wheeler (1981) Еи 2/7 We ches Formand t OI eon ERE Tek eto 16,38,41,48,51,52,77 weidenmeyeri (see machapoorensis) Wie cl ОВО) н ke err ea D NIU ДИИ 14,15-16, Wheelock Member (Cook Mountain Formation) ....... 50,52,73 We Bilun ОНА ОИ a wee foe wissen ERIS 38,46,47,49 МИ ти (RS ОО) ОА ие 18,37,67,78,81,83,85 УИА) otov ete AER tO PU ИНИ HERBAS 85 WIDER Ое AGS 44,47,49 WileoxiGroup undifferentiated о | ман ТОРИ В 85 Wiley ШО Ш Se сыны REAL RE 14 Willtanisbur Formation ИО ИЕЛИ 55 Wile Pont Formationen Т ar en are 38,41,68 Wilma (Oe (НЕТ e rennen 48 O ТӨТЕ Arta Aboud GONE JOE delit Ms 57 WO odin: CISD S) b me doe АКА TORIS E NUM 30,42 Wis ШЕ 972) а и аан ПЕРИЛ Н ИИ 34 ОООО ТТТ idee ОЬ АИ E Н 11 Woodstock Member (Nanjemoy Formation) ............. 65,66 Woodward (Тво Ш) 4 LT Eo RETTEN TER 45n WAG Аваат е (L9 81b) мины же, Mone RR ER 5 V Te RM LESE E eats СТАЛ er 10 ДОО Formation o cie ee а ЕНЕ А 38,41,47,49 Аа у» а Буу зле» иен от ДАНИР О 11,12,29 ОЛОИ И со ОАА RIN TOS 10,12,29 ТОМАСА а eet РЕЛИКВИИ АЕ 21 ОУ а; oe C О AAA АА АА НЕЕ 7 zand ROn (LIDO Turritella Еа ARA И АНЫ 58:55 PREPARATION OF MANUSCRIPTS Palaeontographica Americana currently appears irregularly, on an average of about one issue every other year. This series is a publication outlet for significant longer paleontological monographs (i.e., more than approx. 200 printed pages) for which high quality photographic illustrations and the large quarto format are required. Submissions are welcome from any author, regardless of institutional or or- ganizational affiliation. Authors must, however, be members of the Paleontological Research Institution at time of publication; annual membership is currently US$25.00. Publication costs of Palaeontographica Americana are heavily subsi- dized by the Institution, but authors are currently required to pay illustration charges at a rate of $120.00 per plate and $35.00 per text-figure. Important references for style and format are 1) Bulletins of American Pale- ontology “Instructions for Authors" (volume 108, number 347, pages 149-153); 2) Chicago Manual of Style (fourteenth edition) 1993. 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